AFRICAN CHIROPTERA REPORT 2020

AFRICAN CHIROPTERA REPORT 2020 LISTED ON EBSCO PUBLISHING DATABASE AFRICAN CHIROPTERA REPORT Published by: AfricanBats NPC, African Chiroptera Project, Pretoria, Republic of South Africa Copyright: © 2020 African Chiroptera Project © 2020 African Chiroptera Report © 2020 African Chiroptera Database Reproduction of this publication for educational or other noncommercial purposes is authorized without prior written permission from the copyright holder provided the source is fully acknowledged. Reproduction of this publication for resale or other commercial purposes is prohibited without prior written permission of the copyright holders. Citation: ACR. 2020. African Chiroptera Report 2020. V. Van Cakenberghe and E.C.J. Seamark (Eds). AfricanBats NPC, Pretoria. i-xviii + 8542 pp. ISSN: 1990-6471 Database development: Victor Van Cakenberghe. Available from: http://www.africanbats.org Online Available: Wednesday, 28 October 2020 Note: The designation of geographical entities in this report, and the presentation of the material, do not imply the expression of any opinion whatsoever on the part of the African Chiroptera Project, managing editors, or other collaborating individuals or organisations concerning the legal status of any country, territory, or area, or of its authorities, or concerning the delimitation of its frontiers or boundaries. Furthermore, the views expressed in this report do not necessarily reflect those of any of the collaborating organisations. This is a print report from the data contained in the 2020 version of the African Chiroptera Database. The resulting views and outputs are based solely on the data contained within the tables in this database. Managing Editors: Victor Van Cakenberghe and Ernest C.J. Seamark Taxonomic Advisory Committee: Victor Van Cakenberghe (University of Antwerp, Belgium), Teresa C. Kearney (Ditsong National Museum of Natural History Museum, South Africa), Peter J. Taylor (University of Venda, South Africa), Petr Benda (National Museum (Natural History), Prague, Czech Republic), Gabor Csorba (Hungarian Natural History Museum, Hungary), and Stéphane Aulagnier (I.N.R.A., Castanet Tolosan Cedex). Contributors: Rudolf HASLAUER Oberkreut 4A D-85309 Poernbach Germany Dr. Teresa KEARNEY Small Mammal Department Ditsong National Museum of Natural History P.O. Box 413 Pretoria, 0001 South Africa Dr. Mark KEITH Eugène Marais Chair of Wildlife Management Mammal Research Institute Faculty of Natural and Agricultural Sciences University of Pretoria, Private Bag X20 Hatfield 0028, South Africa Supporting Organizations: Ernest C.J. SEAMARK AfricanBats NPC 357 Botha Ave Kloofsig, 0157 South Africa Dr. Victor VAN CAKENBERGHE University of Antwerp Department of Biology Labo for Functional Morphology Campus Drie Eiken Universiteitsplein 1 B-2610 Wilrijk Belgium iv ISSN 1990-6471 Funding Organizations: The organizations shown below have contributed funds to support projects that assist with the production and refinement of the African Chiroptera Report 2020. African Chiroptera Report 2020 v Table of Contents: Introduction ............................................................................................................................................ 1 Taxa profiles ........................................................................................................................................ 12 ORDER CHIROPTERA Blumenbach, 1779 ........................................................................................ 12 †Family AEGYPTONYCTERIDAE Simmons, Seiffert and Gunnell, 2016 ...................... 24 †Genus Aegyptonycteris Simmons, Seiffert and Gunnell, 2016 .............................. 24 †Aegyptonycteris knightae Simmons, Seiffert and Gunnell, 2016.................. 24 †Family NECROMANTIDAE Sigé, 2011 ............................................................................ 25 †Genus Necromantis Weithofer, 1887 ...................................................................... 25 †Necromantis fragmentum (Ravel, 2016) ....................................................... 25 SUBORDER PTEROPODIFORMI Van Cakenberghe, Kearney and Seamark, 2007 ...................... 26 INFRAORDER PTEROPODIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 ...... 27 Superfamily PTEROPODOIDEA Gray, 1821 ......................................................................... 27 Family PTEROPODIDAE Gray, 1821 ................................................................................. 27 Subfamily Eidolinae Almeida, Giannini and Simmons, 2016 .................................... 31 Genus Eidolon Rafinesque, 1815 .............................................................................. 31 Eidolon dupreanum (Schegel, 1867) ............................................................... 32 Eidolon helvum (Kerr, 1792) ............................................................................. 36 Eidolon helvum helvum (Kerr, 1792) .................................................................. 50 †Subfamily Propottininae Butler, 1984 ........................................................................ 51 †Genus Propotto Simpson, 1967 ............................................................................... 52 †Propotto leakeyi Simpson, 1967 ..................................................................... 52 Subfamily Pteropodinae Gray, 1821 ............................................................................ 52 Genus Pteropus Brisson, 1762 .................................................................................. 53 Pteropus aldabrensis True, 1893 ..................................................................... 55 Pteropus livingstonii Gray, 1866 ..................................................................... 57 Pteropus niger (Kerr, 1792) .............................................................................. 63 Pteropus rodricensis Dobson, 1878 ................................................................ 68 Pteropus rufus E. Geoffroy St.-Hilaire, 1803 .................................................... 71 Pteropus seychellensis A. Milne-Edwards, 1877 ............................................ 76 Pteropus subniger (Kerr, 1792) ........................................................................ 80 Pteropus voeltzkowi Matschie, 1909 ............................................................... 81 Subfamily Rousettinae Andersen, 1912 ...................................................................... 83 Genus Epomophorus Bennett, 1836 ......................................................................... 84 Epomophorus angolensis Gray, 1870 ............................................................. 85 Epomophorus anselli Bergmans and Van Strien, 2004 ................................... 87 Epomophorus crypturus Peters, 1852 ............................................................ 88 Epomophorus dobsonii Bocage, 1889 ............................................................ 91 Epomophorus gambianus (Ogilby, 1835) ........................................................ 92 Epomophorus gambianus gambianus (Ogilby, 1835) ........................................ 96 Epomophorus gambianus pousarguesi Trouessart, 1904.................................. 97 Epomophorus grandis (Sanborn, 1950) .......................................................... 98 Epomophorus intermedius (Hayman, 1963) ................................................... 99 Epomophorus labiatus (Temminck, 1837)..................................................... 100 Epomophorus minimus Claessen & De Vree, 1991 ...................................... 103 Epomophorus minor Dobson, 1880 ............................................................... 105 Epomophorus pusillus Peters, 1868 ............................................................. 106 Epomophorus wahlbergi (Sundevall, 1846) .................................................. 109 Genus Epomops Gray, 1866 .................................................................................... 117 Epomops buettikoferi (Matschie, 1899) ......................................................... 118 Epomops franqueti (Tomes, 1860) ................................................................ 120 Genus Hypsignathus H. Allen, 1862 ....................................................................... 123 Hypsignathus monstrosus H. Allen, 1862..................................................... 124 Genus Nanonycteris Matschie, 1899....................................................................... 129 Nanonycteris veldkampii (Jentink, 1888) ...................................................... 129 Genus Megaloglossus Pagenstecher, 1885 ........................................................... 131 Megaloglossus azagnyi Nesi, Kadjo and Hassanin, 2012 ............................. 132 Megaloglossus woermanni Pagenstecher, 1885 .......................................... 133 Genus Myonycteris Matschie, 1899 ........................................................................ 136 vi ISSN 1990-6471 Myonycteris angolensis (Bocage, 1898) ....................................................... 137 Myonycteris angolensis angolensis (Bocage, 1898) ........................................ 140 Myonycteris angolensis goliath (Bergmans, 1997) ........................................... 140 Myonycteris angolensis petraea (Bergmans, 1997) ......................................... 141 Myonycteris angolensis ruwenzorii (Eisentraut, 1965) ..................................... 142 Myonycteris angolensis smithii (Thomas, 1908)............................................... 142 Myonycteris brachycephala (Bocage, 1889) ................................................. 143 Myonycteris leptodon K. Andersen, 1908 ...................................................... 144 Myonycteris relicta Bergmans, 1980 .............................................................. 146 Myonycteris torquata (Dobson, 1878) ........................................................... 147 Myonycteris torquata torquata (Dobson, 1878) ................................................ 150 Myonycteris torquata wroughtoni K. Andersen, 1908 ....................................... 150 Genus Plerotes K. Andersen, 1910 .......................................................................... 151 Plerotes anchietae (Seabra, 1900) ................................................................. 151 Genus Rousettus Gray, 1821 .................................................................................. 153 †Rousettus pattersoni Gunnell and Manthi, 2018 ......................................... 155 Rousettus aegyptiacus (E. Geoffroy St.-Hilaire, 1810) .................................. 155 Rousettus madagascariensis G. Grandidier, 1929 ....................................... 176 Rousettus obliviosus Kock, 1978 .................................................................. 181 Genus Casinycteris Thomas, 1910 ......................................................................... 185 Casinycteris argynnis Thomas, 1910 ............................................................ 185 Casinycteris campomaanensis Hassanin, 2014 ........................................... 187 Casinycteris ophiodon (Pohle, 1943) ............................................................ 188 Scotonycteris bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, 2015 ................................................................. 190 Scotonycteris bergmansi bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, 2015 ............................................ 191 Scotonycteris bergmansi congoensis Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, 2015 ............................................ 191 Scotonycteris occidentalis Hayman, 1947.................................................... 191 Scotonycteris zenkeri Matschie, 1894 ........................................................... 192 Genus Stenonycteris Gray, 1870 ............................................................................ 194 Stenonycteris lanosus (Thomas, 1906) ......................................................... 194 †Genus Turkanycteris Gunnell and Manthi, 2018 ................................................... 196 †Turkanycteris harrisi Gunnell and Manthi, 2018 .......................................... 196 INFRAORDER RHINOLOPHIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 ... 196 Superfamily RHINOLOPHOIDEA J. E. Gray, 1825 ............................................................. 197 Family HIPPOSIDERIDAE Lydekker, 1891 ..................................................................... 197 Genus Asellia Gray, 1838 ........................................................................................ 200 Asellia italosomalica de Beaux, 1931 ............................................................ 200 Asellia patrizii de Beaux, 1931 ....................................................................... 201 Asellia tridens (E. Geoffroy St.-Hilaire, 1813) ................................................. 202 Genus Doryrhina Peters, 1871 ................................................................................ 206 Doryrhina camerunensis (Eisentraut, 1956) .................................................. 206 Doryrhina cyclops (Temminck, 1853) ............................................................ 208 Genus Hipposideros Gray, 1831 ............................................................................. 210 †Hipposideros africanum Ravel, 2016 .......................................................... 214 †Hipposideros amenhotepos Gunnell, Winkler, Miller, Head, El-Barkooky, Gawad, Sanders and Gingerich, 2015 ............................................................. 214 †Hipposideros kaumbului Wesselman, 1984 ................................................ 215 †Hipposideros vetus (Lavocat, 1961) ............................................................ 215 Hipposideros abae J.A. Allen, 1917 ............................................................... 215 Hipposideros beatus (K. Andersen, 1906) ..................................................... 217 Hipposideros caffer (Sundevall, 1846) ........................................................... 219 Hipposideros cf. centralis Andersen, 1906 ................................................... 226 Hipposideros curtus G.M. Allen, 1921 ........................................................... 226 Hipposideros fuliginosus (Temminck, 1853) ................................................ 227 Hipposideros jonesi Hayman, 1947 ............................................................... 229 Hipposideros lamottei Brosset, 1985 ............................................................. 230 Hipposideros marisae Aellen, 1954 ............................................................... 232 African Chiroptera Report 2020 vii Hipposideros megalotis (Heuglin, 1861) ....................................................... 233 Hipposideros ruber (Noack, 1893) ................................................................. 234 Hipposideros tephrus (Cabrera, 1906) .......................................................... 239 Genus Macronycteris Gray, 1866............................................................................ 240 †Macronycteris besaoka (Samonds, 2007) ................................................... 241 Macronycteris commersoni (E. Geoffroy St.-Hilaire, 1813) .......................... 241 Macronycteris cryptovalorona (Goodman, Schoeman, Rakotoarivelo and Willows-Munro, 2016) ....................................................................................... 245 Macronycteris gigas (Wagner, 1845) ............................................................. 246 Macronycteris vittatus (Peters, 1852) ............................................................ 249 †Genus Palaeophyllophora Revilliod, 1917 ............................................................ 252 †Palaeophyllophora tunisiensis (Ravel, 2016) ............................................. 253 Family MEGADERMATIDAE H. Allen, 1864 ................................................................... 253 Genus Cardioderma Peters, 1873 ........................................................................... 254 †Cardioderma leakeyi Gunnell, Butler, Greenwood and Simmons, 2015 ...... 254 Cardioderma cor (Peters, 1872) ..................................................................... 254 Genus Lavia Gray, 1838 ........................................................................................... 257 Lavia frons (E. Geoffroy St.-Hilaire, 1810) ...................................................... 257 Genus Megaderma E. Geoffroy St.-Hilaire, 1810 ..................................................... 260 †Megaderma gaillardi (Trouessart, 1898) ...................................................... 260 †Megaderma gigas (Lavocat, 1961) ............................................................... 261 †Megaderma jaegeri Sigé, 1976 ..................................................................... 261 †Genus Saharaderma Gunnell, Simons and Seiffert, 2008 ..................................... 261 †Saharaderma pseudovampyrus Gunnell, Simons and Seiffert, 2008 ......... 261 Genus Rhinolophus Lacépède, 1799 ...................................................................... 263 †Rhinolophus maghrebensis Gunnell, Eiting and Geraads, 2011 ................ 268 †Rhinolophus mellali Lavocat, 1961 .............................................................. 269 Rhinolophus adami Aellen and Brosset, 1968 ............................................... 269 Rhinolophus alcyone Temminck, 1853 .......................................................... 270 Rhinolophus blasii Peters, 1867 .................................................................... 272 Rhinolophus capensis Lichtenstein, 1823 ..................................................... 276 Rhinolophus clivosus Cretzschmar, 1828 ..................................................... 279 Rhinolophus cohenae Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 .................................................................................................... 286 Rhinolophus damarensis Roberts, 1946 ....................................................... 288 Rhinolophus darlingi K. Andersen, 1905 ....................................................... 289 Rhinolophus deckenii Peters, 1868 ............................................................... 292 Rhinolophus denti Thomas, 1904 .................................................................. 293 Rhinolophus eloquens K. Andersen, 1905 .................................................... 296 Rhinolophus euryale Blasius, 1853 ............................................................... 298 Rhinolophus ferrumequinum (Schreber, 1774) ............................................ 303 Rhinolophus fumigatus Rüppell, 1842 .......................................................... 310 Rhinolophus gorongosae Taylor, MacDonald, Goodman, Kearney, Cotterill, Stoffberg, Monadjem, Schoeman, Guyton, Naskrecki and Richards, 2018 ..... 313 Rhinolophus guineensis Eisentraut, 1960..................................................... 314 Rhinolophus hildebrandtii Peters, 1878 ........................................................ 316 Rhinolophus hilli Aellen, 1973........................................................................ 319 Rhinolophus hillorum Koopman, 1989 .......................................................... 320 Rhinolophus hipposideros (Bechstein, 1800) ............................................... 321 Rhinolophus horaceki Benda and Vallo, 2012 .............................................. 325 Rhinolophus kahuzi Fahr and Kerbis Peterhans, 2013 ................................. 326 Rhinolophus landeri Martin, 1838 .................................................................. 327 Rhinolophus lobatus Peters, 1852 ................................................................ 329 Rhinolophus mabuensis Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 .................................................................................................... 331 Rhinolophus maclaudi Pousargues, 1898 ..................................................... 332 Rhinolophus maendeleo Kock, Csorba and Howell, 2000 ............................ 333 Rhinolophus mehelyi Matschie, 1901 ............................................................ 334 Rhinolophus mossambicus Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 ............................................................................................. 339 viii ISSN 1990-6471 Rhinolophus rhodesiae Roberts, 1946 .......................................................... 340 Rhinolophus ruwenzorii J. Eric Hill, 1942 ...................................................... 341 Rhinolophus sakejiensis Cotterill, 2002 ........................................................ 342 Rhinolophus silvestris Aellen, 1959 .............................................................. 343 Rhinolophus simulator K. Andersen, 1904 .................................................... 344 Rhinolophus smithersi Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 .................................................................................................... 347 Rhinolophus swinnyi Gough, 1908 ................................................................ 350 Rhinolophus willardi Kerbis Peterhans and Fahr, 2013 ................................ 352 Rhinolophus ziama Fahr, Vierhaus, Hutterer and Kock, 2002 ....................... 352 Family RHINONYCTERIDAE J. E. Gray, 1866 ................................................................ 353 †Genus Brachipposideros Sigé, 1968 .................................................................... 354 †Genus Brevipalatus Hand and Archer, 2005 ......................................................... 354 Genus Cloeotis Thomas, 1901 ................................................................................. 354 Cloeotis percivali Thomas, 1901 .................................................................... 355 Genus Paratriaenops Benda and Vallo, 2009 ......................................................... 358 Paratriaenops auritus (G. Grandidier, 1912).................................................. 358 Paratriaenops furculus (Trouessart, 1907) .................................................... 360 Paratriaenops pauliani (Goodman and Ranivo, 2008) .................................. 362 Genus Triaenops Dobson, 1871 .............................................................................. 364 †Triaenops goodmani Samonds, 2007 .......................................................... 364 Triaenops afer Peters, 1877............................................................................ 365 Triaenops menamena Goodman and Ranivo, 2009 ...................................... 366 Superfamily RHINOPOMATOIDEA Dobson, 1872 ............................................................. 370 Family RHINOPOMATIDAE Dobson, 1872 ..................................................................... 370 †Genus Qarunycteris Gunnell, Simons and Seiffert, 2008 ..................................... 371 †Qarunycteris moerisae Gunnell, Simons and Seiffert, 2008........................ 372 Genus Rhinopoma E. Geoffroy St.-Hilaire, 1818 ..................................................... 372 Rhinopoma cystops Thomas, 1903 ............................................................... 373 Rhinopoma macinnesi Hayman, 1937 ........................................................... 377 Rhinopoma microphyllum (Brünnich, 1782).................................................. 378 †Family TANZANYCTERIDIDAE Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 .................................................... 381 †Genus Tanzanycteris Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 ............................................... 381 †Tanzanycteris mannardi Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 .......................... 381 SUBORDER VESPERTILIONIFORMI Van Cakenberghe, Kearney and Seamark, 2007.............. 383 INFRAORDER NOCTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 ..... 384 Superfamily NOCTILIONOIDEA Gray, 1821 ....................................................................... 384 Family MYZOPODIDAE Thomas, 1904 ........................................................................... 385 Genus Myzopoda Milne-Edwards and A. Grandidier, 1878 ..................................... 385 †Myzopoda africana Gunnell, Butler, Greenwood and Simmons, 2015 ........ 386 Myzopoda aurita Milne-Edwards and A. Grandidier, 1878 ............................. 386 Myzopoda schliemanni Goodman, Rakotondraparany and Kofoky, 2007 .... 388 †Genus Phasmatonycteris Gunnell, Simmons and Seiffert, 2014 .......................... 390 †Phasmatonycteris butleri Gunnell, Simmons and Seiffert, 2014................. 390 †Phasmatonycteris phiomensis Gunnell, Simmons and Seiffert, 2014 ....... 391 INFRAORDER NYCTERIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 .......... 391 Superfamily NYCTEROIDEA Van der Hoeven, 1855 ......................................................... 391 Family EMBALLONURIDAE Gervais, 1855 .................................................................... 392 †Genus Dhofarella Sigé, Thomas, Sen, Gheerbrant, Roger and Al-Sulaimani, 1985 ................................................................................................................................... 393 †Dhofarella sigei Gunnell, Simons and Seiffert, 2008 .................................... 393 †Dhofarella thaleri Sigé et al., 1994 ............................................................... 394 Subfamily Emballonurinae Gervais, 1855 ................................................................. 394 Genus Coleura Peters, 1867 .................................................................................... 394 †Coleura muthokai Wesselman, 1984 ........................................................... 395 Coleura afra (Peters, 1852) ............................................................................. 395 Coleura gallarum Thomas, 1915 .................................................................... 399 African Chiroptera Report 2020 ix Coleura kibomalandy Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, 2012 ................................................. 399 Coleura seychellensis Peters, 1868 .............................................................. 401 Genus Paremballonura Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, 2012 ..................................................................... 404 Paremballonura atrata (Peters, 1874) ............................................................ 404 Paremballonura tiavato (Goodman, Cardiff, Ranivo, Russell and Yoder, 2006) .......................................................................................................................... 406 †Genus Pseudovespertiliavus Ravel, 2016 ........................................................... 407 †Pseudovespertiliavus parva Ravel, 2016 .................................................... 407 Subfamily Taphozoinae Jerdon, 1867 ....................................................................... 408 Genus Saccolaimus Temminck, 1838 ..................................................................... 408 †Saccolaimus abitus (Wesselman, 1984) ...................................................... 408 †Saccolaimus kenyensis Gunnell and Manthi, 2018 ..................................... 409 Saccolaimus peli (Temminck, 1853) .............................................................. 409 Genus Taphozous E. Geoffroy St.-Hilaire, 1818 ...................................................... 410 †Taphozous incognita (Butler & Hopwood, 1957) ......................................... 412 Taphozous hamiltoni Thomas, 1920 .............................................................. 412 Taphozous hildegardeae Thomas, 1909 ....................................................... 413 Taphozous mauritianus E. Geoffroy St.-Hilaire, 1818 ................................... 414 Taphozous nudiventris Cretzschmar, 1830 ................................................... 418 Taphozous perforatus E. Geoffroy St.-Hilaire, 1818 ...................................... 421 Taphozous perforatus perforatus E. Geoffroy St.-Hilaire, 1818 ....................... 425 Taphozous perforatus sudani Thomas, 1915 ................................................... 425 †Genus Vespertiliavus Schlosser, 1887 ................................................................. 425 †Vespertiliavus aenigma (Ravel, 2016) ......................................................... 426 †Vespertiliavus kasserinensis (Ravel, 2016) ................................................ 426 Family NYCTERIDAE Van der Hoeven, 1855 ................................................................. 426 †Genus Khoufechia Ravel, 2016 ............................................................................. 427 †Khoufechia gunnelli Ravel, 2016 ................................................................. 427 Genus Nycteris G. Cuvier and E. Geoffroy, 1795 .................................................... 427 Nycteris arge Thomas, 1903 ........................................................................... 429 Nycteris aurita (K. Andersen, 1912) ............................................................... 431 Nycteris cf. parisii De Beaux, 1924 ................................................................ 432 Nycteris gambiensis (K. Andersen, 1912) ..................................................... 433 Nycteris grandis Peters, 1865 ........................................................................ 434 Nycteris hispida (Schreber, 1774) .................................................................. 437 Nycteris intermedia Aellen, 1959 ................................................................... 440 Nycteris macrotis Dobson, 1876 .................................................................... 441 Nycteris madagascariensis G. Grandidier, 1937 .......................................... 445 Nycteris major (K. Andersen, 1912) ............................................................... 446 Nycteris nana (K. Andersen, 1912) ................................................................. 448 Nycteris parisii (de Beaux, 1924) ................................................................... 449 Nycteris thebaica E. Geoffroy St.-Hilaire, 1818 .............................................. 450 Nycteris vinsoni Dalquest, 1965 ..................................................................... 460 Nycteris woodi K. Andersen, 1914 ................................................................. 461 INFRAORDER VESPERTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 .................................................................................................................................................... 462 Superfamily MOLOSSOIDEA Gervais, 1856 ...................................................................... 463 Family MOLOSSIDAE Gervais, 1856 .............................................................................. 463 Subfamily Molossinae Gervais, 1856 ......................................................................... 465 Genus Mops Lesson, 1842 ....................................................................................... 466 †Mops kerio Gunnell and Manthi, 2018 .......................................................... 467 †Mops rusingae (Arroyo-Cabrales, Gregorin, Schlitter and Walker, 2002) .... 467 †Mops turkwellensis Gunnell and Manthi, 2018 ............................................ 467 Subgenus Mops (Chaerephon) Dobson, 1874 ..................................................... 468 Mops (Chaerephon) aloysiisabaudiae (Festa, 1907) ................................... 470 Mops (Chaerephon) ansorgei (Thomas, 1913) ............................................. 471 Mops (Chaerephon) atsinanana (Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, 2010) ...................................................... 473 x ISSN 1990-6471 Mops (Chaerephon) bemmeleni (Jentink, 1879) ........................................... 475 Mops (Chaerephon) bivittatus (Heuglin, 1861) ............................................. 477 Mops (Chaerephon) chapini (J.A. Allen, 1917).............................................. 479 Mops (Chaerephon) gallagheri (Harrison, 1975) ........................................... 481 Mops (Chaerephon) jobimena (Goodman and Cardiff, 2004) ....................... 482 Mops (Chaerephon) leucogaster (A. Grandidier, 1869) ................................ 483 Mops (Chaerephon) major (Trouessart, 1897) .............................................. 485 Mops (Chaerephon) nigeriae (Thomas, 1913)............................................... 487 Mops (Chaerephon) nigeriae nigeriae (Thomas, 1913) ................................... 489 Mops (Chaerephon) nigeriae spillmanni (Monard, 1933) ................................. 490 Mops (Chaerephon) pumilus (Cretzschmar, 1826) ....................................... 490 Mops (Chaerephon) pusillus (Miller, 1902) ................................................... 502 Mops (Chaerephon) russatus (J.A. Allen, 1917) ........................................... 504 Mops (Chaerephon) tomensis (Juste and Ibáñez, 1993) .............................. 505 Subgenus Mops (Mops) Lesson, 1842 ................................................................. 506 Mops (Mops) condylurus (A. Smith, 1833) .................................................... 506 Mops (Mops) congicus J.A. Allen, 1917 ........................................................ 513 Mops (Mops) demonstrator (Thomas, 1903) ................................................. 514 Mops (Mops) leucostigma (G.M. Allen, 1918) ............................................... 515 Mops (Mops) midas (Sundevall, 1843) ........................................................... 517 Mops (Mops) midas miarensis (A. Grandidier, 1869) ....................................... 521 Mops (Mops) midas midas (Sundevall, 1843) .................................................. 521 Mops (Mops) niangarae J.A. Allen, 1917 ....................................................... 522 Mops (Mops) niveiventer Cabrera and Ruxton, 1926 .................................... 523 Mops (Mops) trevori J.A. Allen, 1917 ............................................................. 525 Subgenus Mops (Xiphonycteris) Dollman, 1911 ................................................... 526 Mops (Xiphonycteris) bakarii Stanley, 2009 ................................................. 526 Mops (Xiphonycteris) brachypterus (Peters, 1852) ..................................... 527 Mops (Xiphonycteris) nanulus J.A. Allen, 1917 ............................................ 529 Mops (Xiphonycteris) petersoni (El Rayah, 1981) ........................................ 531 Mops (Xiphonycteris) spurrelli (Dollman, 1911) ........................................... 532 Mops (Xiphonycteris) thersites (Thomas, 1903) .......................................... 533 Genus Mormopterus Peters, 1865 .......................................................................... 535 Mormopterus acetabulosus (Hermann, 1804) .............................................. 536 Mormopterus acetabulosus acetabulosus (Hermann, 1804) ............................ 538 Mormopterus acetabulosus natalensis (A. Smith, 1847) .................................. 539 Mormopterus francoismoutoui Goodman, Jansen Van Vuuren, Ratrimomanarivo, Probst, Bowie, 2008 ............................................................ 539 Mormopterus jugularis (Peters, 1865) ........................................................... 543 Genus Myopterus E. Geoffroy St.-Hilaire, 1818 ...................................................... 545 Myopterus daubentonii Desmarest, 1820 ...................................................... 545 Myopterus whitleyi (Scharff, 1900) ................................................................ 547 Genus Otomops Thomas, 1913 ............................................................................... 548 Otomops harrisoni Ralph, Richards, Taylor, Napier and Lamb, 2015 ........... 548 Otomops madagascariensis Dorst, 1953 ...................................................... 550 Otomops martiensseni (Matschie, 1897) ....................................................... 552 Otomops martiensseni icarus Chubb, 1917 ..................................................... 556 Otomops martiensseni martiensseni (Matschie, 1897) .................................... 556 Genus Platymops Thomas, 1906............................................................................. 556 Platymops setiger (Peters, 1878) ................................................................... 557 Platymops setiger macmillani Thomas, 1906 ................................................... 558 Platymops setiger setiger (Peters, 1878) ......................................................... 558 Genus Sauromys Roberts, 1917 .............................................................................. 559 Sauromys petrophilus (Roberts, 1917) .......................................................... 560 Sauromys petrophilus erongensis (Roberts, 1946) .......................................... 562 Sauromys petrophilus haagneri (Roberts, 1917) .............................................. 563 Sauromys petrophilus petrophilus (Roberts, 1917) .......................................... 563 Sauromys petrophilus umbratus (Shortridge & Carter, 1938) .......................... 564 Genus Tadarida Rafinesque, 1814........................................................................... 564 †Tadarida engesseri Rachl, 1983 ................................................................... 565 African Chiroptera Report 2020 xi Tadarida aegyptiaca (E. Geoffroy St.-Hilaire, 1818) ...................................... 565 Tadarida aegyptiaca aegyptiaca (E. Geoffroy St.-Hilaire, 1818) ...................... 569 Tadarida aegyptiaca bocagei (Seabra, 1900) .................................................. 570 Tadarida fulminans (Thomas, 1903) .............................................................. 570 Tadarida lobata (Thomas, 1891) ..................................................................... 572 Tadarida teniotis (Rafinesque, 1814) ............................................................. 573 Tadarida ventralis (Heuglin, 1861) ................................................................. 578 Superfamily VESPERTILIONOIDEA Gray, 1821 ................................................................. 580 Family CISTUGONIDAE Lack, Roehrs, Stanley, Ruedi and Van den Bussche, 2010 581 Genus Cistugo Thomas, 1912 ................................................................................. 581 Cistugo lesueuri Roberts, 1919 ...................................................................... 582 Cistugo seabrae Thomas, 1912 ...................................................................... 584 †Family INDETERMINATE ............................................................................................... 586 †Genus Chambinycteris Ravel, 2016...................................................................... 586 †Chambinycteris pusilli Ravel, 2016 ............................................................. 586 †Genus Drakonycteris Ravel, 2016......................................................................... 586 †Drakonycteris glibzegdouensis Ravel, 2016 .............................................. 586 Family MINIOPTERIDAE Dobson, 1875 .......................................................................... 587 Genus Miniopterus Bonaparte, 1837 ....................................................................... 588 †Miniopterus horaceki Gunnell, Eiting and Geraads, 2011 ........................... 591 Miniopterus aelleni Goodman, Maminirina, Weyeneth, Bradman, Christidis, Ruedi and Appleton, 2009 ................................................................................ 591 Miniopterus africanus Sanborn, 1936 ............................................................ 593 Miniopterus ambohitrensis Goodman, Ramasindrazana, Naughton and Appleton, 2015 .................................................................................................. 594 Miniopterus arenarius Heller, 1912 ................................................................ 595 Miniopterus brachytragos Goodman, Maminirina, Bradman, Christidis and Appleton, 2009 .................................................................................................. 596 Miniopterus cf. inflatus Monadjem, Shapiro, Richards, Karabulut, Crawley, Broman Nielsen, Hansen, Bohmann and Mourier, 2020 .................................. 599 Miniopterus egeri Goodman, Ramasindrazana, Maminirina, Schoeman and Appleton, 2011 .................................................................................................. 599 Miniopterus fraterculus Thomas and Schwann, 1906 ................................... 600 Miniopterus gleni Peterson, Eger and Mitchell, 1995 .................................... 603 Miniopterus griffithsi Goodman, Maminirina, Bradman, Christidis and Appleton, 2009 .................................................................................................. 604 Miniopterus griveaudi Harrison, 1959 ............................................................ 606 Miniopterus inflatus Thomas, 1903 ............................................................... 608 Miniopterus inflatus inflatus Thomas, 1903 ...................................................... 611 Miniopterus inflatus rufus Sanborn, 1936 ......................................................... 611 Miniopterus maghrebensis Puechmaille, Allegrini, Benda, Bilgin, Ibañez and Juste, 2014 ....................................................................................................... 611 Miniopterus mahafaliensis Goodman, Bradman, Christides and Appleton, 2009 .................................................................................................................. 613 Miniopterus majori Thomas, 1906 ................................................................. 617 Miniopterus manavi Thomas, 1906 ................................................................ 619 Miniopterus minor Peters, 1867 ..................................................................... 621 Miniopterus mossambicus Monadjem, Goodman, Stanley and Appleton, 2013 .......................................................................................................................... 623 Miniopterus natalensis (A. Smith, 1833) ........................................................ 624 Miniopterus newtoni Bocage, 1889 ............................................................... 630 Miniopterus nimbae Monadjem, Shapiro, Richards, Karabulut, Crawley, Nielsen, Hansen, Bohmann and Mourier, 2020 ............................................... 631 Miniopterus petersoni Goodman, Bradman, Maminirina, Ryan, Christidis & Appleton, 2008 .................................................................................................. 632 Miniopterus schreibersii (Kuhl, 1817) ........................................................... 634 Miniopterus sororculus Goodman, Ryan, Maminirina, Fahr, Christidis and Appleton, 2007 .................................................................................................. 639 Miniopterus villiersi Aellen, 1956 ................................................................... 641 Miniopterus "incertae-sedis" ......................................................................... 642 xii ISSN 1990-6471 †Family PHILISIDAE Sigé, 1985 ...................................................................................... 642 †Genus Dizzya Sigé, 1991 ....................................................................................... 643 †Dizzya exsultans Sigé, 1991 ......................................................................... 643 †Genus Philisis Sigé, 1985 ...................................................................................... 643 †Philisis sphingis Sigé, 1985 ......................................................................... 643 †Genus Witwatia Gunnell, Simons and Seiffert, 2008 ............................................. 644 †Witwatia eremicus Gunnell, Simons and Seiffert, 2008 ............................... 644 †Witwatia schlosseri Gunnell, Simons and Seiffert, 2008 ............................. 644 †Witwatia sigei Ravel, Marivaux, Tabuce, Bel Haj Ali, Essid and Vianey-Liaud, 2012 .................................................................................................................. 645 Family VESPERTILIONIDAE Gray, 1821 ........................................................................ 645 Subfamily Kerivoulinae Miller, 1907 .......................................................................... 648 †Genus Chamtwaria Butler, 1984 ............................................................................ 649 †Chamtwaria pickfordi Butler, 1984 ............................................................... 649 Genus Kerivoula Gray, 1842 .................................................................................... 649 Kerivoula africana Dobson, 1878 ................................................................... 650 Kerivoula argentata Tomes, 1861 .................................................................. 651 Kerivoula cuprosa Thomas, 1912 .................................................................. 653 Kerivoula eriophora (Heuglin, 1877) .............................................................. 654 Kerivoula lanosa (A. Smith, 1847) .................................................................. 655 Kerivoula phalaena Thomas, 1912 ................................................................. 657 Kerivoula smithii Thomas, 1880 ..................................................................... 658 Subfamily Myotinae Tate, 1942 ................................................................................... 659 †Genus Khonsunycteris Gunnell, Simons and Seiffert, 2008 ................................ 660 †Khonsunycteris aegypticus Gunnell, Simons and Seiffert, 2008................ 660 Genus Myotis Kaup, 1829 ........................................................................................ 661 †Myotis darelbeidensis Gunnell, Eiting and Geraads, 2011 .......................... 667 Myotis anjouanensis Dorst, 1960 ................................................................... 667 Myotis bocagii (Peters, 1870) ......................................................................... 668 Myotis bocagii bocagii (Peters, 1870) ............................................................... 671 Myotis bocagii cupreolus Thomas, 1904 .......................................................... 671 Myotis capaccinii (Bonaparte, 1837) .............................................................. 671 Myotis dieteri M. Happold, 2005 ..................................................................... 675 Myotis emarginatus (E. Geoffroy St.-Hilaire, 1806) ....................................... 676 Myotis goudoti (A. Smith, 1834) ..................................................................... 679 Myotis morrisi Hill, 1971 ................................................................................. 681 Myotis mystacinus (Kuhl, 1817) ..................................................................... 682 Myotis punicus Felten, 1977........................................................................... 684 Myotis scotti Thomas, 1927 ............................................................................ 689 Myotis tricolor (Temminck, 1832) ................................................................... 690 Myotis welwitschii (Gray, 1866) ..................................................................... 693 Myotis zenatius Ibáñez, Juste, Salicini, Puechmaille and Ruedi, 2019 .......... 696 Subfamily Scotophilinae Van Cakenberghe and Seamark, 2008 ............................ 698 †Genus Scotophilisis Horácek, Fejfar and Hulva, 2006 ......................................... 698 †Scotophilisis libycus Horácek, Fejfar and Hulva, 2006 ............................... 698 Genus Scotophilus Leach, 1821 ............................................................................. 699 Scotophilus altilis G.M. Allen, 1914 ............................................................... 701 Scotophilus andrewreborii Brooks and Bickham, 2014 ................................ 701 Scotophilus borbonicus (E. Geoffroy St.-Hilaire, 1803) ................................ 702 Scotophilus dinganii (A. Smith, 1833) ........................................................... 704 Scotophilus ejetai Brooks and Bickham, 2014 ............................................... 709 Scotophilus leucogaster (Cretzschmar, 1826) .............................................. 709 Scotophilus livingstonii Brooks and Bickham, 2014 ..................................... 712 Scotophilus marovaza Goodman, Ratrimomanarivo and Randrianandrianina, 2006 .................................................................................................................. 713 Scotophilus nigrita (Schreber, 1774) ............................................................. 715 Scotophilus nigritellus de Winton, 1899 ........................................................ 717 Scotophilus nucella Robbins, 1983 ............................................................... 718 Scotophilus nux Thomas, 1904 ...................................................................... 719 Scotophilus robustus A. Milne-Edwards, 1881 ............................................. 721 African Chiroptera Report 2020 xiii Scotophilus tandrefana Goodman, Jenkins and Ratrimomanarivo, 2005 ..... 723 Scotophilus trujilloi Brooks and Bickham, 2014 ............................................ 724 Scotophilus viridis (Peters, 1852) .................................................................. 725 Scotophilus "incertae-sedis" ........................................................................ 727 †Genus Vampyravus Schlosser, 1910..................................................................... 727 †Vampyravus orientalis Schlosser, 1910 ...................................................... 728 Subfamily Vespertilioninae Gray, 1821 ..................................................................... 728 Genus Eptesicus Rafinesque, 1820......................................................................... 730 Subgenus Eptesicus (Cnephaeus) Kaup, 1829 .................................................... 733 Eptesicus (Cnephaeus) bottae (Peters, 1869) .............................................. 733 Eptesicus (Cnephaeus) hottentotus (A. Smith, 1833) .................................. 735 Eptesicus (Cnephaeus) isabellinus (Temminck, 1840) ................................ 740 Eptesicus (Cnephaeus) platyops (Thomas, 1901) ....................................... 744 Subgenus Eptesicus (Rhinopterus) Miller, 1906 ................................................... 745 Eptesicus (Rhinopterus) floweri (de Winton, 1901) ...................................... 746 Genus Glauconycteris Dobson, 1875 ..................................................................... 747 Glauconycteris alboguttata J.A. Allen, 1917 ................................................. 748 Glauconycteris argentata (Dobson, 1875) ..................................................... 749 Glauconycteris atra Hassanin, Colombo, Gembu, Merle, Tu, Görföl, Musaba Akawa, Csorba, Kearney, Monadjem and Ing, 2017 ........................................ 751 Glauconycteris beatrix Thomas, 1901 ........................................................... 751 Glauconycteris cf. beatrix Hassanin, Colombo, Gembu, Merle, Tu, Görföl, Musaba Akawa, Csorba, Kearney, Monadjem and Ing, 2017 .......................... 753 Glauconycteris cf. humeralis Hassanin, Colombo, Gembu, Merle, Tu, Görföl, Musaba Akawa, Csorba, Kearney, Monadjem and Ing, 2017 .......................... 754 Glauconycteris curryae Eger and Schlitter, 2001 .......................................... 754 Glauconycteris egeria Thomas, 1913 ............................................................ 755 Glauconycteris gleni Peterson and Smith, 1973............................................ 756 Glauconycteris humeralis J.A. Allen, 1917 ................................................... 757 Glauconycteris kenyacola Peterson, 1982 .................................................... 758 Glauconycteris machadoi Hayman, 1963 ..................................................... 759 Glauconycteris poensis (Gray, 1842) ............................................................ 760 Glauconycteris superba Hayman, 1939 ........................................................ 761 Glauconycteris variegata (Tomes, 1861) ...................................................... 763 Genus Nyctalus Bowdich, 1825 ............................................................................... 766 Nyctalus azoreum (Thomas, 1901) ................................................................ 767 Nyctalus lasiopterus (Schreber, 1780) .......................................................... 769 Nyctalus leisleri (Kuhl, 1817) .......................................................................... 772 Genus Pipistrellus Kaup, 1829 ................................................................................ 775 Subgenus Pipistrellus (Afropipistrellus) Thorn, Kock and Cuisin, 2007 ................ 778 Pipistrellus (Afropipistrellus) grandidieri (Dobson, 1876) ........................... 778 Subgenus Pipistrellus (Pipistrellus) Kaup, 1829 ................................................... 780 Pipistrellus (Pipistrellus) aero Heller, 1912 .................................................. 780 Pipistrellus (Pipistrellus) hanaki Hulva and Benda, 2004 ............................ 781 Pipistrellus (Pipistrellus) hesperidus (Temminck, 1840) ............................. 784 Pipistrellus (Pipistrellus) hesperidus broomi Roberts, 1948 ............................. 787 Pipistrellus (Pipistrellus) hesperidus hesperidus (Temminck, 1840) ................ 788 Pipistrellus (Pipistrellus) hesperidus subtilis (Sundevall, 1846) ....................... 788 Pipistrellus (Pipistrellus) inexspectatus Aellen, 1959 ................................. 789 Pipistrellus (Pipistrellus) kuhlii (Kuhl, 1817) ................................................ 790 Pipistrellus (Pipistrellus) maderensis (Dobson, 1878) ................................ 798 Pipistrellus (Pipistrellus) nanulus Thomas, 1904......................................... 800 Pipistrellus (Pipistrellus) permixtus Aellen, 1957 ........................................ 802 Pipistrellus (Pipistrellus) pipistrellus (Schreber, 1774) ............................... 803 Pipistrellus (Pipistrellus) pygmaeus (Leach, 1825) ..................................... 807 Pipistrellus (Pipistrellus) raceyi Bates, Ratrimomanarivo, Harrison and Goodman, 2006 ................................................................................................ 809 Pipistrellus (Pipistrellus) rueppellii (J.B. Fischer, 1829) .............................. 810 Pipistrellus (Pipistrellus) rueppellii fuscipes Thomas, 1913 .............................. 814 Pipistrellus (Pipistrellus) rueppellii pulcher (Dobson, 1875) ............................. 815 xiv ISSN 1990-6471 Pipistrellus (Pipistrellus) rueppellii senegalensis Dorst, 1960 .......................... 816 Pipistrellus (Pipistrellus) rueppellii vernayi Roberts, 1932 ................................ 816 Pipistrellus (Pipistrellus) rusticus (Tomes, 1861) ........................................ 817 Pipistrellus *(Pipistrellus)* sp. aff. *kuhlii* Kuhl, 1817 ................................. 820 Pipistrellus (Pipistrellus) "incertae-sedis" ................................................... 820 Genus Scotoecus Thomas, 1901............................................................................. 821 †Scotoecus olduvensis Gunnell, Butler, Greenwood and Simmons, 2015 ... 821 Scotoecus albigula Thomas, 1909 ................................................................. 822 Scotoecus albofuscus (Thomas, 1890) ......................................................... 823 Scotoecus hindei Thomas, 1901 .................................................................... 824 Scotoecus hirundo (de Winton, 1899) ........................................................... 826 Genus Barbastella Gray, 1821 ................................................................................. 828 Barbastella barbastellus (Schreber, 1774) .................................................... 828 Barbastella leucomelas (Cretzschmar, 1826) ................................................ 831 Genus Otonycteris Peters, 1859 ............................................................................. 833 Otonycteris hemprichii Peters, 1859 ............................................................. 834 Genus Plecotus E. Geoffroy St.-Hilaire, 1818 .......................................................... 838 Plecotus balensis Kruskop & Lavrenchenko, 2000 ........................................ 839 Plecotus christii Gray, 1838 ........................................................................... 841 Plecotus gaisleri Benda, Kiefer, Hanak & Veith, 2004 ................................... 843 Plecotus teneriffae Barrett-Hamilton, 1907 .................................................... 844 Genus Hypsugo Kolenati, 1856 ............................................................................... 846 Hypsugo ariel (Thomas, 1904) ....................................................................... 847 Hypsugo bemainty Goodman, Rakotondramanana, Ramasindrazana, Kearney, Monadjem, Schoeman, Taylor, Naughton and Appleton, 2015........................ 849 Hypsugo cf. eisentrauti Hill, 1968 .................................................................. 850 Hypsugo musciculus (Thomas, 1913) ........................................................... 850 Hypsugo savii (Bonaparte, 1837) ................................................................... 852 Genus Laephotis Thomas, 1901 .............................................................................. 856 Laephotis aff. guineensis Bocage, 1889 ....................................................... 858 Laephotis anchietae (Seabra, 1900) .............................................................. 858 Laephotis angolensis Monard, 1935 .............................................................. 860 Laephotis botswanae Setzer, 1971 ................................................................ 863 Laephotis brunneus (Thomas, 1880) ............................................................. 867 Laephotis capensis (A. Smith, 1829) ............................................................. 869 Laephotis guineensis (Bocage, 1889) ........................................................... 876 Laephotis helios (Heller, 1912) ...................................................................... 878 Laephotis humbloti (A. Milne-Edwards, 1881) ............................................... 879 Laephotis isabella (Decher, Hutterer and Monadjem, 2016) ......................... 880 Laephotis malagasyensis (Peterson, Eger and Mitchell, 1995) .................... 881 Laephotis matroka (Thomas and Schwann, 1905) ........................................ 882 Laephotis namibensis Setzer, 1971 .............................................................. 884 Laephotis nanus (Peters, 1852) ..................................................................... 887 Laephotis rendalli (Thomas, 1889) ................................................................ 895 Laephotis robertsi (Goodman, Taylor, Ratrimomanarivo and Hoofer, 2012) 899 Laephotis roseveari (Monadjem, Richards, Taylor and Stoffberg, 2013) ...... 900 Laephotis somalicus (Thomas, 1901) ............................................................ 901 Laephotis stanleyi (Goodman, Kearney, Ratsimbazafy and Hassanin, 2017) .......................................................................................................................... 904 Laephotis tenuipinnis (Peters, 1872) ............................................................. 906 Laephotis wintoni Thomas, 1901 ................................................................... 909 Laephotis zuluensis (Roberts, 1924) ............................................................. 913 Laephotis "incertae-sedis" ............................................................................ 916 Genus Mimetillus Thomas, 1904 ............................................................................. 917 Mimetillus moloneyi (Thomas, 1891) ............................................................. 917 Mimetillus thomasi Hinton, 1920 .................................................................... 918 Genus Nycticeinops Hill & Harrison, 1987 .............................................................. 919 †Nycticeinops serengetiensis Gunnell, Butler, Greenwood and Simmons, 2015 .................................................................................................................. 920 Nycticeinops schlieffenii (Peters, 1859) ........................................................ 920 African Chiroptera Report 2020 xv Nycticeinops "incertae-sedis" ....................................................................... 924 Genus Parahypsugo Hutterer, Decher, Monadjem and Astrin, 2019 ...................... 924 Parahypsugo bellieri (De Vree, 1972) ............................................................ 925 Parahypsugo crassulus (Thomas, 1904) ....................................................... 926 Parahypsugo eisentrauti (Hill, 1968) ............................................................. 927 Parahypsugo happoldorum Hutterer, Decher, Monadjem and Astrin, 2019 . 929 Parahypsugo macrocephalus Hutterer and Kerbis Peterhans, 2020 ........... 929 Acknowledgements ........................................................................................................................... 931 Glossary ............................................................................................................................................. 933 References ......................................................................................................................................... 939 Appendices ...................................................................................................................................... 1136 Appendix 1: Current Taxonomy................................................................................................. 1136 Appendix 2: Voucher specimens............................................................................................... 1145 Appendix 2a: Museum Acronyms and Number of Specimens .......................................... 1145 Appendix 2b: Voucher Specimen Details ............................................................................. 1178 Appendix 2c: Voucher Specimens per Museum ................................................................. 1570 Appendix 2d: Voucher Specimens per Museum and Country ........................................... 1715 Appendix 2e: Voucher Specimens per Country .................................................................. 1802 Appendix 2f: Voucher Specimens from Protected Areas ................................................... 1899 Appendix 2f1: Sorted by Country, Protected Area, Taxon ............................................. 1899 Appendix 2f2: Sorted by Taxon, Country, Protected Area ............................................. 1937 Appendix 2f3: Number of specimens of the given taxa collected in and outside Protected Areas .................................................................................................................. 1984 Appendix 2f4: Number of specimens of the given taxa recently [in the last 20 years] collected in and outside Protected Areas ........................................................................ 1992 Appendix 2f5: Number of localities in which the given taxa was collected in and outside Protected Areas .................................................................................................................. 1998 Appendix 2f6: Number of localities in which the given taxa was recently [in the last 20 years] collected in and outside Protected Areas ............................................................ 2006 Appendix 2g: Distribution Maps per Taxon ......................................................................... 2012 Appendix 2h: Species Richness Maps per Country ............................................................ 2341 Appendix 2i: Distribution per quarter degree ...................................................................... 2401 Appendix 2j: Accumulation curves ....................................................................................... 3162 Appendix 3: Synonyms .............................................................................................................. 3190 Appendix 3a: Synonyms by Name ........................................................................................ 3190 Appendix 3b: Synonyms by Author ...................................................................................... 3213 Appendix 3c: Synonyms by Publication Date ..................................................................... 3246 Appendix 3d: Synonyms by Country of Type Specimen .................................................... 3268 Appendix 3e: Common Names in Alphabetical Order ........................................................ 3291 Appendix 3f: Common Names by Language ....................................................................... 3359 Appendix 3g: Original Descriptions...................................................................................... 3429 Appendix 4: Collector Information ............................................................................................ 3739 Appendix 4a: Follow the Collector ........................................................................................ 3739 Appendix 4b: Chronological .................................................................................................. 4292 Appendix 4c: Per Country and Locality................................................................................ 5656 Appendix 4d: Collector Biographies..................................................................................... 5922 Appendix 5: Keys from the Literature ........................................................................................... 5989 Appendix 5a: Dobson (1877). A Monograph of the Group Molossi. ............................... 5989 Appendix 5b: Dobson (1878). Catalogue of the Chiroptera in the collection of the British Museum. .................................................................................................................................. 5992 Appendix 5c: Lataste (1885). Etude de la faune des Vertébrés de Barbarie (Algérie, Tunisie et Maroc) .................................................................................................................... 6019 Appendix 5d: Monticelli (1889). LIX. Some remarks on the genus Taphozous. .......... 6022 Appendix 5e: Matschie (1895). Die Säugethiere Deutsch-Ost-Afrikas ........................... 6023 Appendix 5f: Sclater (1901). The mammals of South Africa. ........................................... 6025 Appendix 5g: Jameson (1909). LIII. On a collection of mammals from South Africa. 6027 Appendix 5h: Andersen (1912). Catalog of the Chiroptera in the collection of the British Museum. Second Edition. Volume I: Megachiroptera .................................................... 6028 Appendix 5i: Monard (1935). Contribution à la mammalogie d'Angola et prodrome d'une faune d'Angola. ....................................................................................................................... 6044 xvi ISSN 1990-6471 Appendix 5j: Frechkop (1938). Exploration du Parc National Albert. Mission G. F. De Witte (1933 - 1935). Fascicule 10. Mammifères. .............................................................. 6046 Appendix 5k: Dorst (1948). Les chiroptères du genre Triaenops Dobson (Hipposidérines)...................................................................................................................... 6047 Appendix 5l: Malbrant and Maclatchy (1949). Faune de l'équateur Africain français. .. 6048 Appendix 5m: Panouse (1951). Les chauves-souris du Maroc. ...................................... 6053 Appendix 5n: Roberts (1951). The mammals of South Africa ......................................... 6056 Appendix 5o: Malbrant (1952). Faune du Centre African français (Mammifères et Oiseaux). .................................................................................................................................. 6062 Appendix 5p: Sanborn and Hoogstraal (1955). The identification of Egyptian bats. .... 6064 Appendix 5q: Hoogstraal (1962). A brief review of the contemporary land mammals of Egypt (including Sinai). 1. Insectivora and Chiroptera. ................................................... 6066 Appendix 5r: Hill (1963). A revision of the genus Hipposideros. .................................... 6068 Appendix 5s: Hayman et al. (1966). The bats of the Congo and of Rwanda and Burundi .................................................................................................................................................. 6072 Appendix 5t: Hayman and Hill (1971). Order Chiroptera: The mammals of Africa ........ 6079 Appendix 5u: Smithers (1971). The mammals of Botswana. ........................................... 6092 Appendix 5v: Hill (1974). A review of Scotoecus Thomas, 1901 (Chiroptera: Vespertilionidae) ..................................................................................................................... 6095 Appendix 5w: Fenton (1975). Observations on the biology of some Rhodesian bats, including a key to the Chiroptera of Rhodesia. ................................................................... 6096 Appendix 5x: Bergmans (1975). On the differences between sympatric Epomops franqueti (Tomes, 1860) and Epomops buettikoferi (Matschie, 1899), with additional notes on the latter species (Mammalia, Megachiroptera). ............................................................ 6100 Appendix 5y: Hill (1977). A review of the Rhinopomatidae (Mammalia: Chiroptera). ... 6101 Appendix 5z: Jones (1977a). Plecotus rafinesquii ............................................................ 6102 Appendix 5aa: Happold and Happold (1978c). The fruit bats of western Nigeria. Part 3. .................................................................................................................................................. 6103 Appendix 5ab: Rautenbach, I.L. (1975). The mammals of the Transvaal. ...................... 6104 Appendix 5ac: Smithers and Wilson (1979). Check List and Atlas of the Mammals of Zimbabwe Rhodesia ............................................................................................................... 6107 Appendix 5ad: El-Rayad (1980). Systematics of African molossid bats of the subgenus Xiphonycteris of the genus Tadarida (Molossidae Chiroptera). ........................................ 6112 Appendix 5ae: El-Rayah (1981). A New species of bat of the genus Tadarida (Family Molossidae) from West Africa. .............................................................................................. 6113 Appendix 5af: Rautenbach (1982). Mammals of Transvaal. ............................................... 6114 Appendix 5ag: Smithers, R.H.N. (1983). The mammals of the Southern African subregion .................................................................................................................................................. 6117 Appendix 5ah: Wassif et al. (1984). Fauna and Flora of Egypt. 1. On a collection of bats from Egypt....................................................................................................................... 6122 Appendix 5ai: Qumsiyeh (1985). The bats of Egypt.......................................................... 6124 Appendix 5aj: Aulagnier and Thevenot (1986). Catalogue des mammifères sauvages du Maroc ....................................................................................................................................... 6126 Appendix 5ak: Meester et al. (1986). Classification of Southern African mammals ..... 6128 Appendix 5al: Qumsiyeh and Jones (1986). Rhinopoma hardwickii and Rhinopoma muscatellum ............................................................................................................................ 6134 Appendix 5am: Happold (1987). The mammals of Nigeria. .............................................. 6135 Appendix 5an: Boulay and Robbins (1989). Epomophorus gambianus ........................ 6140 Appendix 5ao: Happold and Happold (1989). The bats (Chiroptera) or Malawi, Central Africa: Checklist and keys for identification ........................................................................ 6141 Appendix 5ap: Kowalski and Rzebik-Kowalska (1991). Mammals of Algeria. ............... 6145 Appendix 5aq: Peterson et al. (1995). Faune de Madagascar: Chiroptères ................... 6147 Appendix 5ar: Bergmans (1997). Taxonomy and biogeography of African fruit bats (Mammallia, Megachiroptera).5. The genera Lissonycteris Andersen, 1912, Myonycteris Matschie, 1899 and megaloglossus Pagenstecher, 1.885; General remarks and conclusions; Annex: Key to all species ............................................................................... 6149 Appendix 5as: Gharaibeh (1997). Systematics, distribution, and zoogeography of mammals of Tunisia. .............................................................................................................. 6153 Appendix 5at: Bouchard (1998). Chaerephon pumilus .................................................... 6155 Appendix 5au: Colket and Wilson (1998). Taphozous hildegardeae .............................. 6156 African Chiroptera Report 2020 xvii Appendix 5av: Owen-Ashley and Wilson (1998). Micropteropus pusillus ..................... 6157 Appendix 5aw: Taylor (1998). The smaller mammals of KwaZulu-Natal. ....................... 6158 Appendix 5ax: Dunlop (1999). Mops midas ....................................................................... 6161 Appendix 5ay: Gray et al. (1999). Nycteris thebaica ......................................................... 6162 Appendix 5az: Kwiecinski and Griffiths (1999). Rousettus egyptiacus .......................... 6164 Appendix 5ba: Taylor (1999c). Problems with the identification of southern African Chaerephon (Molossidae ), and the possibility of a cryptic species from South Africa and Swaziland. ................................................................................................................................ 6165 Appendix 5bb: Taylor (2000). Bats of Southern Africa ..................................................... 6166 Appendix 5bc: Russ et al. (2001). Key to the bats of Madagascar .................................. 6172 Appendix 5bd: Fahr et al. (2002). A revision of the Rhinolophus maclaudi species group with the description of a new species from West Africa (Chiroptera: Rhinolophidae). .. 6174 Appendix 5be: Jacobs and Fenton (2002). Mormopterus petrophilus ........................... 6175 Appendix 5bf: Csorba et al. (2003). Horsehoe bats of the World (Chiroptera: Rhinolophidae). ....................................................................................................................... 6176 Appendix 5bg: Decher and Fahr (2005). Hipposideros cyclops ...................................... 6180 Appendix 5bh: Bates et al. (2006). A description of a new species of Pipistrellus (Chiroptera: Vespertilionidae) from Madagascar with a review of related Vespertilioninae from the island. ....................................................................................................................... 6182 Appendix 5bi: Csorba (2008). Taxonomy of the horseshoe bats of the world (Chiroptera: Rhinolophidae). ....................................................................................................................... 6183 Appendix 5bj: Thorn and Kerbis Peterhans (2009). Small mammals of Uganda. .......... 6187 Appendix 5bk: Rambaldini (2010). Glauconycteris variegata .......................................... 6195 Appendix 5bl: Bol et al. (2011). Introduction à l’inventaire des Chauves-souris (Chiroptères) de la ville de Maroua, Extrême-Nord Cameroun. ......................................... 6196 Appendix 5bm: Patterson and Webala (2012). Keys to the bats (Mammalia: Chiroptera) of East Africa. .............................................................................................................................. 6198 Appendix 5bn: Lanza et al. (2015). The bats of Somalia and neighbouring areas. ....... 6208 Appendix 5bo: Van Cakenberghe et al. (2017). The bats of the Congo and of Rwanda and Burundi revisited (Mammalia: Chiroptera). .......................................................................... 6215 Appendix 6: Images of Type Specimens ....................................................................................... 6226 Appendix 7: Summary of Characteristic Information .................................................................. 6262 Appendix 7a: Echolocation call Parameters from Literature ............................................. 6262 Appendix 7b: Methods used in Echolocation Literature .................................................... 6290 Appendix 7c: Karyotype information from Literature ......................................................... 6293 Appendix 8: Abstracts ................................................................................................................ 6297 Appendix 9: Gazetteers .............................................................................................................. 6756 Appendix 9a: Hierarchical Gazetteer .................................................................................... 6756 Appendix 9b: General Alphabetical Gazetteer ..................................................................... 7121 Appendix 9c: Country Alphabetical Gazetteer .................................................................... 7492 Appendix 9d: Quarter Degree Gazetteer .............................................................................. 7844 Appendix 9e: Altitudinal Gazetteer ....................................................................................... 8371 African Chiroptera Report 2020 1 Introduction The purpose of the African Chiroptera Report is to collate published information on, and collate specimen records of, African bats. The advent of the internet provides an opportunity for large amounts of information to be easily and economically updated and accessible, which is particularly important for taxonomic information. The electronic, web-based nature of the information is intended to allow information on African bats to be corrected/updated more frequently than a printed format allows, and to be available to users in an affordable form that can be manipulated to their specific requirements. It is hoped this tool will facilitate research and conservation planning, and possibly stimulate interactions across different areas of research. The report is generated from data collated in the African Chiroptera Database and will hopefully grow and develop with the addition of new and corrected information. The incorporation of information other than taxonomic (see the various section headings in the description of the layout below), is still patchy in its execution across the taxa. Information that may answer specific requirements of a user, i.e. more information about the voucher specimens, or specimen collectors, has been drawn from across the database and is presented in separate appendices. Published identification keys for African bat species, have, where necessary, been updated to include current names, and are presented in appendix 5. In appendix 6 images of type specimens are included as they become available. TAXONOMY In 2006 an African Chiroptera Taxonomic Advisory Committee (ACTAC) was established to guide the development of this resource. This Committee has made valuable contributions regarding the incorporation of suggested changes/corrections, in particular in relation to the taxonomy, into this report. The report is currently most comprehensive in the presentation of information on all known synonyms, for presently accepted taxa of African bat species. Having a high Chiroptera diversity, there are currently 17 families (incl. 4 extinct), 80 genera (24 extinct) and 392 species (53 extinct) recognized as occurring (or having occurred) in Africa. An overview of the number of genera and species per family occurring in Africa is shown in Figure 1. Usually the number of species is larger than the number of genera, but there is one exception: in the Megadermatidae, three genera and two species are mentioned. The reason for this is that only extant taxa are included, and although Megaderma gigas is an extinct species, the genus itself still has representatives in Asia. The graph further shows that the Vespertilionidae contain the largest number of both genera and species, followed by the Molossidae and Pteropodidae in number of species, and by the Pteropodidae and Molossidae in number of genera. The synonym information is still largely based on work by Meester et al. (1986), Koopman (1993a), and Simmons (2005). In general, the taxonomy proposed on the BatNames.org website (Simmons and Cirranello, 2020) is followed for the 2020 issue of the ACR. The report also aims to incorporate information on the whereabouts of type specimens, name combinations, and spelling variations and errors/lapsus calami in names. 2 ISSN 1990-6471 Genera and Species per Family 124 140 120 100 80 60 40 20 0 44 2 1 4 12 4 21 2 3 44 26 1 2 7 1 1 38 15 13 6 1 3 3 16 1 NrGenera NrSpecies Figure 1. Overview of the number of extant genera (yellow) and species (red) per family occurring in Africa (Modified after Srinivasulu et al., 2010: 1008). NEW TAXONOMIC CHANGES SINCE THE 2019 REPORT Definitions of terms for taxonomic changes: The taxonomic changes listed here and explained in the appropriate species accounts are new opinions relating to "recent" systematic works as well as publications dealing with extinct taxa: New taxon – a recently described taxon or a taxon that was for some reason omitted from the previous report. Revised status – re-establishment of the taxon name. Removed – a taxon, which was previously recognized as occurring within Africa, but has been shown to not exist, for various reasons. New Status Epomophorus intermedius (Hayman, 1963) Epomophorus pusillus Peters, 1868 Doryrhina camerunensis (Eisentraut, 1956) Rhinolophus rhodesiae Roberts, 1946 1 Status change 2019 Status PTEROPODIDAE Gray, 1821 Micropteropus intermedius Revised status Hayman, 1963 Micropteropus pusillus Revised status (Peters, 1868) HIPPOSIDERIDAE Lydekker Hipposideros camerunensis Revised status Eisentraut, 1956 RHINOLOPHIDAE Gray, 1825 Revised status Justification1 Previously recognized as a synonym of Rhinolophus swinnyi Gough, 1908 If no explicit justification is given, the change is based on Simmons and Cirranello (2020) [Simmons, N.B. and A.L. Cirranello. 2020. Bat Species of the World: A taxonomic and geographic database. Accessed on 09/28/2020. - https://batnames.org/] African Chiroptera Report 2020 New Status Coleura gallarum Thomas, 1915 Mops (Chaerephon) aloysiisabaudiae (Festa, 1907) Mops (Chaerephon) ansorgei (Thomas, 1913) Mops (Chaerephon) atsinanana (Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, 2010) Mops (Chaerephon) bemmeleni (Jentink, 1879) Mops (Chaerephon) bivittatus (Heuglin, 1861) Mops (Chaerephon) chapini J.A. Allen, 1917 Mops (Chaerephon) gallagheri (Harrison, 1975) Mops (Chaerephon) jobimena (Goodman and Cardiff, 2004) Mops (Chaerephon) leucogaster (A. Grandidier, 1869) Mops (Chaerephon) major (Trouessart, 1897) Mops (Chaerephon) nigeriae (Thomas, 1913) Mops (Chaerephon) nigeriae nigeriae (Thomas, 1913) Mops (Chaerephon) nigeriae spillmanni (Monard, 1933) Mops (Chaerephon) pumilus (Cretzschmar, 1826) Mops (Chaerephon) pusillus (Miller, 1902) Mops (Chaerephon) russatus J.A. Allen, 1917 Mops (Chaerephon) tomensis (Juste and Ibáñez, 1993) Miniopterus arenarius Heller, 1912 Miniopterus nimbae Monadjem, Shapiro, Richards, Karabulut, Crawley, Nielsen, Hansen, Bohmann and Mourier, 2020 Status change 2019 Status EMBALLONURIDAE Gervais, 1855 Revised status 3 Justification1 Previously recognized as a synonym of Coleura afra (Peters, 1852) MOLOSSIDAE Gervais, 1856 Chaerephon Revised status aloysiisabaudiae (Festa, 1907) Chaerephon ansorgei Revised status (Thomas, 1913) Chaerephon atsinanana Revised status Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, 2010 Revised status Revised status Revised status Revised status Revised status Chaerephon bemmeleni (Jentink, 1879) Chaerephon bivittatus (Heuglin, 1861) Chaerephon chapini J.A. Allen, 1917 Chaerephon gallagheri (Harrison, 1975) Chaerephon jobimena Goodman and Cardiff, 2004 Revised status Chaerephon leucogaster (A. Grandidier, 1869) Revised status Chaerephon major (Trouessart, 1897) Chaerephon nigeriae Thomas, 1913 Chaerephon nigeriae nigeriae Thomas, 1913 Revised status Revised status Revised status Chaerephon nigeriae spillmanni (Monard, 1933) Revised status Chaerephon pumilus (Cretzschmar, 1826) Revised status Chaerephon pusillus (Miller, 1902) Chaerephon russatus J.A. Allen, 1917 Chaerephon tomensis (Juste and Ibáñez, 1993) Revised status Revised status MINIOPTERIDAE Dobson, 1875 Revised status New species Previously included in either M. natalensis or M. schreibersii, but see Monadjem et al. (2020: 242). 4 ISSN 1990-6471 New Status Miniopterus villiersi Aellen, 1956 Myotis zenatius Ibáñez, Juste, Salicini, Puechmaille and Ruedi, 2019 Scotophilus altilis G.M. Allen, 1914 Status change 2019 Status Revised status Justification1 Previously recognized as a synonym of Miniopterus inflatus or Miniopterus schreibersi, but see Monadjem et al. (2020: 237, 248) VESPERTIILIONIDAE Gray, 1821 New species Revised status Scotophilus nigritellus de Winton, 1899 Revised status Eptesicus (Cnephaeus) bottae (Peters, 1869) Eptesicus (Cnephaeus) hottentotus (A. Smith, 1833) Eptesicus (Cnephaeus) isabellinus (Temminck, 1840) Eptesicus (Cnephaeus) platyops (Thomas, 1901) Eptesicus (Rhinopterus) floweri (de Winton, 1901) Parahypsugo Hutterer, Decher, Monadjem and Astrin, 2019 Parahypsugo bellieri (De Vree, 1972) Revised status Parahypsugo crassulus (Thomas, 1904) Parahypsugo eisentrauti (Hill, 1968) Parahypsugo happoldorum Hutterer, Decher, Monadjem and Astrin, 2019 Parahypsugo macrocephalus Hutterer and Kerbis Peterhans, 2020 Laephotis anchietae (Seabra, 1900) Laephotis brunneus (Thomas, 1880) Laephotis capensis (A. Smith, 1829) Laephotis guineensis (Bocage, 1889) Laephotis aff. guineensis Laephotis helios (Heller, 1912) Laephotis humbloti (A. Milne-Edwards, 1881) Laephotis isabella (Decher, Hutterer and Monadjem, 2016) Laephotis malagasyensis (Peterson, Eger and Mitchell, 1995) Revised status Revised status Revised status Revised status Revised status Traditionally considered a synonym of Scotophilus leucogaster, but see Vallo et al. (2019). Previously recognized as a synonym of Scotophilus viridis Eptesicus bottae (Peters, 1869) Eptesicus hottentotus (A. Smith, 1833) Eptesicus isabellinus (Temminck, 1840) Eptesicus platyops (Thomas, 1901) Eptesicus floweri (de Winton, 1901) New genus Revised status Revised status Previously recognized as a synonym of Hypsugo crassulus (Thomas, 1904) Hypsugo crassulus (Thomas, 1904) Hypsugo eisentrauti (Hill, 1968) New species New species Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Hypsugo anchietae (Seabra, 1900) Neoromicia brunnea (Thomas, 1880) Neoromicia capensis (A. Smith, 1829) Neoromicia guineensis (Bocage, 1889) Neoromicia aff. guineensis Neoromicia helios (Heller, 1912) Neoromicia humbloti (A. Milne-Edwards, 1881) Neoromicia isabella Decher, Hutterer and Monadjem, 2016 Neoromicia malagasyensis (Peterson, Eger and Mitchell, 1995) African Chiroptera Report 2020 New Status Status change Laephotis matroka (Thomas and Schwann, 1905) Revised status Laephotis nanus (Peters, 1852) Laephotis rendalli (Thomas, 1889) Laephotis robertsi (Goodman, Taylor, Ratrimomanarivo and Hoofer, 2012) Laephotis roseveari (Monadjem, Richards, Taylor and Stoffberg, 2013) Laephotis somalicus (Thomas, 1901) Laephotis stanleyi (Goodman, Kearney, Ratsimbazafy and Hassanin, 2017) Laephotis tenuipinnis (Peters, 1872) Laephotis zuluensis (Roberts, 1924) Pipistrellus (Pipistrellus) aero Heller, 1912 Pipistrellus (Afropipistrellus) grandidieri (Dobson, 1876) Pipistrellus (Pipistrellus) hanaki Hulva and Benda, 2004 Pipistrellus (Pipistrellus) hesperidus (Temminck, 1840) Pipistrellus (Pipistrellus) hesperidus broomi Roberts, 1948 Pipistrellus (Pipistrellus) hesperidus hesperidus (Temminck, 1840) Pipistrellus (Pipistrellus) hesperidus subtilis (Sundevall, 1846) Pipistrellus (Pipistrellus) inexspectatus Aellen, 1959 Pipistrellus (Pipistrellus) kuhlii (Kuhl, 1817) Pipistrellus (Pipistrellus) maderensis (Dobson, 1878) Pipistrellus (Pipistrellus) nanulus Thomas, 1904 Pipistrellus (Pipistrellus) permixtus Aellen, 1957 Pipistrellus (Pipistrellus) pipistrellus (Schreber, 1774) Pipistrellus (Pipistrellus) pygmaeus (Leach, 1825) Pipistrellus (Pipistrellus) raceyi Bates, Ratrimomanarivo, Harrison and Goodman, 2006 Pipistrellus (Pipistrellus) rueppellii (Fischer, 1829) Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status 2019 Status Neoromicia matroka (Thomas and Schwann, 1905) Neoromicia nana (Peters, 1852) Neoromicia rendalli (Thomas, 1889) Neoromicia robertsi Goodman, Taylor, Ratrimomanarivo and Hoofer, 2012 Neoromicia roseveari Monadjem, Richards, Taylor and Stoffberg, 2013 Neoromicia somalica (Thomas, 1901) Neoromicia stanleyi Goodman, Kearney, Ratsimbazafy and Hassanin, 2017 Neoromicia tenuipinnis (Peters, 1872) Neoromicia zuluensis (Roberts, 1924) Pipistrellus aero Heller, 1912 Pipistrellus grandidieri (Dobson, 1876) Pipistrellus hanaki Hulva and Benda, 2004 Revised status Pipistrellus hesperidus (Temminck, 1840) Revised status Pipistrellus hesperidus broomi Roberts, 1948 Revised status Pipistrellus hesperidus hesperidus (Temminck, 1840) Pipistrellus hesperidus subtilis (Sundevall, 1846) Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Revised status Pipistrellus inexspectatus Aellen, 1959 Pipistrellus kuhlii (Kuhl, 1817) Pipistrellus maderensis (Dobson, 1878) Pipistrellus nanulus Thomas, 1904 Pipistrellus permixtus Aellen, 1957 Pipistrellus pipistrellus (Schreber, 1774) Pipistrellus pygmaeus (Leach, 1825) Pipistrellus raceyi Bates, Ratrimomanarivo, Harrison and Goodman, 2006 Pipistrellus rueppellii (Fischer, 1829) 5 Justification1 6 ISSN 1990-6471 New Status Status change Pipistrellus (Pipistrellus) rueppellii fuscipes Thomas, 1913 Pipistrellus (Pipistrellus) rueppellii pulcher (Dobson, 1875) Pipistrellus (Pipistrellus) rueppellii rueppellii (J. B. Fischer, 1829) Pipistrellus (Pipistrellus) rueppellii senegalensis Dorst, 1960 Pipistrellus (Pipistrellus) rueppellii vernayi Roberts, 1932 Pipistrellus (Pipistrellus) rusticus (Tomes, 1861) 2019 Status Revised status Pipistrellus rueppellii fuscipes Thomas, 1913 Revised status Pipistrellus rueppellii pulcher (Dobson, 1875) Revised status Pipistrellus rueppellii rueppellii (J. B. Fischer, 1829) Pipistrellus rueppellii senegalensis Dorst, 1960 Revised status Revised status Pipistrellus rueppellii vernayi Roberts, 1932 Revised status Pipistrellus rusticus (Tomes, 1861) Justification1 GEOGRAPHIC SCOPE This report covers the geographic area of the African continent and adjacent regions, including, Madagascar, and other surrounding islands in the Indian and Atlantic oceans (Figure 3). This and previous issues of the report have focused attention mainly on capturing information from the African continent and the associated islands. The taxa from the Arabian Peninsula are included as far as these are shared with the African continent. SPECIES RICHNESS AND ESTIMATES Figure 2 shows the number of families, genera and species occurring in the various African countries. The most diverse country in number of genera and species is The Democratic Republic of the Congo, followed by Kenya, Cameroon and Tanzania (all over 100 species) (see Figure 3 and 4). Species lists per country together with voucher specimens (evidence) can be located in Appendix 2e. Families,136Genera and Species per Country 123 112 140 120 100 80 60 40 20 0 87 111 94 96 90 68 52 61 65 65 75 70 7079 94 797172 60 57 57 48 48 47 45394038 39 35 32 4133 30 27 35 4033 293420 21 36 20 34 18272528 33 33 33 32 32 16 25 30 12 2121 252 2622 9 15 283 26 152 289 14 231 2715 6 14 6 4 1920 228 11 5 2529 272 2122 4 26 3 121 15 2 1720 1 13 11 1917 10 10 9 8 6 72 4 45 6 3 2 7 1 3 1 42 2 1 1 6 9 1 8 9 9 9101 3108 4 9108 6 9 8 9 9112 9 9 9 6 2115 9 6 1 9 1107 1 8 3 2 7 9 9 9105 9 4 2 9 4 9111010110108 8 5101109 9 69 NrFamilies NrGenera NrSpecies Figure 2. Overview of the number of extant families (blue), genera (yellow) and species (red) occurring in the various countries in Africa (Modified after Srinivasulu et al., 2010: 1008). Species richness maps for individual countries showing all known records have been mapped at a quarter degree level (0.25) (Appendix 2h). African Chiroptera Report 2020 7 Examination of the estimated spatial extent of collection coverage (Figure 4) indicates large areas where there is urgent need for surveys. Within southern Africa – southeastern Angola is in desperate need of preliminary survey work, as well as smaller areas in northern Mozambique, Botswana southwestern Zambia and Northern Cape Province of South Africa. In East Africa - Tanzania and Somalia should be prioritized and in Central Africa - The Democratic Republic of the Congo. Figure 3. African bat species richness map, calculated in half degree grid (0.5) size from the specimen data in the African Chiroptera Database 2020. Light green = 1- 8 species; dark green / black = 9 – 17 species; yellow = 18 - 25 species; orange = 26 - 34 species; red= 35 - 60 species. Figure 4. African bat species richness estimator (Choa 2) was used to calculate in quarter degree (0.25) grid size the specimen data in the African Chiroptera Database 2020, using DIVA-GIS 7.4.0.1 (http://www.diva-gis.org) with a nearest neighbour set at 2. Light green = 1 – 10; dark green / black = 11 – 30; yellow = 31 – 65; orange = 66 – 100; red = 101 – 178. 8 ISSN 1990-6471 All light green areas in species richness maps should be viewed with some caution as these only represent 1 - 10 species. These areas most likely indicate bat records collected as part of larger mammal fieldwork or were collected opportunistically. Therefore, areas that are light green require surveys that will focus attention of the bat species found at these sites. For Sub-Saharan Africa, Monadjem and Schoeman (2013: 104) indicate that the highest species richness is recorded in the Albertine Rift, the northern shores of Lake Victoria in East Africa, and the Upper Guinean forest zone of West Africa. Furthermore, eastern South Africa, southern Mozambique, Zimbabwe, Malawi, coastal Kenya and Tanzania, the Ethiopian rift valley, the Congo basin north of the Congo River and the humid Guinean woodlands north of the rainforest in West Africa have a high species richness. The poorest species richness, they found (not unexpectedly) in the arid zones of south-western Southern Africa, the entire Sahel zone of North Africa, and the Horn of Africa. But the latter might still be the result of (large) collections in various museums, which are still unreported (as was already indicated by Schlitter and Delany, 1985: 47). SPECIMENS At the time the 2020 report was generated the African Chiroptera Database contained 133,602 specimen records of which 117,862 specimens had co-ordinates. This indicates that 88 % of the available records could be plotted. The species distribution maps created in this report probably indicate biases in relation to the collections from which data has currently been obtained. In previous issues of the ACR, it was estimated that about 100,000 to 150,000 specimen records were available for African Chiroptera in collections around the world. This is now believed to be a serious underestimate of potentially available specimens (many European, African and Asian museum collections have not been captured as yet). We currently suspect that a more realistic number of records is possibly between 150,000 – 250,000. Using these revised estimates, we assume that between 53 – 89 % of available records have been captured in the ACD, for 2020. LAYOUT OF ACCOUNTS The taxa are presented in alphabetical order, top down (Order, Suborder, Infraorder…etc.). In the synonyms list, an asterisk (*) in front of the year indicates that the synonym represents the original name for the taxon. This also reveals instances where it is not necessarily the oldest name that has taken precedence. The use of colours denotes the following: red - first name used; blue - name currently in use; green - valid synonyms; black - name combinations, spelling variations and errors/lapsus calami. A question mark (?) in the position of the year indicates that the first use of a particular name combination has not yet been determined. Information relating to the following section headings, or citations to relevant publications, may also be presented in an account, if information relating to these sections has been published or communicated (e.g. "pers. comm." or "unpublished report"), and has been entered in the database. Sections marked as "unknown" indicate that literature data explicitly mention that this information is "not known" (contrary to not having found any information on the topic). Taxonomy: Provides a brief history of previous studies in systematics, giving a summary of different conclusions arising from different methodologies and the resulting taxonomic debates. Much of this information has been taken from Simmons (2005). Common names: Common names used, in a variety of different languages, and found in various sources: e.g. Afrikaans (Roberts, 1951),Chinese (Musila et al., 2018a), Czech (Benda, 2010), Dutch (Dietz, 2017), English (Roberts, 1951; Wilson and Cole, 2000), French (http://www.planet-mammiferes.org; Stéphane Aulagnier, pers. comm.), German (Rudi Hasslauer, pers. comm.), Greek, Albanian and Macedonian (Papadatou et al., 2011), Italian: (Lanza et al., 2015), Maltese (http://www.geocities.com/RainForest/3096/bats.html), Spanish (UNEP-WCMC, 2003; Fa, 1991), Arabian and Hebrew (Ferguson, 2002), Tamil (Kamalakannan and Nameer, 2019). Lina (2017) African Chiroptera Report 2020 9 provided an overview of common names of Europan bats in every possible European language. As a large number of these bats also occur in (northern) Africa, these vernacular names are included too. Etymology of name: Information about the origin/meaning of the name. Similar species: Information about other taxa, with which there has been a recorded mistake in identification. Palaeontological / archaeological records: Any records identified from palaeontological or archaeological sites. Conservation status: This section is split into two categories: Global - provides information on past and present Red data assessments of the global populations of a species and; Regional - provides information on past and present Red data assessments of regional or country specific assessments of a species. General distribution: Distribution of the taxa in Africa, and extra-limital to Africa. Much of this information also comes from Simmons (2005). Additional records are indicated by a reference to their source. Biogeography: Information about the evolutionary processes that have driven speciation of the taxa and also its possible historical distributions in association with sister taxa. Geographic variation: Any known variation within a taxon across its distribution. General description of external morphology: Information about the outward appearance that may distinguish a taxon. General description of cranial and dental morphology: Information about the skull, mandible and teeth that may be distinguishing characters of a taxon. Detailed morphology: More detailed description of some structure of a taxon, i.e. for the baculum, tragus, hyoid, or wingshape. Functional morphology: Information on any work that has related morphological structure to function. Sexual dimorphism: Any known variation within a taxon between males and females. Echolocation: Information about the echolocation call of a taxon. Molecular biology: Information in this section is currently divided into three categories for information on DNA, Karyotype, and Protein/allozyme. The following abbreviations are used in the description of the karyotype: 2n diploid number, FN - fundamental number of autosomal chromosome arms, BA - total number of biarmed autosomal chromosomes, X - female sex chromosome, and Y - male sex chromosome. Habitat: Information about the environment in which a taxon has been recorded. Behaviour: Information about the timing and actions of a taxon. Roost: Indicates where a taxon settles during the day and at night. 10 ISSN 1990-6471 Migration: Any records of period movement from one geographic area to another. Feeding/diet: Records of behaviour in relation to feeding, and what is eaten. Predators: Records of other species know to eat the taxon in question. Population: Information in this section is separated into two categories: structure - in relation to any organization between individuals, and density - in relation to the number of individuals. Lifespan: Records relating to how long individuals of a taxon are known to live for. Reproduction and ontogeny: Information relating to the biology associated with producing further members of a species, and the growth/development over the course of a lifespan. Mating: Information relating to the behaviour associated with producing further members of a species. Post-natal development: Information relating to changes that occur after birth. Parasites: Information relating to both ectoparasites (parasites living outside the body), and endoparasites (parasites living within the body). Viruses: Any known viruses carried by a taxon. The taxonomy followed is that by the ICTV (International Committee on Taxonomy of Viruses - http://www.ictvonline.org/index.asp). Utilization/significance: Information about how a taxon may be used or viewed by people. Distribution map: The species distribution maps should still be viewed with some caution, since these are based on voucher specimen records currently included in the African Chiroptera Database. Hence, the actual distribution of the taxon might be more extensive than indicated on the maps, and the voucher specimen classification/identification may not be accurate. On the other hand, it might also be possible that the actual distribution is more restricted than indicated on the maps. This might be due to the inclusion of historical data, which indicate localities where the animals might have occurred a 100 (or less) years ago, when the habitat was favorable for the animals, but which currently is no longer the case (e.g. increased desertification, destruction of rain forest…). Additionally, it needs to be pointed out that due to variety of reasons not all voucher specimen localities have been plotted, i.e. no X, Y co-ordinates could be found, or these co-ordinates fall outside the country where this locality is supposed to be located. Voucher specimen localities that were not plotted are, however, indicated in the Appendices dealing with voucher specimens. Similarly the position of the point within a country does not necessarily confirm its presence at that spot on the map, as data has currently only been screened to fall within its associated country of representation. Appendix 2g contains a larger version of the maps displayed in the taxa accounts. Distribution based on voucher specimens: This is a calculated paragraph, where information is derived from information in the African Chiroptera Database, and provides a list of countries from which there are voucher specimens. A note of caution in viewing this data is that it is based on voucher specimen information currently in the African Chiroptera Database, hence there may be voucher specimen records that have not yet been included in the African Chiroptera Report 2020 11 database, and voucher specimen classification/identification may not be accurate. Again, if you have any knowledge of omissions or errors in relation to voucher specimens, please send this information to the managing editors of the database (ACD@africanbats.org) for inclusion in the next revision. An additional, calculated query indicates through the use of colour in the country name, the period of time when voucher specimens were last collected from a country. Green indicates that voucher specimens have been collected within the past 20 years, blue within the past 20-50 years, orange within the past 50-100 years, and red if the last voucher specimens were collected more than 100 years ago. Country names in black represent countries for which voucher specimens currently have no associated date of collection. The purpose of this ranking is to assist researchers and conservation planners to focus attention on species and areas where further collecting needs to be done to ascertain data on current presence in an area. REFERENCES In the references section of the report, the author names are colour coded to indicate in which part of the report the reference was mentioned: blue indicates that the reference is used in the main report, green that it is found in the appendices, and red is used for references occurring in both the main report and the appendices. References without colour code have not been used in the main report nor in the appendices, but are included to provide an overview of all publications pertaining to African bats. GIS AND ESRI SHAPEFILES GIS shapefile have been release with this 2020 report that contain all the spatial information to date on the bats in Africa. This is the raw information that was used to create the various maps as well as associated information that is used in the calculation of the various appendicies. Download data: www.africanbats.org/Documents/ACR/2020/ACR_2020_data.zip. Shapefile has been created using geographic projection. Caution must be used when interpreting the information as the X and Y co-ordinates have been calculated using various co-ordinate systems (e.g. DDMMSS, DDMM.SS or center of Quarter Degree), this level of precision varies and therefore the points also contain this level of error. FUTURE OF THE REPORT The current managing editors felt this project may have a better chance of being a successful tool for some time into the future, by encouraging contributions of data and information from interested individuals and organizations. It is hoped the African Chiroptera Report will continue to grow and develop with many more contributions of new and corrected information, i.e. verification of specimens, and specimen voucher records to the database from interested individuals and organizations. Not assigning individual names to authorship/editorship of the report has also been an intentional decision, to try and develop "ownership" of the project among all individuals and organizations who contribute to the database. It is envisaged that each release of the report, being built on the foundation of the previous report, will retain the information of whom and in what capacity they contributed to the project in a particular cycle. Future funding is being sought for the continued support and development of the African Chiroptera Project and the development of additional tools and resources. Some of the tools and resources being explored are contracting in of additional technical skills to develop web access, GIS integration for spatial queries and information dissemination. The saying goes 'Garbage in garbage out', the African Chiroptera Project will continue to refine, verify and interrogate information contained in the database. This will be achieved through the development of teams who will verify and check all information contained in the database – verification of identifications, spatial accuracy of collection sites, and the entering in of recent and historical information from published sources. Any mistakes, misrepresentation of data, or omitted publications can be reported to the managing editors of the database (ACD@africanbats.org). This will allow corrections and updates to be made in the next report (2021). 12 ISSN 1990-6471 Taxa profiles ORDER CHIROPTERA Blumenbach, 1779 330BC. DERMAPTERA Aristotle. - Comments: ca. 330 B.C. Non-Linnean but noted here for the sake of interest. 1767. Vespertiliones Pallas, Vespertiliones in genre. pp. 1–35, in Spicilegia Zoologica quibus novae imprimis et obscurae animalium species iconibus, descriptionibus atque commentariis illustrantur. Tomus 1. Fascicle Tertius. G. A. Lange, Berolini, Germany., 3. Comments: Jackson and Groves (2015: 227) consider this as a synonym of Chiroptera. Originally, the name was proposed without rank, and included the genus Vespertilio Linnaeus, 1758. *1779. CHIROPTERA Blumenbach, Handbuch der Naturgeschichte, 58. - Etymology: Neuter plural scientific Latin substantive made up of the Greek substantives, respectively feminine and neuter "χείρ" (kheír) [genitive "χειρόϛ" (kheirós)] meaning "hand" and "πτερόν" (pterón) meaning "wing", or combined: "with the hand serving as a wing" or "with the hand changed into a wing" (see Lanza et al., 2015: 13). - ZooBank: 06148AD8-596C-45A5-875674BCF5758557. (Current Combination) 1806. Chiropteren Duméril, Zoologie Analytique, 5, 11. - Comments: Jackson and Groves (2015: 227) indicate that Duméril introduced this "family" name to include the genera: Galeopithecus Pallas, 1780 [Order Dermoptera]; Pteropus Brisson, 1762; Noctilio Linnaeus, 1766; Vespertilio Linnaeus, 1758; Rhinolophen [ = Rhinolophus Lacépède, 1799]; and Phyllostomen [ = Phyllostomus Lacépède, 1799]. 1815. Chiropteria Rafinesque, Analyse de la Nature., 51, 54. - Comments: Jackson and Groves (2015: 227) indicate that the name originally included the families Galeopia Rafinesque, 1815 [ = Galeopithecidae Gray, 1821] and Vespertilia Rafinesque, 1815 [ = Vespertilionidae J. Gray, 1821], and is an unjustified emendation of Chiroptera Blumenbach, 1779. (Emendation) 1816. Cheiroptères G. Cuvier, xxx, 121. - Comments: Originally proposed as family name and included the genera Vespertilio Linnaeus, 1758; Pteropus Brisson, 1762; Cephalotes É. Geoffroy, 1810 [ = Nyctimene Borkhausen, 1797]; Dysopes Illiger, 1811: 122 [ = Molossus É. Geoffroy, 1805]; Noctilio Linnaeus, 1766; Phyllostoma G. Cuvier, 1800 [ = Phyllostomus Lacépède, 1799]; Les Megadermes [ = Megaderma É. Geoffroy, 1810]; Rhinolophus Lacépède, 1799; Nycteris É. Geoffroy & G. Cuvier, 1795; Thaphozous [ = Taphozous É. Geoffroy, 1818]; and Plecotus É. Geoffroy, 1818 (see Jackson and Groves, 2015: 227). 1821. CHEIROPTERA Gray, 299. - Comments: Proposed as a class, and originally included the orders Fructivorae J. Gray, 1821)[ = Pteropodidae J. Gray, 1821] and Insectivorae J. Gray, 1821 [ = Yangochiroptera Koopman, 1985 part] (see Jackson and Groves, 2015: 227). Synonymized with Chiroptera by McKenna and Bell (1997: 295). 1822. CHEIROPTERA Fleming, The Philosophy of Zoology, xxxii, 175. - Comments: Originally proposed as order and included the genera Galeopithecus Pallas, 1780 [Order Dermoptera (Illiger, 1811)]; Pteropus Brisson, 1762; Cephalotes É. Geoffroy, 1810b [ = Nyctimene Borkhausen, 1797]; Noctilio Linnaeus, 1766; Phyllostoma G. Cuvier, 1800 [ = Phyllostomus Lacépède, 1799]; Molossus É. Geoffroy, 1805; Stenoderma É. Geoffroy, 1818; Rhinopoma É. Geoffroy, 1818; Rhinolophus Lacépède, 1799; Nycteris É. Geoffroy and G. Cuvier, 1795; Megaderma É. Geoffroy, 1810; Thaphozus [sic = Thaphozous] Bowdich, 1821 [ = Taphozous É. Geoffroy, 1818; Vespertilio Linnaeus, 1758; and Plecotus É. Geoffroy, 1818 (see Jackson and Groves, 2015: 227) . Synonymized with Chiroptera by McKenna and Bell (1997: 295). 1866. Nycterides Haeckel, Generelle Morphologie der Organismen., 2: clx. - Comments: Nomen oblitum. Jackson and Groves (2015: 227) indicated that this name was proposed as suborder for the "families" Gymnorrhina [= Gymnorhina Giebel, 1855] [= Yangochiroptera Koopman, 1985 part] (including the genera Vespertilio, Dysopes, Noctilio) and Histiorrhina [= Histiorhina Van der Hoeven, 1855] [= Rhinolophidae J. Gray, 1825] (including the genera Rhinolophus, Megaderma, Phyllostoma). McKenna and Bell (1997: 301) synonymized the name with Chiroptera. 1872. ANIMALIVORA Gill, Smiths. Misc. Coll., 11 (1): 16. - Comments: Proposed as a suborder, which originally included the following families: Desmodidae Bonaparte, 1845, African Chiroptera Report 2020 1875. 1883. 1889. 1985. 1994. 2014. 13 Phyllostomidae J. Gray, 1825, Mormopidae [= Mormoopidae de Saussure, 1860], Rhinolophidae J. Gray, 1825, Megadermidae Gill, 1872 [ = Megadermatidae H. Allen, 1864], Vespertilionidae J. Gray, 1821, Molossidae Gervais, 1855 and Noctilionidae J. Gray, 1821 (see Jackson and Groves, 2015: 228). Generally synonymized with the Microchiroptera. MICROCHIROPTERA Dobson, Ann. Mag. nat. Hist., ser. 4, 16 (95): 346. - Comments: Originally included the families Rhinolophidae J. Gray, 1825, Nycteridae Van der Hoeven, 1855, Vespertilionidae J. Gray, 1821, Emballonuridae Gervais, 1855, and Phyllostomidae J. Gray, 1825 (Jackson and Groves (2015: 228). - Etymology: From neuter plural scientific Latin substantive made up of the Greek adjective "μικρόϛ" (mikrós) meaning "small" and chiroptera, in opposition to Megachiroptera ("big Chiroptera") (see Lanza et al., 2015: 18). Chiropteri de Rochebrune, Act. Soc. Linn. Bordeaux, 37: 87. - Comments: Unjustified emendation of Chiroptera Blumenbach, 1779. Originally included the families Pteropidae [= Pteropodidae J. Gray, 1821], Megadermidae Gill, 1872 [= Megadermatidae H. Allen, 1864], Nycteridae Van der Hoeven, 1855, Rhinolophidae J. Gray, 1825, Phyllorhinidae de Rochebrune, 1883 [= Hipposideridae Flower & Lydekker, 1891], Taphozoidae Jerdon, 1867 [= Emballonuridae Gervais, 1855b], Molossidae Gervais, 1855, and Vespertilionidae J. Gray, 1821 (see Jackson and Groves, 2015: 228). (Emendation) Ptética Ameghino, Actas Acad. Nac. Cien. Rep. Arg. Cór., 6: xxi, 44, 348. - Comments: Proposed as "Grand Seccion" and included the orders Prochiroptera Ameghino, 1889 and Chiroptera Blumenbach (see Jackson and Groves, 2015: 228). McKenna and Bell (1997: 295) synonymized the name with Chiroptera. Pre-occupied by Ptetica Saussure, 1884 (Orthoptera: Acrididae). Yinochiroptera Koopman, Bat Res. News, 25 (3/4): 26 (for 1984). - Comments: Proposed as infraorder and originally included the superfamilies Emballonuroidea Gervais, 1855 and Rhinolophoidea J. Gray, 1825 (Jackson and Groves, 2015: 228). Chiropteriformes Kinman, The Kinman System., 37. - Comments: Proposed as order. McKenna and Bell (1997: 295) synonymized this name with Chiroptera (Jackson and Groves, 2015: 228). Vespertilioniformes Zagorodniuk, Proc. Theriol. School, 12: 8. TAXONOMY: Higher classification of the Chiroptera follows Alonso-Zarazaga (2005), Eick et al. (2005), Hutcheon and Kirsch (2006), and Teeling et al. (2005). The chiropteran monophyly/diphyly debate that raged over the last decade or so has been quelled by recent molecular studies (Stanhope et al., 1992; Van Den Bussche et al., 1998; Allard et al., 1999; Miyamoto et al., 2000; Murphy et al., 2001a, 2001b; Liu et al., 2001; Butler et al., 2010; Amador et al., 2016) that have unequivocally demonstrated bat monophyly, and thereby dispelled the "flying primate" hypothesis (e.g. Pettigrew, 1986). Molecular analyses reject any close relationship among bats, flying lemurs and tree shrews (once grouped in the cohort Archonta), and instead support a sister-taxon relationship between Chiroptera and Eulipotyphla (Murphy et al., 2001a, 2001b), which together are the sister-group of the Fereuungulata clade (Nikaido et al., 2001, Van Valen, 1979, Feijoo and Parada, 2017: 79), in the supercohort Laurasiatheria (also represented by carnivores, pangolins, cetartiodactyls and perissodactyls) (Murphy et al., 2001a, 2001b; Madsen et al., 2001; Eizirik et al., 2001; Scalley et al., 2001; Nikaido et al., 2001; Springer et al., 2001; Nery et al., 2012; Tsagkogeorga et al., 2013; Foley et al., 2016). On of the most recent reviews (Chen et al., 2017: 1998) suggests a close relationship between Chiroptera and Perissodactyla. The relationship between these two is mentioned as "plausible" by Esselstyn et al. (2017: 2308). However, based on neural findings on 38 mammalian species and 82 characters, Dell et al. (2010: 177) still suggest a diphyletic origin of Chiroptera with Megachiropterans being a sistergroup of the primates. Jackson and Groves (2015) present the following higher order subdivision for the Chiroptera: Class Mammalia Linnaeus, 1758 Subclass Theria Parker and Haswell, 1897 Superlegion Trenchnotheria McKenna, 1975 Legion Yangotheria Chow and Rich, 1982 Sublegion Cladotheria McKenna, 1975 Infralegion Zatheria McKenna, 1975 Infraclass Tribosphenida McKenna, 1975 Supercohort Placentalia Bonaparte, 1838 Cohort Laurasiatheria Waddell et al., 1999 Subcohort Scrotifera Waddell et al., 1999 Order Chiroptera Blumenbach, 1779 They also provide an impressive historical overview of the content and synonyms of these divisions. Studies using molecular techniques to investigate higher level relationships within Chiroptera have 14 ISSN 1990-6471 challenged the monophyly of Microchiroptera, as supported by morphological characters associated with laryngeal echolocation (see list of references in Simmons, 1994), and altered the traditional subordinal division of bats as Megachiroptera and Microchiroptera. Recent analyses of nuclear and mtDNA gene sequences (Teeling et al., 2000; Springer et al., 2001; Van Den Bussche et al., 2002; Teeling et al., 2002; 2003; Hoofer et al., 2003; Hutcheon and Kirsch, 2004b; Teeling et al., 2005; Eick et al., 2005; Teeling et al., 2000; Hahn and Nakhleh, 2015; Khwanmunee et al., 2015; Amador et al., 2016; Feijoo and Parada, 2017; Hawkins et al., 2019) now strongly support an alliance of megachiropterans and rhinolophoids (excluding Nycteridae) in one suborder, and all other microbats and nycteriids in the other suborder. Although these independent studies identified similar subordinal groups, Teeling et al. (2002; 2003; 2005), Springer et al. (2001), AmrineMadsen et al. (2003) and Tsagkogeorga et al. (2013: 2263) named the two rearranged Chiropteran suborders as: “Yinpterochiroptera” (including Pteropodidae, Rhinolophidae - includes Hipposideridae, Megadermatidae, Craseonycteridae and Rhinopomatidae) [Springer et al., 2001, also include Nycteridae in the Yinpterochiroptera] and “Yangochiroptera” (including Emballonuridae, Nycteridae, Noctilionidae, Mormoopidae, Phyllostomidae, Natalidae, Furipteridae, Thyropteridae, Myzopodidae, Vespertilionidae, Mysctacinidae, and Molossidae), whereas Hutcheon and Kirsch (2004b; 2006) and Eick et al. (2005: 1876), following the Principal of Typification (International Code of Zoological Nomenclature, 1999), named the two Chiropteran suborders as: "Pteropodiformes" (including the Pteropodidae, Rhinolophidae, Hipposideridae, Megadermatidae, and Rhinopomatidae) and "Vespertilioniformes" (including Emballonuridae, Nycteridae, Noctilionidae, Mormoopidae, Phyllostomidae, Natalidae, Furipteridae, Thyropteridae, Myzopodidae, Vespertilionidae, Mystacinidae, Miniopteridae and Molossidae). These studies showing microbat paraphyly indicate that the complex suite of morphological innovations for nasal emission of laryngeal echolocation pulses found in rhinolophoid bats either evolved independently at least twice, or once with the subsequent loss in Pteropodidae (Simmons and Geisler, 1998; Springer et al., 2001). This also has implications for the evolution of flight, where many features of flight and ventilation are either directly related or functionally correlated with echolocation (Teeling et al., 2000). Hence, morphological characters previously suggested as synapomorphies of microchiropterans (e.g. features of the inner ear and post-cranial skeleton) have probably also evolved independently, together with the evolution of echolocation (Teeling et al., 2000). laryngeal Based on transcriptome RNA-seq data, Lei and Dong (2016: 1) estimate the split between Yinpterochiroptera and Yangochiroptera at around 63 MYA. Teeling et al. (2000: 190) suggest two additional considerations that support the hypothesis that rhinolophids and megabats are more closely related: First, a sister-group relationship between these two rather than between megabats and all other chiropterans, reduces the duration of the implied ghost lineage for megabats from the early Eocene to the late Eocene. Second, both groups have distributions that are restricted to the Old World. The subdivision into suborders "Pteropodiformes" and "Vespertilioniformes" is also supported by cytogentic data (Volleth et al., 2011: 518), as the order of sub-segments in the HSA 4 homologous segment differs between members of both suborders. Contrary to an earlier suggestion that included Furipteridae, Natalidae, Thyropteridae, and Myzopodidae in the Rhinolophoidea clade (Simmons and Geisler, 1998), the recent molecular studies have confirmed an expanded superfamily, Noctilionoidea, which includes the families Noctilionidae, Furipteridae, Phyllostomidae, Mormoopidae, Mystacinidae, and Thyropteridae (Teeling et al., 2002; Hoofer et al., 2003; Van Den Bussche and Hoofer, 2004b; Teeling et al., 2005; Eick et al., 2005). However, placement of some families is still controversial, since the results of Teeling et al. (2005) showed good support for the basal position of Myzopodidae within Noctilionoidae, while those of Hoofer et al. (2003) did not placed Myzopoda within the Noctilionoidae, but rather in a basal position to all other “yangochiropterans”; Emballonuridae (OR EMBALLONUROIDEA?), Vespertilionoidea, Noctilionoidea. While Eick et al. (2005) also indicated Myzopodidae to be basal to the Vespertilionoidea, Noctilionoidea and Emballonuroidea (see figure 1 in Eick et al., 2005: 1874), their finding apparently had no significant nodal support and alternative topologies could not be rejected by the data (Eick et al., 2005). The other superfamilies recognised by the molecular studies are the Rhinolophoidea (including the families Rhinolophidae, Megadermatidae, Craseonycteridae, and Rhinopomatidae), Emballonuroidea (including the families Emballonuridae and Nycteridae), and Vespertilionoidea (including the families African Chiroptera Report 2020 Vespertilionidae, Molossidae, Natalidae, and Miniopteridae) (Eick et al., 2005; Teeling et al., 2005 [although no miniopterids were included in this study], Springer et al., 2001; Hoofer et al., 2003 [although no miniopterids were included in this study]). Springer et al. (2001) also recognised the superfamily Pteropodoidea. Both Eick et al. (2005) and Teeling et al. (2005) also found similar sister taxa groups between the families, with Furipteridae and Noctilionidae (probably what was meant by Noctilionoidae in Eick et al. (2005)), and Nycteridae and Emballonuridae being sister taxa. Many families can be diagnosed by unique morphological apomorphies and/or combinations of plesiomorphic and apomorphic features that strongly suggest monophyly (Miller, 1897; 1907; Hill, 1974a; Van Valen, 1979; Koopman, 1984c; 1994; Griffiths and Smith, 1991; Corbet and Hill, 1992; Griffiths et al., 1992; Griffiths, 1994). The principal exception is Vespertilionidae, a diverse group that comprises between five and eight subfamilies in different classifications: Kerivoulinae, Miniopterinae, Murininae, Tomopeatinae, Vespertilioninae, Nyctophilinae (included within Vespertilioninae by Koopman (1994) and Volleth and Heller (1994)), Antrozoinae (named as a subfamily by Miller (1897) but subsequently included either within Nyctophilinae (e.g. Hill and Smith, 1984) or Vespertilioninae (e.g. Koopman, 1993a; 1994), and Myotinae (usually included as a tribe within Vespertilioninae (Koopman, 1994) but here raised to subfamily rank following the suggestion of Volleth and Heller (1994). The molecular work of Hoofer and Van Den Bussche (2003) suggested four subfamilies in the family Vespertilionidae: Vespertilioninae, Myotinae, Kerivoulinae and Murininae, with the subfamily Miniopterinae being elevated to family level, Miniopteridae. However, the intron results of Eick et al. (2005) were not able to reject or support alternative hypotheses that Miniopteridae was the sister taxa of Molossidae, and not the sister taxa of Vespertilionidae. Based on upper molar morphology, Fracasso et al. (2011: 422) propose the following tree structure: The Icaronycteridae is positioned at the base, followed by three clades: Yinochiroptera, Noctilionoidea, Yangochiroptera. Within the Yinochiroptera, the Emballonuridae form the basal node, followed by Rhinopomatidae, Nycteridae, Megadermatidae, Rhinolophidae, and Hipposideridae. The Yangochiroptera have a basic split-up between the Nataloidea (including the Myzopodidae [at the base], Thyropteridae, and Furipteridae) and the group formed by the Antrozoinae, Mystacinidae, Molossidae (including 15 Molossinae and Tomopeatinae), and Vespertilionidae (as a poorly supported monophyletic group, including Myotinae, Miniopterinae, Kerivoulinae, and Murininae). If the suggestion to follow the format proposed for standardization of nomenclature of higher taxa by Alonso-Zarazaga (2005) is accepted for renaming of the suborders, then the suggested typified name for the order would be, VESPERTILIONIFORMES, from the oldest generic name of Vespertilio Linnaeus 1758. Zagorodniuk (2014: 8) uses an alternative hierarchy, which starts with the Superorder Chiroptera, which contains the order Vespertilioniformes (Chiroptera auct.), which itself contains two suborders: Vespertilionimorpha (Yangochiroptera) and Pteropodimorpha (Yingochiroptera). Known Chiroptera families: † Aegyptonycteridae Simmons, Seiffert and Gunnell, 2016 † Necromantidae Sigé, 2011 † Philisidae Sigé, 1985 † Icaronycteridae Jepsen, 1966 † Archaeonycteridae Revilliod, 1917 † Onychonycteridae Simmons, Seymour, Habersetzer and Gunnell, 2008 † Palaeochiropterygidae Revilliod, 1917 † Hassianycteridae Habersetzer and Storch, 1987 † Mixopterygidae Maitre, 2008 † Tanzanycterididae Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 Pteropodidae Gray, 1821 Hipposideridae Lydekker, 1891 Megadermatidae H. Allen, 1864 Rhinolophidae Gray, 1825 Rhinonycteridae J. E. Gray, 1866 Craseonycteridae Hill, 1974 Rhinopomatidae Dobson, 1872 Emballonuridae Gervais, 1855 Nycteridae Van der Hoeven, 1855 Myzopodidae Thomas, 1904 Mystacinidae Dobson, 1875 Phyllostomidae Gray, 1825 Mormoopidae Saussure, 1860 Noctilionidae Gray, 1821 Furipteridae Gray, 1866 Thyropteridae Miller, 1907 Natalidae Gray, 1866 Molossidae Gervais, 1856 Cistugonidae Lack, Roehrs, Stanley, Ruedi and Van den Bussche, 2010 Miniopteridae Dobson, 1875 Vespertilionidae Gray, 1821 16 ISSN 1990-6471 Worldwide, six of the extant families (Pteropodidae, Rhinolophidae, Hipposideridae, Phyllostomatidae, Molossidae and Vespertilionidae) contain about 75 % of all species (see Shi and Rabosky, 2015: 1529). COMMON NAMES: Arabian: Bouchaara, Boujlida, ‫الوطواط‬, ‫خفاش‬. Castilian (Spain): Murcielagos. Czech: letouni či netopýři, letaunowé, ptákossavci, upírové, letouni lysoblaní. Dutch: Vleermuizen. English: Bats. French: Chauves-souris. German: Fledermäuse. Italian: Chiròtreri, Pipistrèlli. isiXhosa: ilulwane, idludaka. Kabyle (Algeria): Imtchaghyeye. Portuguese: Morcegos. Somalian: Fidmer, Fidmair, Auo-rnèr, Kibille, Kibillei, Kibil, Fid Mer, Sodan silli, Sod silli (see Lanza et al., 2015: 13). Swahili: Popo. Tanoboase (Ghana): ampane (singular), mmpane (plural) (see Ohemeng et al., 2017: 184). Twi (Ghana): apan (singular), mmpan (plural). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The paleontological record of the Chiroptera in Africa, as in the world, is generally incomplete (Butler, 1978). The first bats appeared somewhere in the Mid- to Late Cretaceous (about 88.7 million years ago), and the diversification began at about 74.9 to 63 MYA (Bininda-Emonds et al., 2007: 509; Lovegrove, 2011: 152). However, dos Reis et al. (2012: 3494) situate the origin of the Chiroptera at 59.1 MYA, hence: after the Cretaceous-Paleogene event. Yu et al. (2014: 2204) suggest that the Chiroptera have an Asian origin and that there was rapid adaptive radiation from the emergence of crown bats until the Early Eocene Climatic Optimum. This was followed by a decreased diversification between 49 and 35 MYA, and another period of high diversification between 35 and 25 MYA. Gunnell et al. (2017: 18) indicate that the earliest known bat fossils came from Europe, India, North Africa and Australia (around 53 - 54 MYA), but they all represent archaic bat groups that have no clear phylogenetic connections with modern taxa. Modern families of bats began to appear in the late early to middle Eocene of Europe and North Africa. They also pointed out (p. 1) that only 13 placental mammal genera first occurred in the middle to late Paleogene, of which six represented bats, including some extant genera belonging to both Yangochiroptera and Yinpterochiroptera. The rapid diversification at the Paleocene-Eocene thermal maximum might have been the result of the large increase in insect abundance during that period (Eriksson, 2014: 175). Springer et al. (2011: 2493) mention that the oldest bat fossils are from the Early Eocene of North America, and that further Early Eocene bats are also known from Europe, Africa and Australia. Most researchers seem to agree that bats originated in Laurasia, but some see Gondwana as the cradle for the Chiroptera. Teeling et al. (2005) [in Springer et al. (2011: 2493)] suggested North America or Laurasia as the ancestral area for bats, and Asia, Europe or Laurasia as the ancestral area for both Pteropodiformi (Yinpterochiroptera) and Vespertilioniformi (Yangochiroptera). The oldest fossil bat described from Africa is: Vampyravus (= Provampyrus) orientalis, from the Oligocene of the Fayum (Egypt). However, in their abstract on an upper molar and two fragments of lower molars from El Kohol (Algeria), Ravel et al. (2010: 149) mention these belong to a member of the "Eochiroptera" (sensu Van Valen, 1979), dating back to the Early Eocene. Ravel et al. (2016: 359) indicate that these "Eochiroptera" cover six families (Onychonycteridae Simmons, Seymour, Habersetzer and Gunnell, 2008, Icaronycteridae Jepsen, 1966, Archaeonycteridae Revilliod, 1917, Hassianycteridae Habersetzer and Storch, 1987, Palaeochiropterygidae Revilliod, 1917, and Tanzanycteridae Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003), none of which are related to the current day families. The oldest of the current day families start to appear in the lower Eocene of Africa, whereas on the other continents they only appeared in the middle Eocene (and even in the upper Eocene of Australia). This points to the great importance of Africa during the first phase of the development of modern bats (Ravel et al., 2016: 359). Hildebrand et al. (2010: 268, 281) recorded unknown bat species from the Kumali and Koka sites in southern Ethiopia. CONSERVATION STATUS: Worldwide, Amori et al. (2013: 98) state that 177 out of 1,116 bat species reported by Wilson and Reeder (2005) or 15.39 % have an IUCN status of CR, E, or VU. MAJOR THREATS: Sherwin et al. (2012: 173) [referring to Ciechanowski et al., 2007] indicate that aerialhawking species are probably more sensitive to climatic changes as they are dependent of a food supply that is extremely variable in time as well as in space. This is less the case for species foraging above water surfaces, as they rely on stable and spatially concentrated food resources. African Chiroptera Report 2020 "Water Stress" will become an important climatic change risk factor for species living in arid regions, as they depend on permanent water resources close to roosts (see Adams and Hayes, 2008 [in Sherwin et al. (2012: 173)]). For southern Africa, Pio et al. (2014: 1543) predict that high concentrations of phylogenetic diversity (PD) will contract considerably by 2080, but that climate change will not result in higher nor lower PD losses, on average, compared to random extinction simulations. They also indicated (p. 1546) that the Kalahari will become very barren in terms of future surviving bat PD. For the Mediterranean area, Maiorano et al. (2011: Table S2) predict that none of the presently occurring species will be extinct by the end of the century, but up to five of the Vespertilionid species might encounter a projected range loss of over 80 %. Between two and fifteen species will probably see a loss between 50 and 80 %, which will also be the case for four of the Rhinolophid species. Between four and eleven of the Vespertilionid species will encounter a loss of between 30 and 50 % of their range, which will also be the case for up to two of the Rhinolophid species. All bat families will contain species which will have a range loss of up to 30 %. One of the Hipposiderid species, one or both of the Rhinopomatid species, and one or two of the Vespertilionid species, however, will gain from the changing. O'Shea et al. (2016) reviewed multiple mortality events in bats and identified nineteen events for Africa: Intentional killing (n = 11), Biotic (n = 5), Accidental (n = 1), Wind Turbines (n = 1), Viral or bacterial disease (n = 1). Some bat populations are killed to prevent them spreading viral diseases. However; Plowright et al. (2016: 16) mention that "culling bats may decrease spillover risk if transmission of the virus increases as population size increases (SIR [susceptible-infectious-recovered] dynamics), whereas culling may increase spillover risk if pulses are driven by viral reactivation after stress (SILI [susceptible-infectious-latent-infectious] dynamics)." Another threat is the bushmeat trade (see Dougnon et al., 2012; Friant et al., 2015; Jenkins and Racey, 2008; Kamins et al., 2010, 2011a, 2011b; Kaswera Kyamakya et al., 2019; Mickleburgh et al. (2009; Musaba Akawa et al., 2017; Waylen, 2004), although this might be (temporarily) reduced as a result of zoonotic outbreaks: e.g. Duonamou et al. (2020) found that 78.3 % of their respondents consumed bat meat 17 prior to the 2013-2016 Ebola outbreak in Guinea, but only 31.9 % consumed it during the outbreak. CONSERVATION ACTIONS: Lisón et al. (2019: 11) pointed out that bat conservation in semi-arid and arid landscapes is receiving little research interest, and that studies in these areas are especially under-represented in Africa and Australia (p. 1). BIOGEOGRAPHY: Maas et al. (2015: 1086) indicate that the Afrotropics (roughly Africa south of the Sahara) is certainly not the most specious zoogeographical region (the Neotropics has 100 species more), but it has with 211 out of 237 species the highest percentage of endemics (89 %), versus 75 % in the second region (Neotropics). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Lucati and López-Baucells (2016: 117) provided a summary of chromatic disorders reported in bats and found that this character is underreported for Africa: Albinism (All-white hairs, pale skin and red eyes) is reported from five localities on the African continent and one in Réunion and Piedbaldism (All-white fur/skin patches, eyes always normally coloured) from two on the continent. Other disorders, such as Melanism (Increase in black or brown pigmentation on the whole body), Partial melanism (Increase in black or brown pigmentation on a part of the body), Leucism (Allwhite or whitish hairs, pale skin, eyes always normally coloured.) and Hypomelanism (Beige, golden, yellowish or reddish fur and skin, eyes always normally coloured) weren't reported from Africa at all. Delmore et al. (2018: 127) studied the pigmentation of the tissues surrounding the testes and suggest that scrotal melanin may protect mature sperm from UV damage, and from oxidative damage in species with male sperm storage. DETAILED MORPHOLOGY: Bahlman et al. (2016: 118) provide a schematic overview of the wing joints and musculature of the various bat families. FUNCTIONAL MORPHOLOGY: Adams and Carter (2017) studied the climbing and walking locomotion of a number of Pteropodids and Vespertilionids and found fundamental differences. The Pteropodids had a far greater ability to climb in a head-up posture than the Vespertilionids, whereas the situation was quite the inverse when walking on the ground was compared. The Pteropodids behaved much more 18 ISSN 1990-6471 like other arboreal mammals (e.g. two-toed sloths - Choloepus hoffmanni Peters, 1858). Stanchak et al. (2019: 1591) compared evolutionary models of calcar length and the corresponding disparity-through-time. They found that these indicate that the calcar diversified early in the evolutionary history of Chiroptera, as the variation in calcar length and its relative proportion to tibia and forearm length is of functional relevance to flight-related behaviours, leading to a large phylogenetic diversification. ECHOLOCATION: Fenton (1994: 28) states "Echolocation profoundly affects the biology of bats, their behaviour and ecology and may have been central to their evolution". MOLECULAR BIOLOGY: The following statement has been made by Dell et al. (2013: 65): "So DNA sequence results in bats may have to be accepted with caution in view of the possibility that DNA is perhaps as susceptible to convergent evolution as morphological systems.", indicating that the results of these stateof-the-art techniques need to be interpreted with caution. Dool et al. (2016a: 196) furthermore suggest: "We caution against the indiscriminate use of mtDNA in phylogenetic studies and advocate for pilot studies to select nuclear introns. The selection of marker type and number is a crucial step that is best based on critical examination of preliminary or previously published data. Based on our findings and previous publications, we recommend the following markers to recover phylogenetic relationships between recently diverged taxa (< 20 My) in bats and other mammals: ACOX2, COPS7A, BGN, ROGDI and STAT5A." HABITAT: Reardon and Schoeman (2017: 273) investigated the taxonomic and functional diversity of the assemblage structure of insectivorous bat communities along the Mount Nimba elevational gradient in view of rising global temperatures. They found that there was a taxonomic shift along the gradient, but that functionally, the assemblage remained largely intact. In their study on the bat fauna of the southern African arid region, Monadjem et al. (2017a: 5) found that at a regional scale, landscape features might be more relevant for bats living in arid regions, as opposed to aridity gradients. They also found that open air foragers are restricted in the southern African arid region, although they could not determine the factor behind this. Furthermore, they were unable to determine the ecosystem services provided by bats in arid environment (as opposed to limiting pest insects in agro-ecosystems). DIET: Webala et al. (2019a: 260-261, 266) divide the equatorial African bat fauna into four groups: frugivores, forest-interior insectivores, forest-edge insectivores and open-space insectivores. They suggest that forest-interior insectivores are strongly associated with the interior of the larger, more intact forests, and are more vulnerable to habitat fragmentation and degradation. The three other groups are more strongly associated with edges and smaller, more degraded forest fragments. Every night, insectivorous bats consume approximately 1/4 to 2/3 of their body mass (Lacher et al., 2019: 948). A large proportion of their prey consists of pest insects, e.g. stink bugs. Taylor et al. (2017a: 376) indicated that the percentage of Hemiptera in the diet of bats (at Levubu, RSA) varied between 6-20 % in Nycteris thebaica to 50 % in Pipistrellus hesperidus. They also calculated (p. 378) that the South African madacamia industry (covering about 25,000 ha) avoided a yearly cost between 56.81 and 138.91 USD/hectare as a result of this. Linden et al. (2019: 2069) performed exclusion experiments in macademia orchards in South Africa and found that the benefits of allowing birds and bats access to the trees economically well outweighed the disadvantages caused by crop raiding by monkeys (approximately +5 000 USD per ha/year versus -1 600 USD per ha/year). PREDATORS: Black et al. (1979: 19) reported that the prey of a bat hawk (Macheiramphus alcinus Bonaparte, 1850) observed in Zambia most likely consisted of Cloeotis percivali, Rhinolophus simulator, Hipposideros caffer, and Nycteris thebaica. Mikula et al. (2016: Supplemental data) mention the following birds preying on unspecified bats: Frances's sparrowhawk (Accipiter francesiae Smith, 1834), Rufous-breasted sparrowhawk (Accipiter rufiventris Smith, 1830), Ovambo sparrowhawk (Accipiter ovampensis Gurney, 1875) [on large bat], Tawny eagle (Aquila rapax (Temminck, 1828)), Mountain buzzard (Buteo oreophilus Hartert and Neumann, 1914), African marsh harrier (Circus ranivorus Daudin, 1800), Sooty falcon (Falco concolor Temminck, 1825) [on insectivorous bats], African hobby (Falco cuvierii Smith, 1830), Dickinson's kestrel (Falco dickinsoni Sclater, 1864) [on small insectivorous bat], Taita falcon (Falco fasciinucha Reichenow & Neumann, African Chiroptera Report 2020 1895); Lilac-breasted roller (Coracias caudata Linnaeus, 1766) [on insectivorous bat], Brownhooded kingfisher (Halcyon albiventris (Scopoli, 1786)) [on insectivorous bat], Woodland kingfisher (Halcyon senegalensis (Linnaeus, 1766)), Southern grey shrike (Lanius meridionalis (Temminck, 1820)), Gabar goshawk (Micronisus gabar (Daudin, 1800)), Green wood hoopoe (Phoeniculus purpureus (J.F. Miller, 1784)), Longtailed hawk (Urotriorchis macrourus (Hartlaub, 1855)). Tapanes et al. (2016: 405, 406) indicate that several primates prey on bats: Pan paniscus Schwarz, 1929 (Bonobo); Papio anubis (Lesson, 1827) (olive baboon, Anubis baboon); Perodicticus potto (Statius Müller, 1766) (Potto); Cercopithecus mitis Wolf, 1822 (Blue monkey) and a C. mitis x C. ascanius hybrid. These observations suggest an alternative pathway for bat-Cercopithecus disease transmission that has implications for zoonotic disease transmission to humans. Of the 13 bat predations they observed, three were on Pteropodidae and one on Molossidae. In four addional events, the bats were small (13 - 18 cm body length) and one large (ca. 50 cm wingspan). ACTIVITY AND BEHAVIOUR: Schoeman (2015: 122, 125) classified South African insectivorous bats in three functional urban groups (based on wing and echolocation characteristics and foraging and/or roosting characteristics): 1) urban avoiders - slow-flying bats that typically hunt prey in forests or dense vegetation and have specific roosting requirements such as obligate cave roosters (e.g. Nycteris thebaica, Rhinolophus simulator); 2) urban adapters - bats that forage mainly along forest edges and adjacent open areas, and readily utilize favourable conditions provided by humans, for example, forage at wastewater treatment plants and/or roost in man-made structures or plants in gardens (e.g. Miniopterus natalensis, Neoromicia capensis, Neoromicia nana, Pipistrellus hesperidus/Hypsugo anchietai, Taphozous mauritianus). 3) urban exploiters - fast-flying, open-air bats often recorded in urbanized landscapes that depend to a certain degree on urban resources such as houses that provide suitable roosting sites and can sustain large bat colonies (e.g. Chaerephon pumilus, Mops condylurus, Otomops martiensseni, Scotophilus dinganii, Tadarida aegyptiaca). 19 PARASITES: BACTERIA: Lutz et al. (2019: 3) investigated the microbial richness of bat's skin, oral cavity and gut and found that the skin contained a significantly higher richness than the two other areas. They also found that the geographical location was more important than the taxonomical relationship to explain the microbial variation. In decreasing order, they found the following groups of bacteria in the gut: Proteobacteria, (Enterobacteriaceae) and Firmicutes (Clostridiaceae and Streptococcaceae). In the oral cavity, they found Proteobacteria (Pasteurellaceae and Neisseriaceae) and Firmicutes (Streptococcaceae and Gemellaceae). No specific bacterial phylum was found overabundant in the skin swaps, but Proteobacteria (Enterobacteriaceae), Actinobacteria (Mycobacteriaceae, Pseudonocardiaceae, Nocardiaceae), Bacteroidetes (Moraxellaceae) and Euryarchaeota (Halobacteriaceae) were most present. HEMOSPORIDIA: Perkins and Schaer (2016: 776) provide an overview of the malaria parasites of bats, which include nine genera, of which the most important ones are: Hepatocystis, Polychromophilus, Nycteria, and Plasmodium (the others are: Bioccala*, Biguetiella*, Sprattiella*, Johnsprentia*, and Dionisia - *: not in Africa). Killick-Kendrick (1973: 647) mentions Plasmodium (Vinckea) voltaicum Van Der Kaay, 1964 from a non-specified bat. Makanga et al. (2017: Suppl.) refers to an Anopheles marshalli infected with Polychromophilus melanipherus, which it could only have received from sucking blood of a bat. Rudoler et al. (2018: 392) found four species of Plasmodium in pools of blood from various Ugandan bats: P. berghei Vincke and Lips, 1948 , P. falciparum Welch, 1897, P. yoelii Landau, Michel and Adam, 1868 and P. chabaudi Landau, 1965. TREMATODA Richard (1966: 416) described Acanthatrium (Acaisthatrium) houini from an unidentified bat from Ranomafana (Madagascar). HEMIPTERA Cimicidae: Loxaspis miranda Rothschild 1912 from Gogrial, Sudan, but no host given (Haeselbarth et al., 1966: 10). Cacodmus ignotus Rothschild 1912 described from a single female without locality data, and a subsequent collection is reported from Uganda (Haeselbarth et al., 1966: 10). In the British Museum there are also specimens of Cacodmus sparsilis Rothschild 1914 20 ISSN 1990-6471 from Pietermaritzbrug, South Africa, without host records (Haeselbarth et al., 1966: 10). Aphrania barys Jordan and Rothschild 1912 described from Maseru, Lesotho, host is not known (Haeselbarth et al., 1966: 11). Aphrania recta Ferris and Usinger 1957 described from Mutir, Uganda, and another record from Mabal, Congo, but hosts unknown (Haeselbarth et al., 1966: 11-12). Leptocimex boueti (Brumpt 1910) has been recorded from a few places in Guinea, the northern Gold Coast, Upper Volta, Dahomey and the former French Sudan, where it was found biting humans, the hosts are bats but no definite species (Haeselbarth et al., 1966: 13). Stricticimex antennatus Ferris and Usinger 1957, found in the Cape Province, South Africa in caves at Bredasdorp and Three Sister’s Rocks, the host-bat not recorded (Haeselbarth et al., 1966: 14). Strictimex brevispinosus Usinger 1959, found in a cave near Lake Victoria, Urundi associated with unidentified bat (Haeselbarth et al., 1966: 14). Strictimex intermedius Ferris and Usinger 1959, was recorded from a cave near Mombasa, Kenya, the host not recorded (Haeselbarth et al., 1966: 14). Stricticimex transverses Ferris and Usinger 1957 was originally described from an unknown bat from Bloemfontein, South Africa (Haeselbarth et al., 1966: 14). Polyctenidae: Eoctenes intermedius (Speiser 1904) found on an unidentified bat in the Sudan (Haeselbarth et al., 1966: 16). Androctenes horvathi Jordan 1912 described form an unidentified bat in Somalia (Haeselbarth et al., 1966: 17). DIPTERA Streblidae: Raymondia huberi Frauenfeld 1856 from unidentified bat in the Rukwa Valley, Tanzania (Haeselbarth et al., 1966: 102). Raymondia setosa Jobling 1930 collected from a “broad-eared bat” at Aden (Haeselbarth et al., 1966: 104). Raymondiodes leleupi Jobling 1954 described form a poorly identified bat (Hipposideros caffer or Miniopterus sp.) from Thysville, Congo (Haeselbarth et al., 1966: 104). Shapiro et al. (2016: 255) indicates that Kessel (1925: 24) mentions Raymondia lobulata Speiser, 1900 "off Megaderma lyra from Ceylon, from bats in Madras and British Somaliland". Nycteribiidae: Nycteribia hoogstraali Theodor 1957 from the Congo and Tanzania host unidentified (Haeselbarth et al., 1966: 109). Basilia meridionalis Theodor 1956 host and exact locality not recorded from South Africa (Haeselbarth et al., 1966: 111). Basilia bouvieri (Falcoz, 1924) originally described without exact locality (West Africa) or host (Haeselbarth et al., 1966: 111). Basilia glabra Theodor 1957 from Aruwimi province in the Congo, but no host is given (Haeselbarth et al., 1966: 112). Dipseliopoda arcuata Theodor 1955, collected from an unknown bat at Addis Ababa, Ethiopia (Haeselbarth et al., 1966: 115). When studying the Nycteribiid abundance on fruit bats and insectivorous bats, Luguterah and Lawer (2015: 4) found that fruit bats were much more infested than insectivorous bats. This could be the result of fruit bats being poorer in nitrogen, which leads to a lack of protein, which in turn inhibits the proper functioning of the immune system. Another explanation might be that insectivorous bats capture their prey on the wing and as such minimize contact with contaminated surfaces. Szentivanyi et al. (2017: 182) found that bat fly species showed a higher diversity towards the equator, and also that bat species assigned to the IUCN LC category hosted a larger number of bat fly specimens than bats assigned to the VU category. SIPHONAPTERA Pulicidae: Echidnophaga aethiops Jordan and Rothschild 1906 host recorded as unidentified Chiroptera and locality not mentioned (Haeselbarth et al., 1966: 135). Ischnopsyllidae: Dampfia grahami equatoris Smit 1954 found on a ‘small insectivorous bat’ in the Kivu district of the Congo (Haeselbarth et al., 1966: 187). ARACHNIDA Argasidae: Morel and Mouchet (1965: 492) report the presence of Carios vespertilionis Latreille, 1796 and Carios boueti Roubaud and ColasBelcour 1933 on non-specified bats in Cameroun. ACARI Trombiculidae: Stekolnikov (2018a: 33) reported Schoengastiella vattierae (Taufflieb, 1964) from unidentified bats from the Central African Republic. He also indicated (p. 116) that Trisetica aethiopica (Hirst, 1926) was described from a non specified bat, and that he found (p. 157) Microtrombicula machadoi Taufflieb, 1965. VIRUSES: A study by Afelt et al. (2018: 115) showed that Africa is the third most intensively studied continent as far as virus research is concerned (after Asia and North America). They also reported (p. 119) that worldwide, 33 % of virus related publications deal with Rhabdoviridae, 31 % with Coronaviridae, 10 % with Paramyxoviridae, 6 % with Astroviridae, 3 % with Reoviridae and Adenoviridae, 2 % with Circoviridae, Herpesviridae and Flaviviridae, and 1 % with Parvoviridae, African Chiroptera Report 2020 Picornaviridae, Polyomaviridae, Papilliomaviridae and Filoviridae. Luis et al. (2013, 2015) and O'Shea et al. (2014: 741) indicate that bats are more likely to be infected with more zoonotic viruses per host species than were rodents, and therefore adding weight to the suggestion that bats might in some way be unique as sources of emerging zoonoses. Luis et al. (2015) found that gregarious bats are more likely to share more viruses and regionally migrating bats are important for spreading viruses through the "viral sharing network". O'Shea et al. (2014: 742) also indicate that body temperatures of flying bats are elevated (in the range of typical fever temperatures), and they suggest that these daily high temperatures might arm bats against some pathogens during the early stages of infection. This hypothesis, however, was rejected by Schountz et al. (2017: 2) as fever response to a virus infection is much more complex than simply raising the body temperature. These authors also point to the "allways on" IFN (interferon) system and the immunoglobulin repertoires of bats. Reusken and Heyman (2013: 99) [referring to Weiss et al. (2012b)] indicate that bats might be the primordial reservoir for the majority of viruses pathogenic to humans. Brierley et al. (2016: "5") report that - based on viral richness alone - the risk of bat-human virus sharing showed prominent hotspots in Central/South America, sub-Saharan Africa, and parts of Southeast Asia. Yinda et al. (2019) analysed the gut virome of Cameroonians and found a genetic relatedness between orthoreovirus, picobirnavirus and smacovirus found in humans and those of bats and/or other animals. The accounts below represent only viruses where the host (e.g. bat) is unknown. For further information see taxa accounts where the virus is associated with a particular taxon. Also see Chen et al. (2014: 1) for an online database on batassociated viruses. Furthermore, Moratelli and Calisher (2015: suppl. Table II) provide an overview of bat host genera and their associated virus genera, and Young and Olival (2016: suppl. Table S2) provide details on species level for both bat hosts and viruses. See also http://www.mgc.ac.cn/DBatVir/ for more details on bats and viruses (Nieto-Rabiela et al., 2019: 657). Astroviridae From a study on 962 bats (Hipposideros cf. ruber, Hipposideros gigas, Coleura afra, Miniopterus inflatus, and Rousettus aegyptiacus) from Gabon, Rougeron et al. (2016: 386) found that bat astroviruses form a group that is genetically distinct from astroviruses infecting other mammals 21 (including humans). These viruses also showed an important genetic diversity and low host restriction in bat species. Bunyaviridae Hantavirus Gu et al. (2014: 1904) argue that primordial hantaviruses may have emerged in an early common ancestor for all bats as bat species hosting these viruses belong to both currently accepted suborders of the Chiroptera (Pteropodiformi [Hipposideridae + Rhinolophidae] and Vespertilioniformi [Nycteridae + Vespertilionidae]). This led Yanagihara et al. (2014: 9) to suggest that primordial hantaviruses may have infected an early common ancestor of bats. Yanagihara et al. (2014: 9) also indicate that the five known bat hantaviruses are among the most genetically diverse described to date. Coronaviridae - Coronaviruses Severe Acute Respiratory Syndrome-associated coronavirus (SARS-CoV or hu-SARS-CoV) Balboni et al. (2012: 8) indicate that SARS-like CoVs have possibly originated in Africa as the viruses themselves has been found in several species of the genera Hipposideros and Chaerephon, and antibodies against SARS-CoV antigens have been found in Hypsignathus, Lyssonycteris, Miniopterus, Mops, Myonycteris, Rhinolophus and Rousettus species. Specifically for Coronaviruses worldwide, Anthony et al. (2017b: 7) found that a large number of bat host species were not infected by these viruses, but they also remarked that none of these species were sampled extensively. They found that all species with sample sizes >110 individuals were positive for one or more CoVs, suggesting that CoVs may be detected in some of the negative species if sampling effort were increased. They also calculated (p. 9) that with 154 host individuals, an average of one CoV would be detected, and with 397 individuals, up to five CoVs would be detected. As they did not find any bat species with more than five viral sequence clusters, sampling 397 bats should capture all of the CoVs in a specific bat species. Filoviridae - Filo viruses Han et al. (2016) reported on traits that discriminate host and non-host (=filovirus-positive) bat species, and they provide a list of bat species most likely to be filovirus-positive on the basis of intrinsic trait similarity with known filovirus-positive bats (See also Sylla et al., 2015). In their study, Schuh et al. (2017c) point to the accumulating number of links between filovirus 22 ISSN 1990-6471 hemorrhagic fever (FHF) index cases and prior exposure to environments inhabited by bats. Becker et al. (2019: 4) point out that strong temporal dynamics exist in both Marburg and Ebola virus outbreaks, which may be linked to bat reproduction or food availability. Marburgvirus Peterson and Holder, 2012: 1831) analyzed fifteen Marburg virus sequences from bats, and found no indication from the topology of the tree or the branch lengths that the bat-derived filoviruses represent a population distinct from viruses collected from human outbreaks. They suggest that bat sequences do not form a monophyletic group, nor are they associated with long or “deep” branches in the tree. This situation could indicate that the bats are the reservoir population, or that high gene flow exists between virus populations in bats and those in some other taxon (see Towner et al., 2009). Given the short sequence lengths obtained from bat filovirus (about 302 nucleotides of the VP35 gene for most samples), it is also possible that Peterson and Holder, 2012: 1831) lacked the power to detect subtle evidence for population structuring. Thus, firm conclusions about gene flow between the "bat" filovirus populations and the true source of human infections will require more data. Ebola virus Most papers related to Ebola (e.g. Alexander et al., 2015) do not provide any details on the bat species involved ("fruit bats", "possibly fruit bats") or refer to the "usual suspects" (Myonycteris torquata, Hypsignathus monstrosus, Eidolon helvum". Murray (2015: 802) indicates that the transmission of Ebola virus from bats to humans must be uncommon because the potential reservoir of Ebola virus is huge. Buceta and Johnson (2017) provide a model of the Ebola zoonotic dynamics suggesting a bidirectional coupling between the available resources and the dynamics of the bat population, and indicating that environmental pressure is the main driving force for bats' migration. Based on the fact that the human seropositive zone is wider than the distribution area of M. torquata, H. monstrosus and E. franqueti, Formella and Gatherer (2016: 3125) suggest that E. helvum, which has a wider distribution, may be involved. They also point out that some (early) Ebola outbreaks, which lie beyond the distribution range of the fruit bats, may have been overlooked. An extensive survey of Ebola viruses in 4,022 bats from Guinea, Cameroon and the DRC by De Nys et al. (2018) showed antibodies in Mops sp., Eidolon helvum, Epomophorus sp., Hypsignathus monstrosus, Lissonycteris angolensis, Micropteropus pusillus and Rousettus aegyptiacus. No antibodies were found in Hipposideros sp., Miniopterus sp., Chaerephon sp., Nycteris sp., Rhinolophus sp., Glauconycteris sp., Kerivoula sp., Myotis bocagii, Neoromicia sp., Scotophilus sp., Epomops sp., Megaloglossus woermanni, Myonycteris torquata or Scotonycteris zenkeri. A phylogenetic analysis, combined with phylogeographical data led Hassanin et al. (2016: 518) to conclude that only three bat species are able to disperse directly ZEBOV (Zaire Ebola) from the Congo Basin to Upper Guinea: E. helvum, H. monstrosus, and R. aegyptiacus. de La Vega et al. (2015: 6) [referring to Biek et al. (2006)] indicate that the L gene of bat-derived EBOV showed that it could have recently experienced a genetic bottleneck-type event. Biek et al. (2006) proposed a number of scenario's, of which the first suggests that the bat population (around 1999 when the most recent common ancestor to all currently known Ebola viruses existed) decreased significantly, causing a reduction in the viral population size. A second scenario suggested that infected bats could have introduced EBOV in the region near Congo and Gabon circa 1999, and a final scenario is that the the fruit bat species identified by Leroy et al. (2005) is not the primary reservoir, and that EBOV was possibly introduced by another (bat) species. Analysing the outbreak data and the distribution area of the various bat species, made Pigott et al. (2016: 2) select four species of bats as explaining more of the variation than the rest: Hypsignathus monstrosus, Epomops franqueti (from both of which Ebolavirus RNA has been isolated), Otomops martiensseni and Epomophorus labiatus (both identified through trait-based machine learning approaches as being similar to species already reporting filoviral infection, as suggested by Han et al. (2016)). However, Leendertz (2016: 1) finds it questionable whether these viruses are circulating regularly in bat populations as both fruit and insectivorous bats have a long lifespan and can fly over large distances, which means that seropositive specimens could be sampled for a long period of time and at large distances from the place where they could have been exposed to the Ebolavirus. Furthermore, the various types of Ebola seem to be linked with different river basins, which do not form barriers for fruit bats. She furthermore suggests to look at insects (e.g. mayflies) as possible vectors. Additionally, Spengler et al. (2016: 958) point out that EBOV has yet to be isolated in nature from any bat species (or non- African Chiroptera Report 2020 human primates). Furthermore, EFSA AHAW Panel (EFSA Panel on Animal Health and W (2017: 6) remark that natural shedding of the virus has not been reported and that the suspected links between bats and human outbreaks have not been confirmed and further research is necessary to confirm or identify reservoir species. A similar conclusion is drawn by Holmes et al. (2016: 198), who indicate that it is likely that other host species exist, which may have a major bearing on epidemiological dynamics. Schmidt et al. (2019: 1) indicated that provisional results suggest that some insectivorous bat genera, Old World monkeys and forest antelopes should receive priority in Ebolavirus survey efforts. The extensive survey by Ayouba et al. (2019), however, supports the reservoir role of bats (see also Hayman (2019) for a further discussion). Hranac et al. (2019: "1") indicate that no full Ebolavirus RNA has been isolated in bats yet, but RNA viral fragments have been found in fruit bats (Epomops franqueti, Hypsignathus monstrosus and Myonycteris torquata) and in Miniopterus inflatus. They (p. "8") were able to find some statistical support for a correlation between the time of births in fruit bats and the outbreaks of Ebola Virus Disease. Gale (2017: 1678) argues that although previous studies failed to detect the presence of EBOV in arthropods, these studies might not have included the correct arthropod species: "This is because filoviruses are not ‘arboviruses’ in the sense that arboviruses are transmitted in a defined cycle which alternates between the vertebrate host and the arthropod vector. Instead, filovirus transmission pathways appear to be more random in terms of host species. Indeed, if these arthropod species were the reservoir for EBOV, then EBOV would be an arbovirus with a vertebrate host defined by the host preference of the arthropod." If an arthropod is the intermediate host, then an insectivorous bat would rather be the host than any of the Pteropodids. Gale (2017: 1678) also indicates that the higher body temperature of the bat (40°C in Mops condylurus, compared to that of an arthropod, which is the ambient temperature) would activate the virus reproduction in the bat, and hence, its increased probablity to infect humans. However, the bats will still need to ingest a large quantity of the virus to start off with, but this could happen during an ephemeral mass emergence of mayflies. Li et al. (2016: 127) also pointed out that as large quantities of bat meat are consumed every year, the virus must be very rare in its reservoir, otherwise more frequent outbreaks of the virus would be observed, which led them to the 23 suggestion that strict restrictions on bushmeat are not recommended for local citizens. Leendertz et al. (2015: 22) suggest that bats are intermediate hosts, which are occasionally exposed via another intermediate host or unknown reservoir and that viral emergence might be more related to environmental factors and other hosts than bats themselves. See also Caron et al. (2018: 5) who discuss a number of hypothetical transmission pathways between a maintenance host of EBOV and bats: Air-borne, vector-borne, food-borne (insects or fruits), water-borne, direct and environmental. Dick and Dittmar (2014: 150) indicate that nycteribiid and streblid bat flies are known to infest the above mentioned fruit bats. These flies readily move from bat to bat, feeding on multiple individuals during the course of days and weeks. Possibly, the flies can uptake viral pathogens, which might remain viable within bat flies. If this is the case, host-specific flies can transfer such viruses among host bats within a population. Babayan et al. (2018: 578) worked out a model that analyses virus genomes and links these to probable reservoir hosts. For the four Ebolavirues, the model showed greater support for Pteropodiformi as reservoir than Vespertilioniformi, but Bundibugyo and Tai Forest ebolaviruses had equal or stronger support for primate reservoirs. Analysing the literature data, Ponce et al. (2019: 3133) pointed out that for only three of the 23 outbreaks (13.0 %) the source animal was confirmed in a laboratory. Of the suspected source animals, 10 were primates and six bats (with three fruit bats and three insectivorous bats). On 24 January 2019, a series of newspaper articles was published, where several scientists from EcoHealth Alliance were quoted (Simon J. Anthony, Jonathan Epstein) stating that in a mouth swab one Liberian Miniopterus inflatus (out of 150 tested) RNA material of the Zaire Ebolavirus was found. Epstein, however, pointed out that if the bat was a natural host for the virus, it would have been found in more than one bat. He also pointed out that the bat could have become infected by another bat species living in the same habitat. Fabian Leendertz (veterinary epidemiologist at the Robert Koch Institute - not involved in the study) indicated that the virus itself wasn't isolated but only about one-fifth of its genome (Grady, 2019; Kupferschmidt, 2019; Su, 2019). Schuh et al. (2019: 2) also confirmed that no infectious virus was yet isolated from any bat species. This would indicate that the bats had either cleared virus 24 ISSN 1990-6471 infection prior to sampling or are 'dead-end' virus hosts. However, it might also be possible that current filovirus isolation techniques lack the sensitivity to recover infectious virus from specimens with low viral loads. Rhabdoviridae Lyssavirus - Rabies related viruses Munang'andu et al. (2011: 23) summarize rabies status in Zambia for the period 1985-2004, where only one single case is reported from an unidentified bat. See Adedeji et al. (2010) for an overview of the history, and epidemiology of rabies and rabies-like viruses. DUV - In 2007 a bat flew against the face of a Dutch woman in Tsavo West National Park in Kenya. She noticed two superficial bleeding wounds. Twenty-three days after the incidence she presented herself with symptoms in a hospital in the Netherlands and died of rabies. The virus was identified as Duvenhage virus (Van Thiel et al., 2009). UTILISATION: Kamins et al. (2011b: S102) report that in southeastern Ghana nearly one half of the population may eat bats, although bats do not appear in other studies on wildlife meat markets. ANTHROPOPHILOUS: Willett (1988) discusses the use of bat symbols in West African art. ZOOBANK: 06148AD8-596C-45A5-8756-74BCF5758557 DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Egypt, Gabon, Kenya, Liberia, Madagascar, Malawi, Mauritius, Senegal, Somalia, South Africa, Tanzania, Togo, Uganda, Zambia, Zimbabwe. †Family AEGYPTONYCTERIDAE Simmons, Seiffert and Gunnell, 2016 *2016. Aegyptonycteridae Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 4, 8. Publication date: 9 May 2016. - Comments: Type genus Aegyptonycteris Simmons, Seiffert and Gunnell 2016. TAXONOMY: The supra-familial relationships for this family are yet unknown. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: These fossil bats have a late Eocene (Priabonian, ca. 37 Mya) age. †Genus Aegyptonycteris Simmons, Seiffert and Gunnell, 2016 *2016. Aegyptonycteris Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 6, 8. Publication date: 9 May 2016. - Etymology: From Aegyptus, Latinized Greek for "Egypt," and nycteris, Greek for "bat." The genus name refers to the country in which this new taxon was discovered (Simmons et al., 2016: 8). †Aegyptonycteris knightae Simmons, Seiffert and Gunnell, 2016 *2016. Aegyptonycteris knightae Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 6 8, figs 3 - 5. Publication date: 9 May 2016. Type locality: Egypt: Western Desert: Fayum Depression, Birket Qarun Formation: Fayum Quarry BQ-2: 23 m level [Goto Description]. Holotype: CGM 83740: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Fragment of right maxilla with M2 and M3 (see: Simmons et al., 2016: 8). - Etymology: Named in honour of Mary Knight, Managing Editor of the American Museum of Natural History Scientific Publications, in recognition of the enormous contributions she has made to dissemination of the results of scientific research over the years, as well as her lifelong devotion to the people and culture of Egypt (Simmons et al., 2016: 8). African Chiroptera Report 2020 COMMON NAMES: Dutch: Knight's Egyptische vleermuis. Knight's Egyptian bat English: 25 PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Priabonian (37.97 - 33.23 mya) (see Brown et al., 2019: Suppl.). †Family NECROMANTIDAE Sigé, 2011 *2011. Necromantidae Sigé, Bull. Soc. Hist. Nat. Toulouse, 147: xxx.. - Comments: Type genus †Necromantis Weithofer, 1887. †Genus Necromantis Weithofer, 1887 1888. Necromanter Lydekker, Zool. Rec., London, 71 - 56 [for 1887]. *1888. Necromantis Weithofer, Sber. Akad. Wiss. Wien, math. naturw. Kl., 96 (1) 5: 353 (for 1887). 1903. Necronycteris Palmer, Science, N.S. 17: 873.. Publication date: 29 May 1903. †Necromantis fragmentum (Ravel, 2016) *2016. ?Necromantis fragmentum Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 363, figs 4, 5. Publication date: 30 September 2016. Type locality: Tunisia: Kassérine province: Djebel Chambi National Park: Chambi [35 14 03 N 08 45 29 E, 630 m] [Goto Description]. - Etymology: From the Latin "fragmentum" meaning fragment or debris, referring to the fragmented character of the material attributed to this species (see Ravel et al., 2016: 364). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. 26 ISSN 1990-6471 SUBORDER PTEROPODIFORMI Van Cakenberghe, Kearney and Seamark, 2007 1821. FRUCTIVORAE Gray, London Med. Repos., 15 (1): 299. - Comments: Proposed as order and originally included the families Pteropidae J. Gray, 1821 and Cephalotidae J. Gray, 1821, both currently representing Pteropodidae J. Gray, 1821. Jackson and Groves (2015: 230) included it as synonym of the Pteropodidae. 1855. FRUGIVORA Giebel, Die Säugethiere …, xii, 991. - Comments: Originally included the genera Hypoderma É. Geoffroy, 1828 [= Dobsonia Palmer, 1898]; Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797]; Macroglossus F. Cuvier, 1824 [1821-1825]; and Pteropus Brisson, 1762 (see Jackson and Groves, 2015: 229). McKenna and Bell (1997: 295) synonymized it with Megachiroptera. 1866. Pterocynes Haeckel, Generelle Morphologie der Organismen., 2: clx. - Comments: Nomen oblitum. Proposed as suborder and originally included the "families" Pteropodida (including the genera Pteropus and Macroylossus) and Hypodermida (including the genus Hypoderma) Jackson and Groves, 2015: 229). McKenna and Bell (1997: 295) synonymized it with Megachiroptera. 1875. MEGACHIROPTERA Dobson, Ann. Mag. nat. Hist., ser. 4, 16 (95): 346. - Comments: Originally included the groups Pteropi (containing the genera Pteropus Brisson, 1762; Cynopterus F. Cuvier, 1824 [1821-1825]; Cynonycteris Peters, 1852 [= Rousettus J. Gray, 1821]; Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797]; Epomophorus E. Bennett, 1836; and Cephalotes É. Geoffroy, 1810 [= Nyctimene Borkhausen, 1797]) and Macroglossi (containing the genera Macroglossus F. Cuvier, 1824 [1821-1825]; Eonycteris Dobson, 1873; and Notopteris J. Gray, 1859) (see Jackson and Groves, 2015: 228). Etymology: From the neuter plural scientific Latin substantive made up of the Greek adjective "μέγα" (mega) "big" and chiroptera, i.e. "big Chiroptera", in opposition to Microchiroprera ("small chiroptera") (see Lanza et al., 2015: 18). 2001. YINPTEROCHIROPTERA Springer, Teeling, Madsen, Stanhope and De Jong, Proc. Natl. Acad. Sci. USA, 98 (11): 6423. - Comments: In part. 2004. PTEROPODIFORMES Hutcheon and Kirsch, J. mamm. Evolut., 11 (1): 44. - Comments: Originally included the families Pteropodidae J. Gray, 1821, Hipposideridae Flower & Lydekker, 1891, Rhinolophidae J. Gray, 1825, Megadermatidae H. Allen, 1864, and Craseonycteridae Hill, 1974 (see Jackson and Groves, 2015: 229). *2007. PTEROPODIFORMI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the suborder is based on Pteropus Brisson, 1762 (type of Pteropodidae Gray, 1821). (Current Combination) 2014. Pteropodimorpha Zagorodniuk, Proc. Theriol. School, 12: 8. 2018. Yinpterachiroptera: Skirmuntt and Katzourakis, Virus Research, 270 (197645): 2. Publication date: 1 July 2018. (Lapsus) TAXONOMY: The suborder name follows the typified name used by Hutcheon and Kirsch (2004b, 2006), but with the ending changed to follow the format proposed for standardization of nomenclature of higher taxa by Alonso-Zarazaga (2005). '-formes' indicates ordinal level, while ending '-formi' indicates subordinal level. and found a very high median rate between the superfamilies Pteropoidea and Rhinolophoidea (2.77) and between the families Pteropodidae and Rhinolophidae (2.32). Ao et al. (2007: 257) provide the first cytogenetic signature rearrangement that supports the grouping of Pteropodidae and Rhinolophoidea in a common clade (i.e. Pteropodiformes or Yinpterochiroptera). Currently two Infraorders are recognized within the suborder PTEROPODIFORMI from Africa: PTEROPODIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 and RHINOLOPHIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007. In their phylogenetic study on Bartonella, McKee et al. (2016: 390) estimated the posterior state transition rate for gltA sequences in this bacterium Based on transcriptome RNA-seq data, Lei and Dong (2016: 2) estimated the split between Rhinophoidea and Pteropoidae at about 60 MYA. COMMON NAMES: Czech: kaloňotvaří. African Chiroptera Report 2020 DETAILED MORPHOLOGY: The echolocating bats belonging to the Pteropodiformi can be distinguished from the Vespertilioniformi by the presence of a pectoral ring. This bony structure is formed by the fusion of the top bone of the breast bone (manubrium), the first two ribs and the last cervical and first two thoracic vertebrae (Jacobs (2016: 66-67). MOLECULAR BIOLOGY: Volleth (2010: 308) reports the detection of two cytogenetic characters serving as synapomorphies for Pteropodiformi (i.e. Rhinolophoidea plus Pteropodidae), and indicates 27 that the cytogenetic basis is a small paracentric inversion hardly detectable with banding techniques, but clearly detectable with FISH (fluorescence in-situ hybridization) techniques. VIRUSES: Osborne et al. (2011: 8) indicate that "Yinpterochiroptera" [=Pteropodiformi] is the only suborder in which Betacoronaviruses are found, but they indicate that their conclusion might be biased as they only examined a subset of species of the New World, where no representatives of this suborder are occurring. INFRAORDER PTEROPODIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 *2007. PTEROPODIFORMACEI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the Infraorder is based on Pteropus Brisson, 1762 (type of Pteropodidae Gray, 1821). (Current Combination) TAXONOMY: Currently only a single superfamily is recognised within the infraorder PTEROPODIFORMACEI: PTEROPODOIDEA Gray, 1821. Superfamily PTEROPODOIDEA Gray, 1821 *1821. PTEROPODOIDEA Gray, London Med. Repos., 15 (1): 229. - Comments: The name of the superfamily is based on Pteropus Brisson, 1762 (type of Pteropodidae Gray, 1821), and originally included the genera Pteropus Brisson, 1762; and Rousettus J. Gray, 1821. (Current Combination) 2008. Pteropoidea Serra-Cobo, López-Roig, Bayer, Amengual and Guasch, Ratpenats. Ciència i mite., 31. TAXONOMY: Known families PTEROPODOIDEA: 1821. of the superfamily PTEROPODIDAE Gray, COMMON NAMES: Czech: kaloňovci, kaloni. Family PTEROPODIDAE Gray, 1821 1821. Cephalotidæ Gray, London Med. Repos., 15 (1): 299. - Comments: Type genus: Cephalotes É. Geoffroy, 1810b [= Nyctimene Borkhausen, 1797]. *1821. Pteropidæ Gray, London Med. Repos., 15 (1): 299. Publication date: 1 April 1821. Comments: Jackson and Groves (2015: 230) explains that the stem of the Greek pous (=foot) is pod-, so the stem of the latinised Pteropus is ‘pteropod-’, leading to the corrected name Pteropodidae introduced by Bonaparte (1837-41: 112). 1825. Pteropina Gray, Ann. Philos., 10 (5): 338. Publication date: November 1825. Comments: Proposed as tribe and originally included the genera Pteropus Brisson, 1762; Cynopterus F. Cuvier, 1824 [1821-1825]; Macroglossum [sic = Macroglossus] F. Cuvier, 1824 [1821-1825]; Cephalotis [sic = Cephalotes] É. Geoffroy, 1810b [= Nyctimene Borkhausen, 1797]; and Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797] (see Jackson and Groves, 2015: 230). McKenna and Bell (1997: 295) synonymized it with 28 ISSN 1990-6471 1838. 1842. 1866. 1866. 1866. 1893. Pteropodidae. Jackson and Groves (2015: 236) also mention it in the synonymy of the subfamily Pteropodinae. PTEROPODIDAE Bonaparte, Syn. Vert. Syst., in Nuovi Ann. Sci. Nat., Bologna, 2: 111. Comments: Type genus: Pteropus Brisson, 1762. (Alternate Spelling) Harpyidae C. H. Smith, An introduction to the Mammalia, 15: 115. - Comments: Type genus: Harpyia Illiger, 1811 [=Nyctimene Borkhausen, 1797). Originally included the genera Pteropus Brisson, 1762; Pachysoma I. Geoffroy, 1828 [= Cynopterus F. Cuvier, 1824 [1821-1825]; Macroglossus F. Cuvier, 1824 [1821-1825]; Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797]; and Cephalotes É. Geoffroy, 1810 [= Nyctimene Borkhausen, 1797] (see Jackson and Groves, 2015: 230). Simmons (2005: 313) synonymized this name with Pteropodidae. Hypodermida Haeckel, Generelle Morphologie der Organismen., clx. - Comments: Originally included the genus Hypoderma É. Geoffroy, 1828 [= Dobsonia Palmer, 1898] (see Jackson and Groves, 2015: 230). McKenna and Bell (1997: 295) synonymized it with Pteropodidae. Pteropi Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 1865: 256. - Comments: Proposed as family name and originally included the genera Pteropus Brisson, 1762; Cynonycteris Peters, 1852 [= Rousettus J. Gray, 1821]; Pterocyon Peters, 1861 [= Eidolon Rafinesque, 1815]; Cynopterus F. Cuvier, 1824 [1821-1825]: 248; Megaerops Peters, 1865; Epomophorus E. Bennett, 1836; Macroglossus F. Cuvier, 1824 [1821-1825]; Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797]; Cephalotes É. Geoffroy, 1810b [= Nyctimene Borkhausen, 1797]; and Notopteris J. Gray, 1859. Pteropodida Haeckel, Generelle Morphologie der Organismen., clx. - Comments: Originally included the genera Pteropus Brisson, 1762; and Macroglossus F. Cuvier, 1824 [1821-1825] (see Jackson and Groves, 2015: 230). McKenna and Bell (1997: 295) synonymized it with Pteropodidae. Pteropi Winge, Samling af Afhandlinger. E Museo Lundii, 2 (1): 53. - Comments: Proposed as tribe(?) and originally included the genera Cynonycteris Peters, 1852b: 25 [= Rousettus J. Gray, 1821]; Pteropus Brisson, 1762; Pteralopex Thomas, 1888; Epomophorus Bennett, 1836; Cephalotes É. Geoffroy, 1810 [= Nyctimene Borkhausen, 1797]; Cynopterus F. Cuvier, 1824 [1821-1825]; and Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797] (see Jackson and Groves, 2015: 231). TAXONOMY: The taxonomy of the Pteropodidae follows that of Bergmans (1988, 1989, 1991, 1994, 1997) and Simmons (2005: 313 - 350). Corbet and Hill (1992: 55) refer to Handley (1980) for a discussion on the correct formation of the family name. Between two and six subfamilies of Pteropodidae have been recognized (Simmons, 2005: 313) including: Cynopterinae K. Andersen, 1912, Epomophorinae K. Andersen, 1912, Harpionycterinae Miller, 1907, Nyctimeninae Miller, 1907, Macroglossinae Gray, 1866, Rousettinae Andersen, 1912, and Pteropodinae Grey, 1821 (Bergmans, 1997; Corbet and Hill, 1980, 1991, 1992; Hill and Smith, 1984; Koopman, 1993a, 1994; McKenna and Bell, 1997). Recent phylogenetic studies agree that Macroglossinae and Pteropodine sensu (Koopman, 1993a, 1994; McKenna and Bell, 1997) are not monophyletic (Álvarez et al., 1999; Giannini and Simmons, 2003; Hollar and Springer, 1997; Hood, 1989; Juste B. et al., 1997; Kirsch et al., 1995; Romagnoli and Springer, 2000; Springer et al., 1995). Monophyly of cynopterines and empomophorines has also been questioned (Álvarez et al., 1999; Hollar and Springer, 1997; Kirsch et al., 1995; Romagnoli and Springer, 2000). Phylogenetic studies based on DNA hybridization and DNA sequences have found support for a large clade of endemic African taxa including genera previously placed in several different subfamilies/tribes based on traditional taxonomic groupings (Álvarez et al., 1999; Giannini and Simmons, 2003; Hollar and Springer, 1997; Kirsch et al., 1995; Romagnoli and Springer, 2000). Relationships among pteropodid genera are not yet fully resolved, however, and questions remain concerning the position of Nyctimene; Paranyctimene, Eidolon, and several SE Asian endemic genera (Simmons, 2005) . Based on a phylogenetic study of newly generated sequences of four nuclear loci, Almeida et al. (2011b: 86) found a basal split of Pteropodidae into 7 (maybe 8) main clades, but the relationships between them are not well resolved. The analyses performed by Almeida et al. (2011a: 7) revealed six principal clades and one independent lineage (for Eidolon). Some of their principal clades agree with African Chiroptera Report 2020 previously proposed subfamilies: Cynopterinae, Harpyionycterinae, and Nyctimeninae. The other clades agree with Macroglossini, Epomophorinae + Rousettini, and Pteropodini + Melonycteris. Almeida et al. (2016: 73) recognize four main African clades: Eidolon, Scotonycterini (including the genera Casinycteris and Scotonycteris), African Rousettus (with three species) and the previously identified 'endemic African clade' (with nine genera: (Stenonycteris, Plerotes, Myonycteris, Megaloglossus, Hypsignathus, Nanonycteris, Epomops, Epomophorus, and Micropteropus), and suggest that a new classification on subfamilial and tribal levels is probably needed. Simmons (2005: 313) suggests the relationships among the pteropodid genera are not yet fully resolved, and no subfamilial or tribal groups were recognized pending a thorough reevaluation of pteropodid classification. Almeida et al. (2016: 83) provide the following classification: Harpyionycterinae (including the genera Dobsonia, Aproteles, Boneia, Harpyionycteris, Rousettinae (including the tribes Rousettini (Rousettus), Eonycterini (Eonycteris), Scotonycterini (Scotonycteris, Casinycteris), Epomophorini (Epomophorus, Epomops, Hypsignathus, Nanonycteris, Micropteropus), Stenonycterini (Stenonycteris), Myonycterini (Myonycteris (incl. Lissonycteris), Megaloglossus), and Plerotini (Plerotes)), and Eidolinae (Eidolon). Hassanin et al. (2016: 521) indicate that two subfamiles occur in Africa: Pteropodinae (with one genus: Eidolon) and Rousettinae. The latter subfamily is subdivided in three tribes: Scotonycterini (Scotonycteris and Casinycteris), Rousettini (Rousettus), and Epomophorini, which is further divided in four subtribes: Epomophorina (Epomophorus, Epomops, Hypsignathus, Micropteropus, and Nanonycteris), Myonycterina (Myonycteris and Megaloglossus), Plerotina (Plerotes), and Stenonycterina (Stenonycteris). They also estimate that the three tribes diversified about 7 - 8 Mya. They also indicate (p. 524) that this family has its origin in Southeast Asia, which is supported by: 1) More than half of the species (94) are found in this region, 2) The highest species density is found on the islands of Sumatra, Borneo and Sulawesi, 3) the oldest fossil of fruit bats has been described in the Late Eocene/Early Oligocene of Thailand, and 4) The molecular analyses indicate that all African species have a derived position with respect to their congeners from Asia and Oceania. Almeida et al. (2020: 11) recognized eight subfamilies: Cynopterinae Andersen, 1912, 29 Macroglossusinae Almeida et al., 2020, Harpyionycterinae Miller, 1907, Rousettinae Andersen, 1912, Eidolinae Almeida et al., 2016, Notopterisinae Almeida et al., 2020, Nyctimeninae Miller, 1907, and Pteropodinae Gray, 1821. Currently (Simmons and Cirranello, 2020) recognized genera of Pteropodidae: Acerodon Jourdan, 1837; Aethalops, Thomas, 1923; Alionycteris Kock, 1969; Aproteles Menzies, 1977; Balionycteris Matschie, 1899; Boneia Jentink, 1879; Casinycteris Thomas, 1910; Chironax K. Andersen, 1912; Cynopterus F. Cuvier, 1824; Desmalopex Miller, 1907; Dobsonia Palmer, 1898; Dyacopterus K. Andersen, 1912; Eidolon Rafinesque, 1815; Eonycteris Dobson, 1873; Epomophorus Bennett, 1836; Epomops Gray, 1870; Haplonycteris Lawrence, 1939; Harpyionycteris Thomas, 1896; Hypsignathus H. Allen, 1861; Latidens Thonglongya, 1972; Macroglossus F. Cuvier, 1824; Megaerops Peters, 1865; Megaloglossus Pagenstecher, 1885; Melonycteris Dobson, 1877; Micropteropus Matschie, 1899; Mirimiti Helgen, 2005; Myonycteris Matschie, 1899; Nanonycteris Matschie, 1899; Neopteryx Hayman, 1946; Notopteris Gray, 1859; Nyctimene Borkhausen, 1797; Otopteropus Kock, 1969; Paranyctimene Tate, 1942; Penthetor K. Andersen, 1912; Plerotes K. Andersen, 1910; † Propotto Simpson, 1967; Ptenochirus Peters, 1861; Pteralopex Thomas, 1888; Pteropus Brisson, 1762; Rousettus Gray, 1821; Scotonycteris Matschie, 1894; Sphaerias Miller, 1906; Stenonycteris Gray, 1870; Styloctenium Matschie, 1899; Syconycteris Matschie, 1899; Thoopterus Matschie, 1899. COMMON NAMES: Czech: kaloňovití, upírové, létací psi, bejložraví netopýři, létací lišky. Dutch: Grote fruitvleermuizen, Vliegende honden. English: Old World Fruit bats. Finnish: Hedelmälepakot. French: Prétopodidés. German: Flughunde, Flederhunde. Italian: Pteropòdidi, Pteròpidi. Norwegian: Flyvende hunder, kalong. Russian: Крылановые. Ukrainian: Летючі лисиці. Vietnamese: Họ dơi qụa. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Gunnell and Simmons (2013: 63) indicate that fossil records for Pteropodid bats are extremely poor. There are hardly any records older than Late Pleistocene-Holocene, although molecular data suggest that Pteropodids diverged from other bats no later than in the Early Eocene (ca. 50 million years ago). They suggest that this is probably due to the fact that most of the Pteropodid fossils are considered to be from Primates rather than from bats. 30 ISSN 1990-6471 The family's stem and crown ages were reported by Shi and Rabosky (2015: 1537) to be respectively 56.6 and 40.2 mya. Foley et al. (2014: 320) indicate that the Rhinolophoidea and Pteropodidae diverged approximately 59 Ma. DETAILED MORPHOLOGY: Dentition - See Adeniyi et al. (2010). Tongue - Morphological studies on the tongue structure of Old World fruit bats suggest that these bats are highly adapted to fruit and nectar diets and are able to change between a fruit and nectar diet when their preferred food is not available (Morrison, 1980; Birt et al., 1997). Old World fruit bat species possess filiform papillae at the tip of the tongue, which is thought to increase the surface area for nectar collection (Birt et al., 1997). Filiform papillae increase the gripping of food on the tongue surface, thus efficiently moving food toward the pharynx (Birt et al., 1997). Bats feeding on fruit also have a large tongue surface area, but generally lack tongue tip filiform papillae, and possess tridentate papillae (Birt et al., 1997). Tridentate papillae are thought to be valuable when piercing through the skin of soft fruits (Birt et al., 1997). Morrison (1980) suggested that fruit bats have a widened tongue and consequently larger tongue surface area that is used to squeeze fruit against the hard upper palate, thus releasing fruit juices. See also Adeniyi et al. (2010). SEXUAL DIMORPHISM: Dobson (1873: 251) suggests that the hairs of the male's epaulettes of the Epomophori are long due to glandular excretions, similar to beard of male Taphozous melanopogon. The different colour, however, can (probably) not be explained by this. ECHOLOCATION: Although the use of clicking sounds as means of echolocation by Rousettus aegyptiacus has become well accepted over the past few years, Boonman et al. (2013a: 23) showed that this type of clicks were also produced by representatives of other genera they investigated: (Eonycteris, Cynopterus and Macroglossus, which convinced Boonman et al. (2013a) that the use of echolocation is more widely distributed than previously believed. Boonman et al. (2014: 2962) provided evidence that suggests that all Old World fruit bats can produce clicks with their wings, resulting in a rudimentary echolocation system. MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) mentioned that the 2n value for the species belonging to this family varies between 24 and 58. Jiao et al. (2018: 4477) found that the bitter taste receptor genes (Tas2rs) in the Old World fruit bats underwent a massive loss (-11 genes out of the ancestral 27) in their common ancestor, but the remaining ones underwent subsequent diversification leading to an average number of genes in the present day species (e.g. 23 in Eidolon helvum and 31 in Rousettus aegyptiacus). DIET: In their study on figs as resources for tropical frugivorous vertebrates, Gautier-Hion and Michaloud (1989: 1826) concluded that these fruits are occurring in such distant patches, that only large frugivorous bats can use them as staple food. Pettersson et al. (2004: 166) state that African pteropodid bats apparently feed more commonly from nectar sources visually observable from above the canopy (as compared with pollinating species from the New World). They also suggest that the absence of sulphur compounts in the floral scent of West African C. pentandra (as opposed to South-America) is the result of the different selective regimes of the pollinating bats. Djossa et al. (2015: 286) found that the baobab tree (Adansonia digitate L.) needs pollinators (= fruit bats) to reach its optimal pollination success. They also concluded that the involvement of fruit bats is beneficial as they travel over long distances, which may help conserve the genetic diversity of this plant species, especially keeping in mind that the baobab tree is a multipurpose socioeconomically relevant plant for Sub-Saharan African rural communities. However, a study by Taylor et al. (2020) revealed that - at least in southern Africa - fruit bats rarely pollinate baobab trees. PREDATORS: Mikula et al. (2016: Supplemental data) report the following birds predating on "Pteropodidae": Chestnut-flanked sparrowhawk (Accipiter castanilius Bonaparte, 1853), Little sparrowhawk (Accipiter minullus (Daudin, 1800)), Bat hawk (Macheiramphus alcinus Bonaparte, 1850), African harrier-hawk (Polyboroides typus Smith, 1829), Crowned eagle (Stephanoaetus coronatus (Linnaeus, 1766)). African Chiroptera Report 2020 PARASITES: Fain (1994: 1279) indicated that the Pteropodidae are hosts for two or three genera of Myobiidae: Binuncus (with subgenus Probinuncus) including 19 species of which 18 occur on Pteropodinae only, and one on Macroglossinae. The second genus is Pteropimyobia with five species of which 3 live on Macroglossinae and two on Nyctimeninae. The third genus - Ugandobia contains nine species occuring on Emballonuridae and one species each occurring on Hipposideridae (U. euthrix) and on Pteropodinae (U. balionycteris). However, he believes that these two latter species were reported from the wrong host species due to "contamination of museum jars". Fain (1994: 1280) also mentions that the Myobiidae found on Megachiroptera are more evolved than those found on the Microchiroptera, indicating that the Myobiidae have probably originated on a primitive genus of the Microchiroptera and subsequently have passed to the Megachiroptera. He also reported this as peculiar since mites belonging to the Mesostigmata and Sarcoptidae are more primitive on Megachiroptera than on Microchiroptera. HEMIPTERA Cimicidae: Haeselbarth et al. (1966: 8) recorded Afrocimex leleupi Schouteden 1951 from fruit-bats in the "Grotte de Kakontwe" and "Grotte Dufrenne", Katanga, DRC. DIPTERA Streblidae: Ascodipteron semirasum Maa 1965 from a fruit bat in Kenya (Haeselbarth et al., 1966: 106). 31 VIRUSES: Nesi et al. (2012: 126) suggest that the high nucleotide distance between Ebola virus Côte d’Ivoire and Ebola virus Zaire can be correlated with the Plio/Pleistocene divergence between their putative reservoir host species, i.e., Myonycteris leptodon and Myonycteris torquata, respectively. Akani et al. (2015: 614 - 615) state "... the first Ebola outbreak in Guinea in 2014 occurred in a site where there was recent deforestation due to the planting of massive oil palm plantations (Wallace et al., 2014), exactly as it also occurred in the Democratic Republic of Congo in 2007 (see Leroy et al., 2009). Therefore, it is likely that there is a direct noncasual link between oil palm plantations, fruit bats and Ebola outbreaks (Wallace et al., 2014). Fruit bats may shift from using rainforest to utilizing oil palm plantations as foraging grounds once the forests became disturbed, especially where plantations are adjacent to rainforest patches. The intensified use of oil palm plantations by fruit bats may be due to the high food resource available due to the palm fruits" [Note (30 September 2020): As far as the editors are aware, no proof has yet been provided that fruit bats are indeed the culprits.] DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burkina Faso, Central African Republic, Gabon, Ghana, Kenya, Madagascar, Senegal, Tanzania. Subfamily Eidolinae Almeida, Giannini and Simmons, 2016 *2016. Eidolinae Almeida, Giannini and Simmons, Acta Chiropt., 18 (1): 83, 84. Publication date: June 2016 [Goto Description]. TAXONOMY: This subfamily was created by Almeida et al. (2016: 84) to distinguish monophyletic groups within the Pteropodidae. Currently recognized genera of Eidolinae: Eidolon Rafinesque, 1815. Genus Eidolon Rafinesque, 1815 *1815. Eidolon Rafinesque, Analyse de la Nature, 45. - Comments: Type species: Pteropus stamineus E. Geoffroy Saint-Hilaire, 1803 (= Vespertilio vampyrus helvus Kerr, 1792). Etymology: From the Greek "ειδωλον", meaning "image" or "phantom", evidently in allusion to its movements (see Palmer, 1904: 252; DeFrees and Wilson, 1988: 4). (Current Combination) 1862. Pterocyon Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 1861: 423. - Comments: Type species: Pterocyon paleaceus Peters, 1861 (=Vespertilio vampyrus helvus Kerr, 1792). Etymology: From the Greek "πτερόν", meaning wing and "κύων", meaning dog (see Palmer, 1904: 595). 32 ISSN 1990-6471 1881. 1882. Leiponyx Jentink, Notes Leyden Mus., 3 (15): 60. Publication date: April 1881 [Goto Description]. - Comments: Type species: Leiponyx büttikoferi Jentink, 1881 (=Vespertilio vampyrus helvus Kerr, 1792). - Etymology: From the Greek "λείπω", meaning to leave or to be wanting and "ονυξ", meaning claw, referring to the absence of a claw on the index finger (see Palmer, 1904: 367). Liponyx Forbes, Zoological Record, 18 Mamm.: 13 (for 1881). - Comments: Emendation of Leiponyx Jentink, 1881. Preoccupied by Liponyx Vieillot, 1816; a bird (see Andersen, 1912b: 2; Meester et al., 1986: 27). (Emendation) TAXONOMY: Revised by Bergmans (1991). (2005: 320). See Simmons Almeida et al. (2011a: 11) suggest that Eidolon should be included in a subfamily of its own. Currently (Simmons and Cirranello, 2020) recognized species of the genus Eidolon: dupreanum (Pollen, 1866); helvum (Kerr, 1792). COMMON NAMES: Czech: plaví kaloni. English: Straw-coloured Fruit-bats, Eidolon fruit bats. French: Roussettes des palmiers. German: Palmenflughunde. Italian: Èidoli, Ptèropi fantasrna. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Shi et al. (2014: 285) refer to Howell and Coppens (1974), who reported the only fossil record for Eidolon from the Pliocene (ca. 3 mya) of Ethiopia. They also indicate (p. 279) that the two taxa (helvum and dupreanum) diverged in the mid-tolate Miocene. Almeida et al. (2016: 85) place the origin of the genus in the Miocene or Oligocene. VIRUSES: Flaviviridae Luis et al. (2013: suppl.) mention the following Flaviviridae from "Eidolon": Entebbe bat virus, Uganda S virus, Usutu virus, West Nile virus, Yellow fever virus, and Zika virus. Togaviridae Alphavirus - (Luis et al., 2013: suppl.) mention "from Eidolon". Chikungunya virus - (Luis et al., 2013: suppl.) mention "from Eidolon". Eidolon dupreanum (Schegel, 1867) *1867. Pteropus dupréanus Schlegel, in: Schlegen and Pollen, Proc. zool. Soc. Lond., 1866, III: 419. Publication date: April 1867. Type locality: Madagascar: Nossi Bé island [ca. 13 20 S 48 15 E] [Goto Description]. - Comments: Andersen (1912b: 8) mentions two cotypes in the Leyden Museum collected by Fr. Pollen and C.C. van Dam (see Schlegel, 1867: 419). 2014. Eidolon duprenum: Gunnell, Simmons and Seiffert, PLoS ONE, 9 (2): e86712: 3. Publication date: 4 February 2014. (Lapsus) ? Eidolon dupreanum: (Name Combination, Current Combination) ? Eidolon helvum dupreanum: (Name Combination) TAXONOMY: Koopman (1993a: 140) mentioned Schlegel, 1867 as author, but Allen (1939a: 54), Peterson et al. (1995: 22) and Russ et al. (2001) mention Pollen, 1866, in Schlegel and Pollen, Proc. Zool. Soc. Lond., 1866: 419. Meester et al. (1986: 28), DeFrees and Wilson (1988: 1) mention Schlegel and Pollen, 1867 as authors, but in the references of the latter only Schlegel is mentioned as author. Reviewed by Bergmans (1991) and Peterson et al. (1995). See Simmons (2005: 321 and comments under E. helvum pg. 321). COMMON NAMES: Czech: kaloň malgašský. English: Madagascan Straw-coloured Fruit-bat, Madagascan Fruit Bat, Malagasy Straw-colored Fruit Bat. French: Roussette des palmiers malgache, Roussette malgache couleur paille, Roussette jaune de Madagascar. German: MadagaskarPalmenflughund. Malagasy: Angavo. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Samonds (2007: 43) found samples in deposits dated at 10,000 yrs old or younger from within the Anjohibe cave, Madagascar. Gunnell et al. (2014: 3) also refer to Goodman and Jungers (2013), who report material from the Andrahomana cave. CONSERVATION STATUS: Global Justification African Chiroptera Report 2020 33 This species is listed as Vulnerable (VU A2ad ver 3.1 (2001)) because it is estimated to have undergone a decline exceeding 30% over the past 20 years mainly from extreme pressure from hunting, causing it to abandon roosts, with known examples of local extirpation of roosts sites (Andriafidison et al., 2008b; IUCN, 2009). National Park, even in sacred caves. Roosts that are located in inaccessible rock outcrops pose significant challenges to hunters are relatively protected. Cave roosts are probably subject to highest hunting pressure and conservation measures should be focused at these sites (Andriafidison et al., 2008b; IUCN, 2009). Assessment History Global 2008: VU A2ad ver 3.1 (2001) (Andriafidison et al., 2008b; IUCN, 2009). 1996: LR/lc (Baillie and Groombridge, 1996). Additional research is needed on natural history and in particular roosting ecology and movements (Andriafidison et al., 2008b; IUCN, 2009). Regional None known. MAJOR THREATS: This species is subject to harvesting for bushmeat across its range and hunting occurs both at roosting and foraging sites (Jenkins and Racey, 2008). It is classed as a game species under Malagasy law and although it can only be legally hunted between the first of May and the first of September (Anonymus, 2007b: 6; Durbin, 2007; Randrianandrianina et al., 2010: 412), this legislation is widely ignored and the bats are hunted throughout the year. Its roosts tend to be well protected from bushfires and it is able to survive in landscapes with severely depleted natural food supplies as long as alternative plants are available (Ratrimomanarivo, 2007). Hunting is therefore the main threat and was reported to account for a 30% desertion rate from 60 roosts (MacKinnon et al., 2003). Although hunters report that deserted roosts are recolonized after a few years, there are several confirmed examples where this species has been extirpated at roost sites (MacKinnon et al., 2003). Randrianandrianina et al. (2010: 413) observed that at least 14 E. dupreanum specimens were captured each night at fruiting voandelaka tree (Molinaea sp.). Cardiff and Jenkins (2016: 99) indicate that fires are used to expel E. dupreanum from their rocky roost sites. CONSERVATION ACTIONS: Andriafidison et al. (2008b) [in IUCN (2009)] report this species is known from a number of protected areas (Goodman et al., 2005a) including, Parc National d’Isalo, Parc National de Namoroka, Réserve Spéciale d’Ankarana and Réserve Spéciale de Cape Sainte Marie; however, hunting has been reported from within Réserve Spéciale d’Ankarana (Cardiff, 2006). FernándezLlamazares et al. (2018: 273), similarly, reported hunting in and outside the Tsimanampetsotsa GENERAL DISTRIBUTION: Eidolon dupreanum is endemic to the island of Madagascar where it is widespread, albeit with a patchy distribution (MacKinnon et al., 2003). It is found both along the coast and on the central high plateau and areas from where there are no records probably reflect inadequate survey coverage rather than a genuine absence. There are some areas within its range where the lack of crevices means there are no roosting opportunities (Goodman et al., 2005a). Native: Madagascar. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 52) describe the baculum as a large flat structure, with the distal tip having a serrated or tooth-like structure that is wider than the proximal base; length: 6.59 ± 0.225 (6.41 - 6.84) mm, width: 4.21 ± 0.671 (3.48 - 4.80) mm. ECHOLOCATION: Schoeman and Goodman (2012) identified three distinct call types: social calls 1, social calls 2 and echo clicks. Bats produce the echo clicks while flying towards the entrance of the cave, while the other two calls were emitted at roost sites and near the cave entrance. Based on the preliminary results, Schoeman and Goodman (2012) suggest that the social calls 1 and 2 were employed for social communication, whilst echo clicks may have been used in a sensory context, potentially as incipient echolocation to navigate in the dark cave. HABITAT: Eidolon dupreanum has a patchy distribution which includes humid, dry deciduous and spiny forest is probably related to the availability of suitable roost sites and is rare or absent from a number of forests without rocky outcrops (Goodman et al., 2005a; Jenkins et al., 2007b; Schmid and Alonso, 2007). It continues to survive in highly modified landscapes with very little native vegetation remaining (Ratrimomanarivo, 2007), but appears to use 34 ISSN 1990-6471 native forest vegetation for food in preference to introduced plants (Picot et al., 2007). Dammhahn and Goodman (2013: 108) indicate that mid- to upper-canopy in forest and trees growing in open areas form the foraging habitat of this species. HABITS: Baum (1995: 337) reported that fruit bats (probably E. dupreanum) intensively visited Adansonia suarezensis trees in the forest at Beantely in the hour after dark, but also during the remainder of the night at a much lower frequency. Peak rates were about 14 visits per tree per hour (or ca. 0.5 visits / flower / hr), each lasting between 20 and 30 seconds. During each of the tree stopovers, up to nine flowers were visited. While exiting roost, individuals flew within 50–100 cm of the ceiling, often settling and perching every 5–10 m along their flight patch and displacing in a leapfrog manner towards the cave entrance (Schoeman and Goodman, 2012). ROOST: It is known to roost in tree foliage, but is more usually found in rock fissures and caves (MacKinnon et al., 2003). Its roost sites are usually very high and located in cliffs and rock faces (Jenkins and Racey, 2008). In the Réserve Spéciale d’Ankarana, it used caves that were high and long with a good buffering capacity for temperature and humidity (Cardiff, 2006). See habits for roosting behaviour. MIGRATION: MacKinnon et al. (2003) suggested that E. dupreanum might be migratory because of regular variation in the occupancy and abundance of roosts. DIET: In a study, during three months (February, July and December), at four sites on the central Malagasy highlands, Ratrimomanarivo (2007) found the seeds of 13 plant species. Picot et al. (2007) reported that E. dupreanum ate mainly fruits (65%), although Eucalyptus spp. flowers were also consumed. Thirty plant species (14 identified and 16 unidentified), including six introduced species were utilized in the corridor of rainforest between Ranomafana and Andringitra National Parks, south-eastern Madagascar (Picot et al., 2007). Polyscias spp. fruit (seeds) were the most frequently recorded plant in 70 % of the faecal samples (Picot et al., 2007). Fruit is the main dietary component, but it also eats leaves and other plant parts (Picot, 2005; Picot et al., 2007). Through the ingestion of pollen and small seeds, a significant ecological role is inferred (Picot, 2005; Ratrimomanarivo, 2007). E. dupreanum may be an important pollinator of threatened baobab trees (Baum, 1995). Baum (1995: 339) mentions that E. dupreanum is believed to be one of the major pollinators of endemic baobabs Adansonia grandidieri and Adansonia suarezensis of Madagascar. Picot et al. (2007) suggest that the alimentary tract is one intrinsic factor that influences seed dispersal, as they found a maximum seed size of 7 mm and mean size of 3.2 mm for E. dupreanum and the ecological services provided by this species appear to be limited by the dimensions of its intestinal tract. Jenkins et al. (2007b: 213) [referring to Rakotonirainy (2001)] mention these bats visit fruiting trees of Ficus soroceoides, F. pyrifolia, and F. guatteriifolia. Based on stable isotope analyses, Reuter et al. (2016c) found that Pteropus rufus, Eidolon dupreanum and Rousettus madagascariensis have broadly overlapping diets, although there are differences between P. rufus and E. dupreanum. Furthermore, they found that the diet shifted when these two species occurred sympatrically. POPULATION: Structure and Density:- The population size and local abundance of this species are not well known. Estimates of colony size are difficult to obtain because the bats are hidden during the day, but is usually in the range of between 10 and 500, with a median of 200 individuals (MacKinnon et al., 2003). The maximum colony size of 1,400 is from Réserve Spéciale d’Ankarana (S. G. Cardiff unpubl. [in Andriafidison et al. (2008b)]). Given the estimated adult survival, Brook et al. (2019a: 165) found that the population of E. dupreanum will only be able to persist if the fieldbased fecundity estimates were replaced by higher values reported in the literature for related species. Trend:- 2008: Decreasing (Andriafidison et al., 2008b; IUCN, 2009). REPRODUCTION AND ONTOGENY: Kaudern (1914: 11 - 12) reported that he was unable to shoot any pregnant female or females with young during his travels on Madagascar, which took place in the dry season (May to September), so he assumed they had their reproductive season in Spring or Summer. African Chiroptera Report 2020 PARASITES: Brook et al. (2015: 4) found E. dupreanum to be the host of Cyclopodia dubia (Westwood, 1835) (Nycteribiidae) and Thaumapsylla sp. fleas. In the bat flies (and the bats), DNA of a new Bartonella was found. The bat flies were also reported by Wilkinson et al. (2016) and Ramasindrazana et al. (2017: Suppl). Szentiványi et al. (2019: Suppl.) indicated that thes bat flies were also carrying Enterobacteriales and Wolbachia sp. bacteria. VIRUSES: Reynes et al. (2011: 3) were able to isolate Ife virus from E. dupreanum. Coronaviridae - Coronaviruses BtCoVMAH051F/E. dupreanum/2011 In Madagascar, identified in faecal material (Razanajatovo et al., 2015). Herpesviridae Alphaherpesvirinae Simplexvirus Alphaherpesvirus CS730G - In Madagascar 2006, isolated in Vero cells from throat swabs (Razafindratsimandresy et al., 2009). Paramyxoviridae Henipavirus Cedar (CedPV) - Brook et al. (2019b: 1004) reported the presence of reliable reactive antibodies to tested antigens in serum from E. dupreanum specimens. Hendra (HeV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) found Hendra virus with an infection rate of 19.2%. Nipah (NiV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) found Nipah virus with an infection rate of 19.2%. Kuzmin et al. (2011a: 5) suggest that their exposure to NiV might be the result of contact with infected Pteropus rufus specimens. Tioman (TiV) - In Madagascar during 2003 - 2004, this virus was not detected (Iehlé et al., 2007). 2003). Hunting is undertaken by specialists (due to the roosting high in cliffs) who either access the roosts with wooden ladders (Jenkins and Racey, 2008) or light fires directly underneath the roosts and net or swat flying bats as they emerge (Ranivo, 2001). Alternatively, a hunter is lowered on a rope and hits the bats with a stick (Jenkins and Racey, 2008). Jenkins and Racey (2008) also indicate that guns and slingshots are used. In the rainforests between PN de Ranomafana and PN d'Andringitra, a number of accounts of bat hunting were collected from villages (Picot, 2005). Although familiar with the location of the bat roosts, the local people rarely hunted or consumed the bats. Professional hunters visited a few times a year and occasionally offered bat meat to the village but most of the animals were taken to larger towns and sold for $ 0.4 US to restaurants and hotels. In this area oil residue from cooked bats is used as medicine for whooping cough (Jenkins and Racey, 2008). Further north in the highlands, Ranivo (2001) investigated the hunting of E. dupreanum at roost sites near Ambositra, Antsirabe and Ankazobe, where villages reported purchasing one and four times per year. Pteropus rufus and Eidolon dupreanum appear to be the only species sold in restaurants, where they are either served individually, complete with head and wings, jointed, or are diced up into small pieces and accompanied by rice (Jenkins and Racey, 2008). Ranivo (2001) found that hunting caused the bats to abandon traditional roost sites and local people noted declines in hunting returns. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar. Rhabdoviridae – Rabies like viruses Lyssavirus - Rabies related viruses Andriamandimby et al. (2013: 3) reported that 17 bats testes positive for the detection of antibodies agains Lagos Bat Virus (LBV) in two areas close to Antananarivo. Five of these also tested positive against European Bat Lyssavirus type 1 (EBLV-1). UTILISATION: E. dupreanum is hunted for meat in many parts of Madagascar (Ranivo, 2001; MacKinnon et al., 35 Figure 5. Distribution of Eidolon dupreanum 36 ISSN 1990-6471 Eidolon helvum (Kerr, 1792) *1792. Vespertilio vampyrus helvus Kerr, in: Linnaeus, Animal Kingdom, 1 (1): xvii, 91. Publication date: February 1792. Type locality: Senegal: "Senegal". - Comments: Type locality fixed by Andersen (1907a: 504); see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1), Harrison and Bates (1991). Andersen (1912b: viii) indicates that the type was once in the Museum Leverianum, but is probably no longer in existence. - Etymology: From the Latin "helvus" (=dun-coloured), referring to the straw colour on their shoulders; the back, rump and hind limbs are normally various shades of brown (see DeFrees and Wilson, 1988: 4; Taylor, 2005). Lanza et al. (2015: 27) translate "helvus" to "dull-yellow". 1862. Xantharpyia stramineus: Gerrard, Catalogue of the bones of Mammalia in the collection of the British Museum, 58. (Name Combination) 1866. Xantharpyia leucomelas Heuglin, in: Heuglin and Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 54: sect. 1, pt. 10, p. 544. Type locality: Sudan: Kordofan: Bahr-elAzrak: Bahr-el-Abiad: Sennaar. - Comments: Originally "Upper Nile lands", but subsequently fixed to Senaar by Koopman (1975): see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1). 1867. Cynonycteris straminea: Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 866. - Comments: see Pousargues (1896: 257). (Name Combination) 1895. Cynonicteris straminea: Bocage, J. Sci. mat. phys. nat., ser. 2, 4 (8): 4. Publication date: December 1895. (Name Combination, Lapsus) 1906. R[oussettus (Xantharpyia)] stramineus: Johnston, Liberia, Volume II: 690. (Name Combination, Lapsus) 1920. Rousettus stramineus: Haagner, South African mammals, 23. (Name Combination) 2015. Pterocyon palaeaceus: Lanza, Funaioli and Riccucci, The bats of Somalia and neighbouring areas, 27. (Lapsus) 2019. Eidolon hevlum: Debata, Panda and Palita, Biodivers. Conserv., 28 (8-9 SI): 2386. Publication date: 13 February 2019. (Lapsus) ? Eidolon helvum: (Name Combination, Current Combination) TAXONOMY: Includes sabaeum see Hayman and Hill (1971), Bergmans (1991), Harrison and Bates (1991), Simmons (2005: 321). Does not include dupreanum; see Bergmans (1991), Peterson et al. (1995) and Simmons (2005: 321), but also see Hayman and Hill (1971) who treated dupreanum Schlegel, 1866, from Madagascar and sabaeum Andersen, 1907, from Arabia as subspecies of E. helvum (Meester et al., 1986). African forms reviewed in part by Juste et al. (2000); Palearctic forms reviewed by Horácek et al. (2000). The taxonomic status of populations in the Anr Mountains of Niger is unclear (Simmons (2005: 321). E. dupreanum is here retained as a separate species and sabaeum is considered an extralimital subspecies of E. helvum not occuring in Africa. Juste et al. (2000) found that the population from Annobón is significantly smaller than the continental forms (and the specimens from the other islands in the Gulf of Guinea) and created a new subspecies for these: Eidolon helvum annobonensis. Currently recognized subspecies helvum: of Eidolon E. h. helvum (Kerr, 1792) - Afrotropics. E. h. annobonensis Juste et al. 2000 - islands off Gulf of Guinea. E. h. sabaeum K. Andersen, 1907 - Oriental. COMMON NAMES: Afrikaans: Geel vrugtevlermuis. Castilian (Spain): Zorra Voladora Amarilla. Chinese: 黄毛 果 蝠 . Czech: kaloň plavý, upír bledý, kaloň palmový, kaloň úzkokřídlý. English: African Straw-coloured Fruit-bat, Straw-colored Fruit Bat, Straw-coloured Flying Fox, Pale Xantharpy, Yellow-haired fruit bat. French: Roussette jaune, Roussette des palmiers africaine, Roussette paillée, chauve-souris paillée. German: Afrikanischer Palmenflughund, Palmenflughund, Afrikanischer strohfarbener Flughund. Italian: Èidolo paglierìno, Ptèropo fantàsma paglierìno. Kinande (DRC): Mulima. Portuguese: Morcego frugivoro gigante. Tanoboase (Ghana): ampane ankasa. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Hildebrand et al. (2010: 268) recorded this species from the Kumali site in southern Ethiopia. CONSERVATION STATUS: Global Justification African Chiroptera Report 2020 Listed as Near Threatened (NT ver 3.1 (2001)) because this species is in significant decline (but at a rate of less than 30% over three generations (approximately fifteen years)) because it is being seriously over-harvested for food and medicine, making the species close to qualifying for Vulnerable, Under criterion A2d (Mickleburgh et al., 2008dc; IUCN, 2009). The subspecies E. h. sabaeum (Andersen, 1907) is known only from Yemen and Saudi Arabia and is possibly threatened (see Bergmans, 1999). Assessment History Global 2008: NT ver 3.1 (2001) (Mickleburgh et al., 2008dc; IUCN, 2009). 2004: LC (Mickleburgh et al., 2004bh; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016e). 2004: considered a vagrant not assessed, doesn't breed in SA (Friedmann and Daly, 2004). MAJOR THREATS: In general there are no major threats to this widespread and adaptable species. It is, however, locally threatened in parts of its range by severe deforestation, and more generally across West and Central Africa by hunting for food and medicinal use. It is the most heavily harvested bat for bushmeat in West and Central Africa, and one of the most frequently consumed mammals in this region (P. Racey pers comm. in Mickleburgh et al. (2008dc)). Large pre-migration colonies are considered particularly vulnerable to any threats. In some areas it is considered to be a pest species and roosting locations may be restricted by cutting down trees. Trees are also cut down in order to catch bats for the market (P. Racey pers comm. [in Mickleburgh et al. (2008dc)]). Kamins et al. (2010: 190) report that 41 hunters, they interviewed, killed about 35,000 bats in one year in southeastern Ghana. 95 meat vendors indicated that they sold over 100,000 bats as food in one year. These bats were captured in an area up to 200 km away from the markets where they were sold. Kamins et al. (2011a: 3000) update these figures, based on interviews with 551 Ghanaians, to a minimum 128,000 bats being sold yearly, up to 400 km away from the markets. MacDonald et al. (2011: Suppl. Table 1) indicate that, in rural markets in Cameroon and Nigeria, fresh bats were sold at 2.96 US$, or about 10.6 US$/kg, and smoked bats at 1.63 US$ or 5.82 US$/kg. In Côte d'Ivoire, Niamien et al. (2015a: 233) found that poachers only ate 5 % of their catch themselves, the rest was sold for 0.6 US$ per animal, leading 37 to a daily revenu of 15 US$. They estimated that some 30,600 were killed each month, but luckily for the bats, they made some major calculation errors. On São Tomé, these bats are extensively harvested for household consumption (see Carvalho et al., 2015: 1). Hayman and Peel (2016: 132) indicate that the smaller island populations [from São Tomé, Príncipe and Annobón] may more likely suffer from additive mortality in response to harvesting than populations from the nearby Bioko as these latter can still mix freely with the continental populations. Musaba Akawa et al. (2017) followed the sale of E. helvum at a market in Kisangani (DRC) for one year and found that most of the bats were sold during one of the wet seasons (SeptemberOctober), but low during the second wet season (March-April). They also found that while hunting was allowed from December through July, only 47 % of the total number of bats were sold during that period; 53 % was sold during the period that the hunt was closed. Anonymus (2004f: 51) indicates that their colonies are unwelcome in towns and cities through fruit feeding, defoliation of roost trees, defecation on (commercial) buildings. CONSERVATION ACTIONS: Mickleburgh et al. (2008dc) [in IUCN (2009)] report that there are no known direct conservation measures in place, but in view of the species extensive range it seems probable it is present in some protected areas during migration. There is a need to identify and protect important roosting sites, and a better understanding of the migratory patterns of this species would be beneficial to any conservation activities. The highest priority is to limit the harvesting of this species to sustainable levels. GENERAL DISTRIBUTION: Eidolon helvum is broadly distributed across the lowland rainforest and savanna zones of Africa from Senegal in the west, through to South Africa in the south and Ethiopia in the east (possibly ranging into Djibouti and southern Eritrea). It is also present on the extreme southwest Arabian Peninsula, where it has been recorded from Yemen and Saudi Arabia (Harrison and Bates, 1991). Populations of this bat occur on several offshore islands including the Gulf of Guinea islands (Peel et al., 2013a: 296) and Zanzibar, Pemba and Mafia (off Tanzania) (Bergmans, 1991; Simmons, 2005; O'Brien, 2011: 285). It has also been recorded at sea, 250 km from the nearest land (Anonymus, 2004f: 50). There is a possibly 38 ISSN 1990-6471 disjunct population in the Air Mountains of Niger. Distribution at northern and southern extremes of the range is patchy and erratic. It is also sparse or absent in large areas of the Horn of Africa, central East Africa, and elsewhere (Bergmans, 1991). Eidolon helvum is a migratory species in parts of its range; populations migrate from the West African forest north into the savanna zone during the major wet season. It ranges from sea level to around 2,000 m asl (Ruwenzori Mountains). In Cameroon, Dongmo et al. (2020: 51) reported it from mid-elevated areas (1,220 – 1,320 m). In South Africa, its distribution is mostly affected by precipitation seasonality (Babiker Salata, 2012: 49). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,091,925 km2. It has also been reported from the Cape Verde Islands (Jiménez and Hazevoet, 2010: 116). Native: Angola (Crawford-Cabral, 1989; Lopes and Crawford-Cabral, 1992; Bergmans, 1991; Monadjem et al., 2010d: 551); Benin (Capo-Chichi et al., 2004: 161); Botswana; Burkina Faso (Kangoyé et al., 2015a: 605); Burundi; Cape Verde Islands (Jiménez and Hazevoet, 2010: 116); Cameroon; Central African Republic; Chad; Congo (Bates et al., 2013: 333); Congo (The Democratic Republic of the) (Schouteden, 1944; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 551); Côte d'Ivoire (Henry et al., 2004: 24); Equatorial Guinea; Ethiopia; Gabon; Gambia (Kock et al., 2002: 78); Ghana; Guinea (Fahr and Ebigbo, 2003: 128; Decher et al., 2016: 259); Guinea-Bissau (Bocage, 1892b; Veiga-Ferreira, 1948; Bergmans, 1991; Corbet, 1984: 30) ; Kenya; Lesotho; Liberia (Fahr, 2007a: 103); Malawi (Happold et al., 1988; Ansell and Dowsett, 1988: 28; Bergmans, 1991; Monadjem et al., 2010d: 551); Mali (Meinig, 2000: 104); Mauritania; Mozambique (Smithers and Lobão Tello, 1976; Cotterill, 2001e: 54; Monadjem et al., 2010d: 551); Namibia (Bergmans, 1991; Monadjem et al., 2010d: 551); Niger (Bergmans, 1991 in Cotterill, 2001e, 2001e: 55); Nigeria; Rwanda; Sao Tomé and Principe (Rainho et al., 2010a: 24); Saudi Arabia; Senegal; Sierra Leone; South Africa (Monadjem et al., 2010d: 551); Sudan; Swaziland; Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 58); Yemen; Zambia (Ansell, 1974; Cotterill, 2001e: 54; Monadjem et al., 2010d: 551); Zimbabwe (Monadjem et al., 2010d: 551). Presence uncertain: Djibouti; Eritrea. GEOGRAPHIC VARIATION: Peel et al. (2010b: 245) indicate that the nonmigratory populations on the islands in the Gulf of Guinea (São Tomé and Príncipe, and Bioko and Annobón) show evidence of genetic differentiation from the continental (migratory) populations. They also indicate that two or three colonisation events to the islands have taken place. Juste et al. (2000) found that the population from Annobón is significantly smaller than the continental forms (and the specimens from the other islands in the Gulf of Guinea) and created a new subspecies for these: Eidolon helvum annobonensis. DENTAL FORMULA: Popa et al. (2016: 847) described the replacement of the deciduous (milk) teeth by the permanent ones. The dentition consists of: Upper deciduous dentition: incisors i1, i2; canines c1; premolars pm1, pm2, pm3. Lower deciduous dentition: incisors i1, i2; canines c1; premolars pm1, pm2, pm3. Upper permanent dentition: incisors I1, I2; canines C1; premolars PM2, PM3; molars M1, M2. Lower permanent dentition: incisors I1, I2; canines C1; premolars PM2, PM3; molars M1, M2, M3. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 54) refer to Nwoha et al. (2000: 154), who found that the baculum in juvenile bats consists of two small paddlestone pieces located longitudinally close to each other at the distal third of the penis. In adult bats, the baculum consists of a large distal portion and a narrow proximal portion. The average brain mass for this species is 4.30 g [n = 2, Chawana et al. (2013: 160)]. Huggel (1959) studied the blood pressure in the wings of E. helvum from Côte d'Ivoire. The normal pressure varied between 23 and 30 cm H2O. This pressure was the same for the two types of bloodvessels that are present in the wing (contracting or non-contracting). If there is some blocking, the pressure can double very quickly, and in one case it even trippled. Nwoha (2000: 291) examined the pelvis of this bat and found sexual dimorphism both in the shape and size of these bones as well as in its development. In males, the ossification of the interpubic ligament already start in the juvenile stage, whereas in females this only begins with puberty. Odukoya et al. (2009b: 175) investigated the biochemical differences in the two uterine limbs during pregnancy and found that the uterine limbs actively utilized different metabolic pathways according to the functions they subserve during pregnancy. African Chiroptera Report 2020 Adeniyi et al. (2012: 118) compared the lateral geniculate bodies (LGB) of E. helvum with those of Rattus norvegicus and Manis tricuspis and found some differences, which they attributed to the feeding habits, lifestyle as well as the nature of their habitat. The morphometrics of the reproductive tract of male bats were examined by Danmaigoro et al. (2014a), whereas Danmaigoro et al. (2014b) examined the histology and the histometric anatomy. Abedi-Lartey et al. (2016: 18) found that the average gut passage time for small seeds was 116 ± 112 minutes (4 - 1,143 min). The dispersal distances for the small and large seeds were longer in rural areas than in urban areas, and for large seeds and small seeds in urban areas, they were also longer in the dry season than in the wet season. For small seeds in rural areas, they found that the distances were longer in the wet season than in the dry season. FUNCTIONAL MORPHOLOGY: Church and Warren (1968) indicate that wing membranes of Eidolon helvum are composed of long, prominent strands of elastin with collagen stretched in between, and that 2 x 2 cm holes in wings heal in 24 days. Referring to Riskin et al. (2010), O'Mara et al. (2016: 10) mention that the wing camber of E. helvum shows more variation than in nonmigrating flying foxes. O'Mara et al. (2019a: 1) found that wingbeat frequency has the strongest positive relationship with ODBA (overall dynamic body acceleration) and that there is small, negative correlation between ODBA and airspeed. ODBA is also decreased with increasing tailwind. Igado et al. (2015: 283) studied the tongue, hard palate and buccal cavity of this bat. They found that the overal distribution of lingual papillae was the same for both sexes, but the weight and the length of the tongue was higher in females (although the difference was not significant). Also the number of ridges on the hard palate showed a sexual dimorphism (see also the sexual dimorphism section). The eyes of E. helvum contain approximately 280,000 retinal ganglion cells. The minimum angle of resolution was approximately 0.122° (ca. 4 mm at 1 m distance). Based on a histological investigation, Mainoya and Howell (1979: 162) suggest that the neck skin patch in E. helvum - consisting of long and coarse hairs, and extending from the ventral surface of the 39 neck to the shoulder area - plays a role in scent emission. SEXUAL DIMORPHISM: In Nigeria, Igado et al. (2015: 285) examined the heads of 30 adult males and females and found the males to have larger values for overal weight, weight of the head, thickness of the tongue (at the torus), width of the tongue (at the root), width of the hard palate (at the commissures of the mouth), and width of the hard palate (at the anterior region and at the posterior half). Additionally, the number of palatine ridges was higher in males (♂♂: 8.33 ± 0.577; ♀♀: 7.667 ± 0.577). The width of the tongue at the apex was similar in both species, and the width of the tongue at the torus was significantly larger in females. In a previous study, Igado et al. (2012: 172) already found that males had higher values for most of the craniofacial and neurometric parameters they examined. The only parameter where females had higher values was the width of the left external nares (♀♀: 5.08 ± 0.55 mm; ♂♂: 5.0 ± 0.39 mm). MOLECULAR BIOLOGY: DNA - Peel et al. (2010a) developed twenty microsatellite loci, from 142 individuals sampled from nine populations across Africa. Teeling et al. (2017: 32) mention the estimated genome size for E. helvum to be 2.03 C, and the assembly size 1.44 Gb. Karyotype - Mattey (1962) and Rushton (1970: 460) reported 2n = 34. Protein / allozyme - Unknown. The phosphate and oxygene binding capacity of the bat's haemoglobin was studied by Fodeke (2017). Stasiak et al. (2018: 386) investigated he role of Hepcidin in the iron metabolism of Eidolon helvum, a species that rarely developed iron storage disease (hemochromatosis). They found that Hepcidin mRNA expression doesn't increase in response to iron administration in healthy animals. HABITAT: This adaptable species has been recorded from a very wide range of habitats. It is commonly found in moist and dry tropical rain forest, including evergreen forest habitats in the form of coastal (including mangrove) and riverine forest, through moist and dry savanna and mosaics of these and similar habitat types. Populations can persist in modified habitats and the species is often recorded in urban areas, such as wooded city parks (Mickleburgh et al., 2008dc). 40 ISSN 1990-6471 HABITS: Calderón-Capote et al. (2020: 1) tracked the movements in four colonies (ranging in size from 4,000 to 10,000,000 animals) and found that average distance covered per night (9 - 99 km), number of foraging sites visited per night (2 - 3) as well as foraging and commuting times were largely independent of colony size. In general, all the bats returned to the same day roost each night. ROOST: Roosting trees often become defoliated due to the large number of bats inhabiting them and are therefore only used on a relatively limited temporal scale (Kunz, 1996: 42). The extent of the damage caused by the bats at the Awolowo University Campus, Ile-Ife, Nigeria, was studied by Ayoade et al. (2012: 405), who reported that premature defoliation, loss of branches and hence reduction in canopy foliage of the host trees led to the decline in the ability of the trees to provide many environmental and social services that contribute to the quality of life in cities and to serve as effective wind barrier. Ayensu (1974: 716) mentions that Ghanese E. helvum roosting in the palms do not feed on the fruits of the same palm tree, but fly to another palm to feed. In Accra, the bats roost in large mahogany and neem trees at the 37 Military Hospital, the Department of Parks and Gardens and the 37 Military Barracks (Ohemeng et al. (2018: 3). Lawson et al. (2018: 110) also found them roosting in huge silk cotton trees (Ceiba pentandra). In Côte d'Ivoire, Niamien et al. (2017a: 70) found that E. helvum prefers to roost in Hevea brasiliensis Müll. Arg. (Pará rubber tree Euphorbiaceae) and Mangifera indica L. (Mango Anacardiaceae) trees. For western Kenya - the only colony site in Kenya - Webala et al. (2012: 3) suggested that tree density was an important factor in roosting-camp selection for E. helvum, and the removal of roost trees could have serious ramifications on their conservation in the region. This is especially important as colonies of E. helvum are rarely found in protected areas, but rather within human habitation. Webala et al. (2014: 85) found that, besides the density of the trees, also the number of branches on a tree was an important factor when selecting a roost site. Peel et al. (2017: 77) provided information on distribution, migration patterns, roost size, age and seks composition of 29 E. helvum roosts from nine countries across tropical Africa. They indicate (p. 89) that roosts in urban areas are in zones where hunting is discouraged due to their human usage: hospital grounds, embassies, botanical gardens, hotels, palace gardens, zoos, and military, private or corporate compounds. These zones are usually locations where sufficient numbers of large, tall trees are present. MIGRATION: Cosson et al. (1996: 370) reported that E. helvum is not a year-round inhabitant of Mauritania, and is likely a migrant species. Lelant and Chenaval (2011d: 14) reported an impressive cloud consisting of thousands of animals in Nouakchott on 9 November 2010. Hayman et al. (2010) who radio tracked an individual (EBOV-seropositive, see viruses below), showed typical migratory movements in this region of West Africa. The drivers, routes and patterns of migration are unknown (Peel et al., 2010a). It is unclear whether E. helvum has a panmictic population structure across a continous distribution that shifts according to seasonal resource availability, or whether a finer substructure and specific migration routes exist (Peel et al., 2010a). Daïnou et al. (2010) report that the distribution of Milicia excelsa (Moracea) is largely determined by the migration behaviour of the species main seed disperser E. helvum. Based on stable hydrogen isotopes in the fur of 88 examined museum specimens from Sub-Saharan Africa, Ossa et al. (2012: 1) report that 22 % had migrated at least 250 km, with a mean distance of 860 km, and a maximum of over 2,000 km (for seven specimens). For Zambia, Richter (2004: 70) suggests that trees of the genera Uapaca and Syzygium may be key in timing this species' migration, although she also indicates that the exact migratory cues for the species are unknown. Isotopic data captured by Ossa Gómez (2012: 22) suggest that E. helvum has a core distribution around the Gulf of Guinea, with migration towards the north (Mauritania, Niger) during the months of May to September and towards the south (Tanzania, Zimbabwe, Zambia) during the months of October and December. Pakula et al. (2017: 2) report on a Straw-coloured fruit bat that was observed on a ship sailing along the coast of West Africa, indicating that these bats can be transported over long distances. African Chiroptera Report 2020 DIET: Doutre and Sarrat (1973: 280) indicated that in West Africa, E. helvum has an outspoken preference for (over)ripe local fruits such as mangos, guavas, figs and bananas. They never saw them eat green fruits. Ayensu (1974) reports on E. helvum visiting the following plants when foraging: Parkia clappertonia (Dawadawa), Ceiba pentandra (SilkCotton tree or Kapop tree), Adansonia digitata (Baobab), Anacardium occidentale (Cashew tree), Ficus umbellata (Fig tree), Psidium guajava (Guava tree), Carica papaya (Papaya or pawpaw), and Azadirachta indica (Neem tree). Daïnou et al. (2010) mention that E. helvum feeds on the fruits of the Iroko trees (Milicia excelsa) in Cameroon, and suggest that the migratory behaviour of E. helvum is the primary mechanisim (seed dispersal) for the distribution of Milicia excelsa (Moraceae). In Accra (Ghana), Dechmann et al. (2010: 121) found that seven tracked bats mainly fed on introduced and/or cultivated fruits (primarily neem - Azadirachta indica, but also on papaya, mango, oil palm and banana). Only two animals ate wild figs. They also found that the core foraging area was only 3.9 ha (2.2 - 4.9 ha) and included only a few food trees. This area was on average 18.6 ± 11.8 km (max: 37 km) from their roost. Fahr et al. (2015: 1) found that E. helvum primarily fed on fruits of introduced trees during the wet season low population numbers, whereas nectar and pollen of native trees made up the major part of their diet during the dry season when population numbers peak. O'Brien (2011: 264) mentions that Syzygium is a staple food for mainland African E. helvum, but also refers to DeFrees and Wilson (1988: 4), who indicate that they are also fond of Ceiba pentandra. Investigations by Kankam and Oduro (2011: 22) in the Bia Biosphere Reserve in Ghana showed that Antiaris toxicaria [bark cloth tree, antiaris, false iroko, false mvule or upas tree] fruits processed by E. helvum still had some fruit pulp around them, and resulted in a far less degree of germination than seeds, which were processed by monkeys and were cleared of fuit pulp. In Congo, Kipalu (2009: 16) observed E. helvum visiting and/or consuming fruits of the African custard-apple (Annona senegalensis), Borassus palm (Borassus aethiopum), Heart Fruit (Hymenocardia acida), African peach (Nauclea 41 diderichii), African corkwood tree (Musanga cecropioides), and (Caloncoba welwitschii) Lavin et al. (2009: 336) report on the prevention of ISD (Iron Storage Disease of hemochromatosis) in captive E. helvum specimens, which can result in organ damage and possibly fatalities. The animals might succumb due to the extremely low iron concentrations in the wild diet as compared with the diet they have in captivity. In Abidjan, Côte d'Ivoire, Niamien et al. (2009: 235) reported the following fruits to be present in the bat's droppings: Anacardiaceae: Mangifera indica L. (Mango); Arecaceae: Elaeis guineensis Jacq. (African oil palm); Caricaceae: Carica papaya L. (Papaya); Mimosaceae: Parkia biglobosa (Jacq.) R.Br. ex G.Don (African locust bean) [flowers]; Moraceae: Ficus benjamina L. (Weeping fig, Benjamin fig), Ficus exasperata Vahl (Sandpaper tree, forest sandpaper fig, white fig, or sandpaper leaf tree), Ficus lutea Vahl (giant-leaved fig, Lagos rubbertree), Ficus polita Vahl (Heart-leaved fig), Ficus spp., Ficus sur Forssk (Cape fig, broom cluster fig), Ficus umbellata Vahl; Myrtaceae: Psidium guajava L. (common guava, yellow guava, lemon guava) as well as leaves of Araucaria excelsa (Norfolk Island Pine, Star Pine Araucariaceae). Niamien et al. (2017b: 187) added the fruits of Eugenia jambos L. (Rose apple, Malabar plum) and Eugenia malaccensis L (Malay apple) (Myrtaceaea) and leaves of Platycerium sp. (Polypodiaceae). In western Kenya, Webala et al. (2012: 12) reported that E. helvum helped to disperse seeds of over 32 plant species belonging to 17 families: Anacardiaceae: Mangifera indica (Mango); Apocynaceae: Saba comorensis (Bungo fruit); Canellaceae: Warburgia ugandensis (Uganda greenheart); Caricaceae: Carica papaya (Papaya); Cucurbitaceae: Momordica foetida; Flacourtiaceae: Flacourtia indica (Madagascar plum, batoka plum, flacourtia, governor's plum, Indian plum); Guttiferae: Garcinia buchananii (Granite Mangosteen); Lauraceae: Persea americana (Avocado); Malvaceae Ceiba pentandra (Kapok); Meliaceae: Melia azedarach (white cedar); Moraceae: Ficus amadiensis, Ficus asperifolia, Ficus lutea (Giant-leaved Fig), Ficus natalensis (Natal Fig), Ficus ovalifolia, Ficus ovata, Ficus sur (Cape Fig), Ficus sycomorus (Sycamore Fig), Ficus thonningii (Strangler Fig, Common Wild Fig, Bark-cloth Fig), Ficus vallis-choudae, Morus alba (White Mulberry); Myrtaceae: Psidium guajava (Apple or common guava), Syzygium cordatum (Water berry), Syzygium guineense (Water berry); Fabaceae: Crotolaria lanchnocarpoides; Rosaceae: Eriobotrya japonica (Loquat), Rubus apetalus; Rubiaceae: Vangueria 42 ISSN 1990-6471 apiculata (Mutugundo Fruit); Rutaceae: Teclea nobilis (Small fruited Teclea), Teclea trichocarpa (Furry-fruited Teclea), Toddalia asiatica (Orange climber). Webala et al. (2014: 90) removed Ceiba pentandra from the above list and added Allophylus ferrugineus (Sapindaceae). Makau (2016: xii) found it to be the most effective disperser of guava seeds in western Kenya. Seltzer et al. (2013: table S2) provides an overview of the plant species found beneath bat feeding roosts. For E. helvum these include Elaeis guineensis Jacq. (African Oil palm - Arecaceae), Magnistipula butayei De Wild (Chrysobalanaceae), Parinari excelsa Sabine (Guinea Plum - Chrysobalanaceae), Antiaris toxicaria Lesch. (upas tree - Moraceae), Ficus natalensis Hochst (Natal Fig - Moraceae), Milicia excelsa (Welw.) C. C. Berg (African Iroko Moraceae), Psidium guajava L. (apple guava Myrtaceae), Syzygium guineense (Willd.) D.C. ("water-berry", Snake bean tree - Myrtaceae) and Maesopsis eminii Engl. (Umbrella tree Rhamnaceae). Corlett (2017: 13) (referring to Abedi-Lartey et al., 2016) reports that gut passage times for small seeds in caged bats had a median of 72 min and a maximum of 19 h. Combined with the long commutes these bats undertake, this results in a potential dispersal distance of 10 to 100 km. However, they also indicate that the estimated median dispersal distances (for larger seeds) is about 50 m. van Toor et al. (2019: R238) found that the bats dispersed seeds over distances of up to 95 km, with averages ranging from 12.6 to 21.4 km, depending on colony and season. For the colony in Accra (Ghana), they calculated that during the dry season the 152,000 bats accounted for 338,000 dispersal events during a single night and produced an average seed rain of 26.2 dispersal events per km 2. During the wet season, when only 4,000 bats are present, there are only 5,500 dispersal events per night. In Senegal, Sow et al. (2020: 4) used molecular analyses to identify the presence of Heliocheilus albipunctella de Joannis) (Lepidoptera, Noctuidae) in the faeces of E. helvum foraging in millet fields (Pennisetum glaucum (L.) R. Br. (Cyperales, Poaceae)). PREDATORS: Mikula et al. (2016: 162) report that the Amur falcon (Falco amurensis Radde, 1863) and the Eurasian hobby (Falco subbuteo Linnaeus, 1758) were observed chasing Eidolon helvum bats in Kasanka forest, Zambia. Furthermore (supplemental data), they mention the following avian predators: Steppe eagle (Aquila nipalensis (Hodgson, 1833)), African hawk eagle (Aquila spilogaster (Bonaparte, 1850)), Common buzzard (Buteo buteo (Linnaeus, 1758)), Greater spotted eagle (Clanga clanga (Pallas, 1811)), Lesser spotted eagle (Clanga pomarina (Brehm, 1831)), Lanner falcon (Falco biarmicus Temminck, 1825) [with "Eidolon sp" from countries on the African continent as prey], Palm-nut vulture (Gypohierax angolensis (Gmelin, 1788)), White-backed vulture (Gyps africanus Salvadori, 1865), African fish eagle (Haliaeetus vocifer (Daudin, 1800)), Ayres's hawk-eagle (Hieraaetus ayresii Gurney, 1862), Bat hawk (Macheiramphus alcinus Bonaparte, 1850), Black kite (Milvus migrans (Boddaert, 1783)), Martial eagle (Polemaetus bellicosus (Daudin, 1800)), Crowned eagle (Stephanoaetus coronatus (Linnaeus, 1766)), Pied crow (Corvus albus Statius Muller, 1776). POPULATION: Structure and Density In general this is a common species forming large colonies of thousands to even millions of individual. Hayman et al. (2011b: 1; 2012c: 1393) report on a colony of up to one million individuals being present in Accra, Ghana for approximately 6 months during the dry season, and Sørensen and Halberg (2001: 213) report 1.5 million fruit bats leaving the roost at sunset in the Kasanga National Park, Zambia. Racey (2004a: 2) indicates that the number bats at the same locality peak near the end of November to about 8 million (representing a total mass of 3,000 metric tons in one hectare of forest and consuming nearly 6,000 tons of fruit each night), after which they begin to fall rapidly. Within colonies they form tight clusters of up to 100 animals, although in particularly large colonies this clustering may not be so obvious. Colonies may show extreme roost-site fidelity. During migration this species disperses into small groups. There is evidence of a widespread decline (P. Racey pers. comm. in Mickleburgh et al., 2008dc). A wellknown colony in Kampala (Uganda) declined in numbers over a forty-year period from ca. 250,000 animals to 40,000 in 2007 (Monadjem et al., 2010d). A trend which continued over the following years (Akite, 2009: 28-29). Based on emergence data for one roost on Principe (10,500 - 14,200 bats), Dallimer et al. (2006: 50) estimated a population density between 82 and 111 bats/km 2 for the entire island. On Annobón, Peel et al. (2012: 3) reported colony sized of up to 1,500 animals. Wood et al. (2012: 2885) indicate that while colonies in West Africa can be very large, these in African Chiroptera Report 2020 East Africa appear to be smaller and more fragmented, and also seem to be less migratory. Magloire et al. (2017: 890) reported on a colony from Abidjan (Côte d'Ivoire) counting on average 180,000 bats. They found that the colony size increased from August to November (from 200,000 to 300,000), stabilized from November to February and decreased from March to July (to 12,000). These fluctuations were linked to food diversity and abundance. For their study on the distribution of Ebola viruses, Fiorillo et al. (2018: 8) estimated the density of a E. helvum colony to be about 180 bats/km 2. On a much larger timescale, Chattopadhyay et al. (2019: 7) suggested that E. helvum would have taken advantage of the decreased aridity from the Last Interglacial (approximately 110 000 to 130 000 years ago) to the Last Glacial Maximum (approximately 20 000 years ago), which led to an increased population size. They also estimated the bat's suitable habitat percentage to 100 at LGM, 27.34 at LIG, 36.67 at Mid-Holocene (ca. 6 000 years ago) and currently 44.31. Trend 2008: Decreasing (Mickleburgh et al., 2008dc; IUCN, 2009). 2004: Decreasing (Mickleburgh et al., 2004bh; IUCN, 2004). LIFESPAN: Hayman et al. (2012c: 1399) examined the tooth cementum annuli, and found three old specimens with an estimated age range from 13 to 15 years. They found that adult bats had a survival probability of about 0.83, whereas this was 0.43 for juveniles. Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 21.8 years. Hayman and Peel (2016:134) calculated that the mean life expectancy estimates across the colonies they studied to be ranging from 2.3 to 6.8 years, and that individuals may live up to 30 years of age. Peel et al. (2017: 85) report on regional differences in the livespan of E. helvum: Tanzania (Dar es Salaam: mean 6.4 yrs, max 15 yrs, n = 53 adults; Morogoro: 4.5, 8, 24), Ghana (6.0, 14, 76), Bioko (2.4, 6, 16), Príncipé (4.2, 10, 23), and São Tomé (4.8, 13, 57). ACTIVITY AND BEHAVIOUR: In Nigeria, E. helvum was reported to fly at least 24 km each night and often stopped at a number of locations between leaving the roost and arriving 43 at the foraging site (Okon, 1974). Richter and Cumming (2006) reported flights of over 15 km for E. helvum in Kasanka National Park, Zambia. Richter and Cumming (2008: 172) reported that bats foraged as far as 59 km away from their roost. During their migration, one animal travelled 370 km in one night, but on average they travelled 29 km/day. The largest distance travelled by one bat, reported in that study, is 2,518 km in 149 days. Sapir et al. (2014: 1) found that, in Ghana, E. helvum modified its flight in response to the wind vector (e.g. they decreased their airspeed in response to tailwinds so that their groundspeed remained nearly constant). In Ghana too, Fahr et al. (2015: 1) found that E. helvum foraged locally during the wet season population low (ca. 4,000 animals traveling between 3.5 and 36.7 km) in urban areas with low tree cover. During the dry season population peak, some 150,000 individuals travelled between 24.1 and 87.9 km. Carpenter (1986: 90) and McGuire and Guglielmo (2009: 1291) reported that the respiratory quotient for this species indicates that fat provided 77 - 92 % of the fuel during flight, and a decline in respiratory quotient during long flights may prove that fat is indeed the only fuel source following initial carbohydrate use. Carpenter (1986: 93) also indicates that at airspeeds of 5.5 to 6.5 m/s the heart beats at a frequency of 576 - 601 beats per minute, and at airspeeds between 5.5 and 7.0 m/s, these bats take 295 to 316 breadths per minute. From flight corridor tests, Riskin et al. (2012: 2946) determined the average flight speed to be 4.24 ± 0.91 m/s. In captive groups, Carter and Leffer (2015: 3) found that 6 males spent 0.52 % of their awake time with social grooming. A group consisting of six females and two castrated males did not perform any social grooming at all. REPRODUCTION AND ONTOGENY: For Uganda, Mutere (1967, 1968) reported pregnancies between October and February and births occurring between February and March. For Congo, Bergmans (1979a: 165) reported two pregnant females on 6 December 1972. Krutzsch (2000: 109) reports that various authors found that E. helvum has a monoestrous reproduction cycle. Simbauni and Bernard (2001: 63) indicate that the reproductive cycle of E. helvum is characterized by a three-month period of delayed implantation (see 44 ISSN 1990-6471 also Badwaik and Rasweiler, 2000: 228; Racey and Entwistle, 2000: 383) and that the pregnancy may last for 10 months. At birth, the young has about 18.4 % of the mother's body mass (48.6 g versus 263.5 g; see Kurta and Kunz, 1987: 82). Peel et al. (2013b: 7) indicate that E. helvum is panmictic across its continental African range, meaning that all individuals from over the entire range are potential partners. They found no indication that gene flow was more likely to occur among neighbouring populations than among distant populations (> 4,500 km). They conclude that E. helvum, therefore, has the largest reported panmictic unit of any mammal. Szekely et al. (2015: Suppl.) report a gestation period of 275 days, and indicate that weaning occurs after 65 days. At birth the young weighs 50 g and after gestation its weight has increased to 135 g. In their overview, Peel et al. (2017: 82) report the estimated start of births in different countries: Annobón (mid-September), Malawi (midNovember), Principé (November-December), Tanzania (mid-December), São Tomé and Ghana (February-March), Uganda (March-April), Bioko (mid-April). They also indicate (p. 83) that females are reproductively active between the ages of 2 and 14 years. PARASITES: BACTERIA Gram-negative bacterium - Pinus and Müller (1980) and Sara (2002: 41) reported Citrobacter freudii, Escherichia coli, Klebsiella oxytoca, and Klebsiella pneumoniae. Nowak et al. (2017: 6) tested three bats from the Republic of Congo of which two tested positive for E. coli. Gram-positive bacterium - Akobi et al. (2012: 1) found that 500 faecal samples of bats at the university of Ife-Ife, Nigeria were colonized by 107 isolates of Staphylococcus aureus strains with genotypic characteristics that are rarely found in humans. Held et al. (2016: 118) refer to a report from Nigeria, indicating that E. helvum can be colonized by S. schweitzeri. Spirochaetales - Reviewed in McCoy (1974), who listed Borrelia sp. Ogawa et al. (2015a: 143) report on the presence of at least two other spirochaete bacteria: Leptospira borgpetersenii and L. kirschneri in E. helvum which migrated from the Democratic Republic of the Congo to Zambia. Bartonellae - Bacteria Bartonella - Kosoy et al. (2010: 1877) and Bai and Kosoy (2012: 58) reported a prelevence of Bartonella spp. In E. helvum from Kenya of 23/88 (26.1%) cultured from blood samples. The level of gltA sequence identified between 16 Bartonella genotypes, but varied greatly, eight genotypes (sequence variants) were clustered around 4 clades (Kosoy et al., 2010: 1878). Three novel Bartonella species have been isolated from E. helvum (Kosoy, 2010: 719). See also Kosoy (2010: 719) for further information. Morse et al. (2012: 1719) refer to Billeter et al. (2012) for Bartonella sp. found in the bat fly Cyclopodia greefi, parasiting on E. helvum in Ghana. Billeter et al. (2012: 325) collected batflies (Cyclopodia greefi greefi) from E. helvum specimens from Ghana (see also Morse et al., 2012), Bioko, and Annobón. Of these bat flies, 56.5 % (Ghana) to over 71 % (Bioko, Annobón) were carriers of Bartonella bacteria. Genetically, 65.9 % of the Bartonella sequences were identical or similar to sequences obtained in other studies (e.g. Kosoy et al., 2010). One sequence was very similar to one obtained from a E. helvum specimen collected in Kenya (Billeter et al., 2012: 326 - 327). Kamani et al. (2014: 628) tested 79 E. helvum specimens from Nigeria and found at least 44 testing positive for Bartonella DNA. 79 isolates studied by Bai et al. (2015: 7) revealed five genogroups (E1 to E5) for ribC. An additional lineage (Ew) did not contain a fragment for ribC. Mannerings et al. (2016: 923) report that the following Bartonella species were already isolated from E. helvum bats: B. henselae, B. quintana, B. clarridgeiae, B. vinsonii vinsonii, B. elizabethae, and Bartonella strain E1–105. Olatimehin et al. (2018: 3) found Staphylococcus aureus, Staphylococcus schweitzeri and Staphylococcus argenteus in feacal samples from bats at the Obafemi Awolowo University, Ile-Ife, Nigeria. Di Cataldo et al. (2020: 2) examined 51 Nigerian Eidolon bats and found 13 (45 %) to be infected by hemoplasma bacteria. HAEMOSPORIDIA Boundenga et al. (2018: 10581) reported 1 out of six examined E. helvum to be infected by Hepatocystis (16.67%). In a study performed in Ibadan (Nigeria), Li et al. (2019a: 21) found six out of 109 E. helvum specimens (5.5%) infected with Cryptosporidum sp. and 16 (14.7%) infected by Enterocytozoon bieneusi; none were infected by Giardia duodenalis. African Chiroptera Report 2020 ACARI Laelapidae: Taufflieb (1962: 112) indicated that Chelanyssus aethiopious Hirst, 1921 was found once on a female Eidolon helvum, but this was never confirmed by subsequent findings. He considered this one time occurrence as accidental. Sarcoptidae: Nycteridocoptes pteropodi Rodhain and Gedoelst, 1923 was reported by Fain (1958: 236) from E. helvum specimens from Astrida (=Butare), Rwanda. Spinturnicidae: Taufflieb (1962: 112) reported Meristaspis kenyaensis Radford, 1947. Demodicidae: Bianco et al. (2019: 547) reported the first case of demodicosis, caused by Demodex sp. mites from bats in a captive colony at the Accra Zoological Gardens (Ghana). HEMIPTERA Cimicidae: Haeselbarth et al. (1966: 8) recorded Afrocimex constrictus Ferris & Usinger, 1957 associated with E. helvum near Ruiru, Kenya. DIPTERA Nycteribiidae: Penicillidia fulvida (Bigot, 1885) (see Haeselbarth et al., 1966: 114). Eucampsipoda africanum Theodor, 1955, widely distributed over the Ethiopian region and recorded from localities in Senegal to the Sudan and southwards to the Cape (Haeselbarth et al., 1966: 115). Dipseliopoda setosa Theodor, 1955 from Kenya and Tanzania (Haeselbarth et al., 1966: 116). C. greeffi was originally described by Karsch (1884) from an E. helvum specimen from São Tomé, and the adjacent islet of Rojas (see Urich et al., 1922: 471). Urich et al. (1922) furthermore mentioned additional specimens from this bat fly, reported by Rodhain and Bequaert from E. helvum specimens from Leopoldville [= Kinshasa], DRC. Cyclopodia greeffi are found in a belt across central Africa, between lat. 10o south and north, while in West Africa it occurs further north up to lat. 20o (Haeselbarth et al., 1966: 117). Billeter et al. (2012: 325) collected bat flies (Cyclopodia greefi greefi) from E. helvum specimens from Ghana (see also Morse et al., 2012), Bioko, and Annobón (see above for bacteria results from this study). Szentiványi et al. (2019: Suppl.) indicated that these bat flies were also carrying the arthropod Eupelmus urichi (on São Tomé) and Bartonella bacteria (in Ghana). SIPHONAPTERA Ischnopsyllidae: Thaumapsylla breviceps Rothschild, 1907, described from Kenya and the DRC (Haeselbarth et al., 1966: 186). Thaumapsylla dina Jordan, 1937 from the mountains of the Ruwenzori, Congo and Kenya (Haeselbarth et al., 1966: 186). Lagaropsylla consularis Smit, 1957, traditional host being 45 molossid bats, but it has also been recorded from this taxon (Haeselbarth et al., 1966: 190). Teinocoptidae: Teinocoptes eidoloni Fain, 1959 was reported by Fain (1967: 367) from E. helvum from Astrida (=Butare), Rwanda. VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Eidolon helvum: Adenoviruses, Astroviruses, Coronaviruses, Flaviviruses, Paramyxoviruses and Polyomavirus. Willoughby et al. (2017: Suppl.) report the following viruses: Bundibugyo ebolavirus, Bunyamwera orthobunyavirus, Entebbe bat virus, Hendra henipavirus, Ife virus, Influenza A virus, Lagos bat lyssavirus, Mokola lyssavirus, Nipah henipavirus, Ntaya virus, Sudan ebolavirus, Tai forest ebolavirus, Usutu virus, West Caucasian bat lyssavirus, West Nile virus, Yellow fever virus, Zaire ebolavirus, Zika virus. Nieto-Rabiela et al. (2019: Suppl.) reported the following viruses: Achimota virus, Bat coronavirus, Bat kobuvirus, Bat mastadenovirus, Bat paramyxovirus, Bat pegivirus, Bat simplexvirus, Chiroptera tetraparvovirus, Eidolon helvum papillomavirus, Eidolon helvum retrovirus, Eidolon paramyxovirus, Ife virus, Kumasi rhabdovirus, Lagos bat virus, Polyomavirus, Poxvirus, Rotavirus A. Yinda et al. (2016: 6) identified 5 divergent novel bat RVA strains from faeces collected in Cameroon. Four of these were genetically similar to each other and the last one was related to the Kenyan bat strain. They did not notice any diarrhea or other obvious signs of sickness in the bats. Peel (2018: 1) estimated that maternally-derived immunity to Lagos bat virus and African henipavirus has a mean duration of about six months, which is much less than the acquired immunity for these viruses: 12 and 4 years respectively. Yinda et al. (2018: 2) pooled 87 fecal samples of Eidolon helvum and Epomophorus gambianus from Cameroon into 25 groups. One group containing fecal samples of E. gambianus only yielded mainly phage viral reads, leading them to conclude that all the eukaryotic viral reads they encountered were from E. helvum. The viruses they found belonged to the following families: Alphaflexiviridae, Astroviridae, Baculoviridae, Caliciviridae, Caulimoviridae, Circoviridae, Coronaviridae, Dicistroviridae, Geminiviridae, Hepeviridae, Herpesviridae, Iflaviridae, 46 ISSN 1990-6471 Marnaviridae, Nodaviridae, Other Picornavirales, Papillomaviridae, Partitiviridae, Parvoviridae, Picobirnaviridae, Picornaviridae, Polydnaviridae, Reoviridae, Retroviridae, Secoviridae, Tombusviridae, Totiviridae, Tymoviridae, Unclassified eukaryotic virus, and Virgaviridae. Most of the identified partitiviruses and densoviruses constitute putative novel genera in their respective families. In bats from Saudi Arabia, Mishra et al. (2019: 5) found evidence for the following viruses: Bat astrovirus, Bat papilloma virus, Bat Coronavirus, Bat polyomavirus, Me Hepe-Astro virus, Parechovirus, Rotavirus, Sapovirus, Teschovirus, Adenoviridae Sequences of Eidolon helvum adenovirus 1 attributed to the genus Mastadenovirus were found by Baker et al. (2013a: 98) in samples from bats from Ghana. Conrardy et al. (2014) tested 9 bats from Kenya, none of which tested positive. Ogawa et al. (2017: 2) isolated a new adenovirus from Zambian bats: E. helvum adenovirus 06-106 - EhAdV 06-106, which they also tentatively called: Bat mastadenovirus H. Caliciviridae Sapovirus (SaV) From Cameroonian bats, Yinda et al. (2017a: 2) identified the complete genome of six highly divergent SaVs and one partial SaV, which probably represent two novel genogroups, that are only distantly related to any other known SaVs. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, six individuals for antibody to SARS-CoV in sera from Bandundu Province, DRC, none were found to be positive. In February 2008, Pfefferle et al. (2009) tested 212 fecal samples from bats from Ghana and all were negative for coronavirus (CoV) RNA. Tong et al. (2009) found 6/10 bats collected in 2006 positive for coronavirus RNA in Kenya (Kenya bat coronavirus BtKY20, Kenya bat coronavirus BtKY23, Kenya bat coronavirus BtKY24). 21 % (38 out of 181) of the Kenyan bats tested by Tao et al. (2017: 3) were positive for coronaviruses. In Nigeria during June 2008, Quan et al. (2010) screened the gastrointestinal tissue for the presence of coronaviruses by PCRs, none tested positive. Anthony et al. (2017b: Suppl.) mention betacoronaviruses Eidolon_bat_CoV and Kenya_CoV_BtKY56. Leopardi et al. (2016: 575) report on the first occurrence of CoV outside of Kenya: Nigeria. Nziza et al. (2019: 156) found Eidolon bat coronavirus/Kenya/KY24/2006 in a rectal swap from Rwandan bats. Joffrin et al. (2020: 7) identified the Kenyan viruses as Beta-D coronavirus. Filoviridae - Filoviruses Olival and Hayman (2014: 1770) found that E. helvum has a high seroprevalence of antibodies against several RNA viruses, but not filoviruses, especially when compared to other species in the same locations. They suggest that the migratory lifestyle of this species might prevent filovirus cirulation. Peterson et al. (2007: 1547) indicate that the geographic distribution of E. helvum covers the entire distribution range of both Ebola and Marburg viruses. Hayman (2015: 6) points to the single, highly synchronous birth pulse per year, which prevents persistence of the virus in E. helvum. Ebolavirus Hayman et al. (2010) tested the blood of 262 individuals from Ghana for Ebola virus using indirect flurescent tests for antibodies against EBOV subtype Zaire [= ZEBOV], one positive was found in a pregnant female. The positive reactivity for EBOV was confirmed using western blot against a recombinant nucleocapsid protein of EBOV-Zaire (Hayman et al., 2010). Funk and Piot (2014: 2) erroneously refer to Gire et al. (2014), who should have claimed that the 2014 Ebola outbreak in West Africa probably originated from a child infected by a strawcoloured fruit bat. However, as indicated by Jakob Fahr in the discussion following this paper, Gire et al (2014) never mention bats. The reference was then changed to Baize et al. (2014), but that paper (p. 1424) only refers to the "usual suspects": Hypsignatus monstrosus, Epomops franqueti and Myonycteris torquata. To date, there is no proof that the West African Ebola outbreak is directly related to bats. In Zambia, Ogawa et al. (2015b: "1") found some serum samples that showed distinct specificity for Reston ebolavirus, which has thus far been found only in Asia. Studying E. helvum genes, Ng et al. (2015: 2) found that the NCP1 gene, which encodes a protein that mammals need in order to move cholesterol within their cells, also turns out to be a receptor that the filoviruses must bind to before they can infect the cells. Marburgvirus Towner et al. (2007) tested 35 individuals from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and 33 individuals for antiMarburg virus IgG antibodies by ELISA; no positives were found. Hayman et al. (2010) African Chiroptera Report 2020 tested the blood of 262 individuals from Ghana for Marburg virus using indirect flurescent tests for antibodies against MARV subtype Leiden, no positives were found. Flaviviridae Flavivirus - In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against West Nile virus (WNV) and non-specific Flaviviruses. Malmlov et al. (2019: 3) refer to Simpson et al. (1968) who experimentally inoculated three E. helvum specimens with Zika virus, of which two became viremic and seroconverted. Pegivirus (BPgv) - 0 out of 8 bats from Cameroon, 1 out of 17 from the Democratic Republic of the Congo, 0 out of 11 from Kenya and 1 out of 50 animals from Nigeria examined by Quan et al. (2013: Table S5) were infected by clade H type Pegivirus (overal 2.3 %). For the same bats examined, they also found 1, 0, 1, and 5 specimens infected by clade K type Pegivirus. Herpesviridae Alphaherpesvirinae Simplexvirus Dak An Y 7 – (3 isolates) Cameroon, 1971, isolated from pooled organs (Razafindratsimandresy et al., 2009). Betaherpesvirinae Eidolon helvum herpesvirus 1 and 3 - Baker et al. (2013a: 96) identified 539 sequences related to herpesviruses, primarily from throat samples: 366 belonging to betaherpesvirinae. Gammaherpesvirinae Eidolon helvum herpesvirus 2 - Baker et al. (2013a: 96) identified 539 sequences related to herpesviruses, primarily from throat samples: 171 to gammaherpesvirinae. Nairoviridae Orthonairovirus Of the 167 specimens from Ghana tested by Müller et al. (2016: 3), 1 was positive for Crimean Congo hemorrhagic fever virus (CCHFV) Orthomyxoviridae Freidl et al. (2015) conducted serological studies on 100 E. helvum serum samples collected in Ghana in 2009/10. About 30% of the screened animals showed serological evidence to avian influenza subtype H9. According to Brunotte et al. (2016: 117), this suggests that bats can be naturally infected with classical IAV, although they did not find any evidence to support this hypothesis. 47 Papillomaviridae (PV) Baker et al. (2013a: 97) found 408 sequences belonging to multiple papillomaviruses, of which two represented new forms. García-Pérez et al. (2013: 51) cloned and sequenced the complete genome of a novel PV, EhelPV1, isolated from hair bulbs from a captive straw-colored fruit bat. Mengual-Chuliá et al. (2012) reported on PV sequences from hair follicles of captive bats. Paramyxoviridae - Paramyxoviruses Nine individuals from Kenya were tested for paramyxoviruses by Conrardy et al. (2014). All nine were negative for paramyxovirus RNA. Paramyxovirinae Henipavirus Hayman et al. (2008b) detected antibodies to both Nipah (NiV - 39 %) and Hendra (HeV - 22 %) [both members of the genus Henipavirus (see Hayman et al. (2011b: 1)] in bats collected in Ghana in 2007. Peel et al. (2012: 2) found antibodies against these viruses in bats on Annobón. Drexler et al. (2009) detected henipavirus RNA and three novel viruses were identified. Drexler et al. (2012a: Suppl. Table 1) indicated that 42 out of 722 (5.8 %) bats collected in Ghana, the Central African Republic and Gabon tested positive for Henipavirus. 4.0 % (29) tested positive for Rubulavirus and 1.8 % (13) for Pneumovirus. Pernet et al. (2014: 3) screened 44 Cameroonian bat serum samples for anti-NiV cross-neutralizing activity and found 21 (ca. 48 %) testing positive. Pararubulavirus Baker et al. (2012: 1348) describe two new Rubulaviruses: Achimota virus 1 (AchPV1) and Achimota virus 2 (AchPV2). They developed specific serological assays for these two viruses and found evidence of infection by both in E. helvum across sub-Saharan Africa and on islands in the Gulf of Guinea. Baker et al. (2011b: 855) indicate that a coevolutionary relationship might exist between Chiroptera and Paramyxoviridae, based on an analysis of urine samples of E. helvum from Accra, Ghana. Weiss et al. (2012a: suppl. 1) report on Paramyxovirinae sequences obtained from liver and urine samples from bats collected near Brazzaville (Republic of Congo), organs of one individual presented with two diverse paramyxovirus sequences. From Zambia, Muleya et al. (2013: 611) report five new paramyxoviruses closely related to the genus Henipavirus and two related to the unclassified Bat paramyxoviruses from Ghana and Congo Brazzaville. 48 ISSN 1990-6471 With the use of a seroneutralization assay, Pernet et al. (2014) found cross-neutralizing antibodies to Nipah virus in 48 % of sampled E. helvum bats collected in Cameroon. Mbu'u et al. (2019: "1") mention the presence of Ghanaian bat henipavirus (designated Ghana virus - GhV) in Ghana. Glennon et al. (2019: 1) followed a captive colony in Ghana for nine years and found that the observed dynamics of the henipavirus serology could only be explained by reinfection of the bats. They also found that the acute infectious period is extremely short (hours to days), and that immunity can last for about one to two years. Orthorubulavirus Achimota virus: Baker et al. (2012) describes two new paramyxoviruses isolated from E. helvum: Achimota virus 1 (AchPV1) and Achimota virus 2 (AchPV2). Several rubulaviruses and rubula-related viruses were reported in several E. helvum individuals sampled from several African countries (Drexler et al., 2012). Some of these sequences proved to be closely-related to human mumps virus and simian virus 5. Kohl et al. (2018: 11) indicate that a number of yet unknown rubulaviruses might be present in these bats. Markotter et al. (2020: 6) mentions Mumps virusrelated viruses occurring in Eidolon helvum. Pneumovirinae Pneumovirus Several individuals sampled in Ghana tested positive for a pneumovirus closely related to pneumonia virus of mice, and grouped in a sister clade to human respiratory syncytial virus. contig from a lung sample, and related to the genus Enterovirus, were reported by Baker et al. (2013a: 100). In the faeces of Cameroonian bats, Yinda et al. (2017b: 3) identified 11 new picorna-like viruses of which some might represent new families [*] within the order Picornavirales: bat sapelovirus, bat kunsagivirus and bat crohivirus, bat iflavirus [Iflaviridae], bat posalivirus [Posa-like virus; *], bat fisalivirus [Fisa-like virus; *], bat cripavirus [Dicistroviridae], bat felisavirus [fesa-like virus; *], bat dicibavirus [bat dicistrovirus; *] and bat badiciviruses 1 and 2 [anagram from bat dicistrovirus; *]. Zeghbib et al. (2019: Suppl.) reported Hepatovirus from a bat from Ghana, and Crohivirus, Kunsagivirus and an unassigned (Sapelo-like) virus from Cameroon. Polyomaviridae Baker et al. (2013a: 100) assembled a sequence from a throat sample. Eckerle et al. (2014: 4) mention the presence of several, unassigned viruses of this family. Conrardy et al. (2014: 259) reported that one of the nine specimens tested in the Vihiga district (Kenya) tested positive for Polyomavirus. Tao et al. (2012) identified two novel species termed Eidolon PyV1 and 2, from Kenya in 2006. Poxviridae 38 contigs related to Molluscum contagiosum [MC]), a human contagion were reported by Baker et al. (2013a: 99). Baker and Murcia (2014: 1567) indicate that the bats do not show any clinical signs. Peribunyaviridae Orthobunyavirus Fagre and Kading (2019: 4) report that Bunyamwera virus (BUNV) was isolated from this bat or that molecular and serologic evidence of it was found. Reoviridae - Rotaviruses Rotavirus Esona et al. (2010: 1844) reported genetic characterization of a bat rotavirus (Bat/KE4852/07) detected in the feces of E. helvum, which is a novel Rotavirus A species. This isolate contained a VP4 gene that probably originated from a human rotavirus, an NSP4 gene of likely human or animal rotavirus origin, and otherwise a unique genetic background requiring establishment of new genotypes for at least 7 genes (Esona et al., 2010: 1845). Sasaki et al. (2018: 104) compared genome segments of RVA from bats from Cameroon and Zambia and found these to be 97 to 99 % identical, suggesting that E. helvum is capable of dissimating RVA across long distances. Picornaviridae Eidolon-helvum-Picornavirus-1 contigs related to the genus Kobuvirus from 6 urine samples and one Orbivirus Ife virus - In Nigeria 1971, the virus was isolated from the salivary glands, brain and blood (Kemp et Parvoviridae Canuti et al. (2011: 3) report a new virus (Eidolonhelvum-bat-Parvovirus-1 [Eh-BtPV-1]) from bats captured in Kumasi, Ghana. Baker et al. (2013a: 100) found 10 sequences, belonging to both the mammalian-infecting Parvovirinae (related to the genera Erythrovirus and Betaparvovirus) and the invertebrate-infecting Densovirinae. African Chiroptera Report 2020 al., 1988). Also reported by de Jong et al. (2011: 11). Fagre et al. (2019: 2) indicated that this virus was also found in bats from Cameroon and the Central African Republic. Retroviridae Baker et al. (2013a: 100) reported on 292 sequences related to retroviridae, that were recovered from lung samples. Rhabdoviridae No rhabdoviruses were detected from 9 E. helvum bats sampled in Kenya Conrardy et al. (2014). Lyssavirus - Rabies related viruses 2006 in Nigeria, Dzikwi et al. (2010: 269) tested 223 brains by direct fluorescent antibody (DFA) and mouse inoculation test (MIT) which tested negative for lyssavirus antigens. In Nigeria during June 2008, Quan et al. (2010) did not detect any lyssavirus-specific antigens from the brains by use of direct fluroescent antibody testing. Lagos bat virus (LBV) - Isolated in Nigeria (Boulger and Porterfield, 1958: 424 [1st detection]; Shope et al., 1970; Kemp, 1975: 616; Hayman et al., 2011a: 88; Fisher et al., 2018: 242) and Senegal (Institute Pasteur, 1985; Hayman et al., 2011a: 88). Antibodies detected in Ghana (Hayman et al., 2008a); Kenya (Kuzmin et al., 2008a; Kuzmin et al., 2011b: 1467; Nadin-Davis et al., 2011: 238), Annobón (Peel et al., 2012: 3). In 2006, 105 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), where 24 tested positive (22.86 %) for neutralizing antibodies (Dzikwi et al., 2010: 269). Hayman et al. (2010), who found an individual which tested positive for EBOV, also tested positive for LBV, but no antibodies against Mokola (MOKV). Restif et al. (2012: 1091) report on how emerging infectious disease research can be undertaken when little prior knowledge exists, based on the occurrence of LBV in a population of E. helvum in Accra, Ghana. Freuling et al. (2015: 44) point out that Lagos bat viruses belong to two phylogenetic lineages and that the presence of neutralizing antibodies might be missed if the test is performed with the most commonly used isolate. Kalemba et al. (2017: 409) reported that out of 18 bats examined from the DRC, six tested positive for lyssavirus neutralizing antibodies (LBV). Hayman et al. (2018: 1) constructed a model to determine how host traits (e.g. maternally-derived antibodies, seasonal birthing) and viral traits (e.g. incubation periods) interact to allow LBV viruses to persist within bat populations. 49 Mokola virus (MOKV) - King et al. (1990 168) found monoclonal antibodies for MOKV in bats from Nigeria and Senegal. In 2006, 105 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), where 2 tested positive (1.9 %) for neutralizing antibodies (Dzikwi et al., 2010: 269). Shimoni bat virus (SHIBV) - Kuzmin et al. (2011b: 1467) found 2 individuals testing seropositive on 9 animals tested between 2009 and 2010 in Kenya. Kalemba et al. (2017: 409) found four out of 19 bats examined from the DRC tested positive for SHIBV neutralizing antibodies. Horton et al. (2014: Table S1) tested 8 Kenyan E. helvum specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). Kumasi Rhabdovirus (KRV) - This new virus was detected by Binger et al. (2015: 4588) in the spleen and sera of E. helvum specimens from Kumasi in Ghana. By means of Capture-Mark-Recapture (CMR) procedures, Hayman et al. (2012b: 2163) found that survival probabilities for lyssavirus seronegative and seropositive Eidolon helvum in Ghana were not significantly different. King et al. (1990: 168) reported monoclonal antibodies for Duvenhage and Denmark bat virus from bats from Nigeria and Senegal. ???: Kemp (1975: 616 - 617) also mentions IbAn 67204 being isolated on 9 April 1971 from this species in Nigeria, and Herpes (YV6) virus in April 1971 and Dakar YV 177 in May 1971, both latter in specimens from Cameroon. In their overview table, Maganga et al. (2014a: 8) report the following viruses were found on E. helvum: Lagos bat virus (LBV), Mokola virus, West Caucasian (WC) virus, Zaire Ebola virus (ZEBOV), Ife virus (Orbivirus), Hendra virus, Nipah virus (NPHV), Rubulavirus, Coronavirus, Rotavirus related, Simplexvirus, Parvovirus. Peel et al. (2016) examined 2,827 bats from 9 countries over 8 years and focused on four viruses: Lagos bat virus (LBV), African henipaviruses, Achimota virus 1 (AchPV1) and Achimota virus 2 (AchPV2). The genomic mining study by Cui and Wang (2015: 5793) revealed bornaviral elements built in the genome of E. helvum 50 ISSN 1990-6471 UTILISATION: In Ghana E. helvum is a food source (Hayman et al., 2008a). Eidolon helvum is the most heavily harvested bat for bushmeat in West and Central Africa, and this is believed to be a major factor in reported population declines (P. Racey pers. comm. in Mickleburgh et al., 2008dc). Kamins et al. (2014) investigated the sociological characteristics of communities who interact with bats (specifically E. helvum in Ghana) to assess their attitudes and perceptions regarding bats and disease. Ohemeng et al. (2017: 184) indicated that local (Ghanese) people prefered eating E. helvum over other bat species, because of their delicious taste. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Madagascar, Malawi, Mauritania, Mozambique, Namibia, Niger, Nigeria, Rwanda, São Tomé and Principé, Senegal, Sierra Leone, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. In Benin, Dougnon et al. (2012: 85 - 86) found an average carcass weight of 89.27 ± 11.41 g (about 43 % of the body mass). The quality of the meat was evaluated (for 100g meat): Total mineral matter: 13.2 g, Lipids: 23.11 g, Protein: 56.8 g, Calcium: 55 mg, Sodium: 39 mg, Phosphore: 83 mg and Potassium: 75 mg, making it more fatty than E. gambianus meat (Lipids: 13.36 g). The authors suggest that the higher fat content is a result of the longer foraging distance covered by E. helvum. See also threats section for additional information on utilization. Figure 6. Distribution of Eidolon helvum Eidolon helvum helvum (Kerr, 1792) *1792. Vespertilio vampyrus helvus Kerr, in: Linnaeus, Animal Kingdom, 1 (1): xvii, 91. Publication date: February 1792. Type locality: Senegal: "Senegal". - Comments: Type locality fixed by Andersen (1907a: 504); see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1), Harrison and Bates (1991). Andersen (1912b: viii) indicates that the type was once in the Museum Leverianum, but is probably no longer in existence. - Etymology: From the Latin "helvus" (=dun-coloured), referring to the straw colour on their shoulders; the back, rump and hind limbs are normally various shades of brown (see DeFrees and Wilson, 1988: 4; Taylor, 2005). Lanza et al. (2015: 27) translate "helvus" to "dull-yellow". 1803. Pteropus stramineus E. Geoffroy Saint-Hilaire, Catalogue des Mammifères du Muséum national d'Histoire naturelle, Paris, 48. Type locality: Sudan: Sennaar [15 35 N 33 38 E, 425 m] [Goto Description]. - Comments: Harrison (1964a: 44) refers to Sherborn to indicate that the 1803 paper has never been published. The next available date for this name is 1810: Ann. Mus. Nat. Hist., Paris, 15: 95. Andersen (1912b: 12) mentions that the description in Geoffroy's 1813 paper was based on three specimens in the Paris Museum (nos 92 and 93 from an unknown locality, presented by Professor Fourcroy, and no 94 "de la collection du Stathouder") but these could not be traced by him. The description in the 1810 paper was based on two specimens, one believed to be from Timor (Péron and Lesueur), and the other without details (and possibly one of the cotypes of the 1803 description). Andersen (1912b: 12) also pointed out that "The 'Cat. Mamm. Mus. Nation. d'Hist. Nat.' (1803) was suppressed by Geoffroy himself, and the name Pteropus stramineus is therefore usually dated from his well-known paper in Ann. Mus. d'Hist. Nat. XV. (1810)." Originally, the type locality was not specified, but it was subsequently fixed as Senaar by Koopman (1975: 357): see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1). Harrison (1964a: 44) indicates that the type locality has been fixed by Temminck (1837b, Mon. Mamm. 2: 84), and Andersen (1912b: 12) mentions that the original type locality was African Chiroptera Report 2020 1809. 1815. 1862. 1862. 1865. 1866. 1881. 1906. 1906. 1953. 1975. 2000. ? 51 "Timor" (based on the 1810 description), but that Temminck already corrected this to "Africa". Vespertilio caninus var. b Goldfuss, Vergl. Naturbeschr., Säugethiere, 98. - Comments: Preoccupied by V. caninus Blumenbach, 1797: see Meester et al. (1986: 28). Pteropus flavus Illiger, Abh. Ak. Berlin, 90, 98. - Comments: Nomen nudum (see Andersen, 1912b: 809). Pterocyon paleaceus Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 1861: 423. Type locality: Sudan: Sennaar [15 35 N 33 38 E, 425 m]. - Comments: Type locality originally as "Africa", but restricted to Senaar by Matschie (1899b: 62): see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1). Pteropus mollipilosus H. Allen, Proc. Acad. nat. Sci. Philad., 13 (11): 159 (for 1861). Type locality: Gabon: "Gabon" [Goto Description]. - Comments: Originally mentioned as "Western Africa" by Allen (1861: 160), but restricted to Gabon (see Andersen, 1912b: 12). The type specimen was collected by Mr. Duchaillu, and is presumably in the Philadelphia Museum (see Andersen, 1912b: 12). Pteropus palmarum Heuglin, Leopoldina (Halle), 5 (3 & 4): 34. Publication date: June 1865. Type locality: Sudan: Sennaar [15 35 N 33 38 E, 425 m]. - Comments: Type locality originally as "Middle and Upper White Nile and between Senaar and Fazogli along the Blue Nile", but subsequently fixed as Senaar by Koopman (1975: 360): see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1). Andersen (1912b: 12) mentions that the type is probably no longer in existence. Xantharpyia leucomelas Fitzinger, in: Heuglin and Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 54 (1) 10: 544. Type locality: Sudan: Kordofan province: Bahr-el-Azrak, Bahr-el-Abiad: Sennaar [15 35 N 33 38 E, 425 m]. - Comments: Type locality originally as "Upper Nile lands", but subsequently fixed to Senaar by Koopman (1975): see Meester et al. (1986: 28), DeFrees and Wilson (1988: 1). Andersen (1912b: 13) mentions that the specimen on which Fitzinger based himself is probably in the Munich Museum, and that a specimen belonging to the same series as the type is in the BM (1847.5.27.28; ad ♀; SK; purchased Parreys, from Sennaar). Leiponyx büttikoferi Jentink, Notes Leyden Mus., 3: 59, 60. Publication date: April 1881. Type locality: Liberia: St. Paul's river: Millsburg [06 33 N 10 38 W] [Goto Description]. Comments: Type in RMNH. R[oussettus (Xantharpyia)] buettikoferi: Johnston, Liberia, Volume II: 690. - Comments: Bakwo Fils (2012: 690 - footnote) indicates that this name refers to the "very doubtfull Leiponyx buettikoferi" from Jentink. (Name Combination, Lapsus) Roussettus buttikoferi: Johnston, Liberia, Volume II: 690 - footnote. (Name Combination) Pterocyon palaeceus: Ellerman, Morrison-Scott and Hayman, Southern African mammals 1758 to 1951, 44. - Comments: Lapsus: see Meester et al. (1986: 28). (Lapsus) Pterocyon palaceus: Koopman, Bull. Am. Mus. Nat. Hist., 154 (4): 357. - Comments: Lapsus: see Meester et al. (1986: 28). (Lapsus) Eidolon helvum annobonensis Juste, Ibáñez and Machordom, Biol. J. Linn. Soc., 71: 373. Type locality: Equatorial Guinea: Annobón Island [01 24 N 05 38 E]. Eidolon helvum helvum: (Current Combination) GENERAL DISTRIBUTION: See species, with the exception of SW Arabia. Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Mali, Niger, Nigeria, São Tomé and Principé, Senegal, Sierra Leone, Sudan, Tanzania, Uganda. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Cameroon, Chad, Congo, Congo (Democratic Republic of the), Equatorial Guinea, †Subfamily Propottininae Butler, 1984 *1984. Propottininae Butler, Palaeovert., 14 (3): 118, 175. Publication date: 15 November 1984 [Goto Description]. - Comments: Recognized as a subfamily. (Current Combination) 52 ISSN 1990-6471 TAXONOMY: Currently recognized genera of Propottininae: Propotto Butler 1984. †Genus Propotto Simpson, 1967 *1967. Propotto Simpson, Bull. Mus. comp. Zool., 136: 50 [Goto Description]. - Etymology: Meant to imply an antecedent but not necessarily ancestral relative of the potto. Although the valid generic name of the potto too is probably Perodicticus, Potto has also been used. The valid specific name of the potto is probably Perodicticus potto although several other names are also in use (Simpson, 1967). (Current Combination) TAXONOMY: Simpson (1967) originally described this as a new genus of primates, within the Lorisiforms. Walker (1969) reexamining the type, noted that it was not a Lorisiform, but rather a fruit bat. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: If Archaeopteropus, from the Oligocene of Italy, is removed from the Megachiroptera following Russell and Sigé (1970), Propotto becomes the oldest known member of the suborder (Butler, 1978). Butler (1984: 178) believed it to be unlikely that Propotto is ancestral to any of the living Pteropodid genera. †Propotto leakeyi Simpson, 1967 *1967. Propotto leakeyi Simpson, Bull. Mus. comp. Zool., 136: 50 [Goto Description]. - Etymology: Named for Dr. Louis Seymour Bazett Leakey, "in gratitude and admiration" (Simpson, 1967). (Current Combination) TAXONOMY: Simpson (1967) originally described this as a new species of primate, within the Lorisiforms. Walker (1969) reexamining the type, noted that it was not a primate, but rather a fruit bat. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Miocene (Burdigalian - Brown et al., 2019: Suppl.), Songhor (Simpson, 1967; Walker, 1969), probably Rusinga (Butler, 1978). Subfamily Pteropodinae Gray, 1821 *1821. Pteropodinae Gray, London Med. Repos., 15: 299. (Current Combination) 1831. Pteropodina Bonaparte. - Comments: Introduced as tribe in the Vespertilionidae. Recognized as subtribe by Koopman and Jones (1970: 23) (see Jackson and Groves (2015: 236). 1835. Pteropodae Bonaparte, Iconogr. Fauna Ital. (Fascicolo 14 under Dysopes cestonii, second of four unnumbered pages). - Comments: Introduced as family (see Jackson and Groves, 2015: 236). 1866. Epomophorina Gray, Proc. zool. Soc. Lond., 1866, I (v): 65. Publication date: May 1866. - Comments: Not included in Simmons (2005) since it was not proposed as family name, but as a subdivision within the Pteropodidiae (Simmons, pers. comm.). 1891. Pteropodinae: Flower and Lydekker, Mammals Living and Extinct, 650. - Comments: Proposed as subfamily. Originally included the genera Epomophorus E. Bennett, 1836; Pteropus Brisson, 1762; Xantharpyia J. Gray, 1843 [= Rousettus J. Gray, 1821]; Boneia Jentink, 1879 [= Rousettus J. Gray, 1821]; Cynopterus F. Cuvier, 1824 [1821-1825]; Harpyia Illiger, 1811 [= Nyctimene Borkhausen, 1797]; Cephalotes É. Geoffroy, 1810 [= Nyctimene Borkhausen, 1797]; Pteralopex Thomas, 1888 (see Jackson and Groves, 2015: 236). 1897. Pteropinae Trouessart, Cat. Mamm. Viv. Foss., 1: v, 77. - Comments: Bergmans (1997: 64) assigned it to Gray, 1821. Jackson and Groves (2015: 236) assigned it to Trouessart, 1897, but also mention a misspelling by Simpson (1945). African Chiroptera Report 2020 1912. 1970. 53 Cynopterinae K. Andersen, Cat. Chiroptera Brit. Mus. Publication date: 23 March 1912. - Comments: Bergmans (1997: 69) considers this a valid subfamily in which he includes the genera Cynopterus, Ptenochirus, Megaerops, Dyacopterus, Balionycteris, Chironax, Thoopterus, Sphaerias, Aethalops, Penthetor, Latidens, Alionycteris, Otopteropus, Haplonycteris. Cynopterini Koopman and J.K. Jones Jr., Classification of Bats. - Comments: Not included in Simmons (2005) since it was not proposed as family name, but as a subdivision within the Pteropodidiae (Simmons, pers. comm.). TAXONOMY: Currently recognized genera of Pteropodinae: Acerodon Jourdan, 1837, Desmalopex Miller, 1907, Macroglossus F. Cuvier, 1824, Melonycteris Dobson, 1877, Mirimiri Helgen, 2005, Neopteryx Hayman, 1946, Notopteris Gray, 1859, Pteralopex Thomas, 1888, Pteropus Brisson, 1762, Styloctenium Matschie, 1899, Syconycteris Matschie, 1899. TRIBE Pteropodini Gray, 1821 *1821. Pteropodini Gray, London Med. Repos., 15: 299. TAXONOMY: Includes the genera Acerodon Jourdan, 1837, Neopteryx Hayman, 1946, Pteropus Brisson, 1762 and Styloctenium Matschie, 1899. Genus Pteropus Brisson, 1762 *1762. Pteropus Brisson, Regnum Animale, 2nd ed: 13, 53 [Goto Description]. - Comments: Type species: Vespertilio vampirus niger Kerr, 1792. Allen (1939a: 59), Corbet and Hill (1992: 57), and Simmons (2005) mention Brisson, 1762 [Règne Animal, ed. 2, pp. 13, 153] as author. This name was long time considered to be "not available" (see Hopwood, 1947). However, Opinion 1894 (ICZN, 1998: 64) ruled that the name (and authorship) is available. Not Pteropus Thunberg, 1815 (Insecta, Phasmatodea, Phylliidae - leaf insects); not Pteropus Jennings, 1828 (Aves, Gruiformes, Heliornithidae); not Pteropus Canestrini and Fanzago, 1878 (Arachnida, Mesostigmata, Spinturnicidae) (see Jackson and Groves, 2015: 238). - Etymology: From the Greek "πτερόπους", meaning wing-footed, referring to the wing membrane which arises from the side of the back and the back of the second toe (see Palmer, 1904: 596; Jones and Kunz, 2000: 5). (Current Combination) 1799. Spectrum Lacépède, Tabl. Div. Subd. Orders Genres Mammifères, 15. - Comments: Type species: Vespertilio vampirus Linnaeus, 1758 (according to Miller and Wilson, 1997: 1; Jackson and Groves (2015: 238) or Vespertilio vampirus niger Kerr, 1792 (according to Andersen, 1912b: 61). Preoccupied by Spectrum Scopoli, 1777 [Lepidoptera, Sphingidae] (see Allen, 1939a: 59; Corbet and Hill, 1992: 57; Pavlinov et al., 1995: 62; Miller and Wilson, 1997: 1; Jones and Kunz, 2000: 1). Not Spectrum Stoll, 1787 (Insecta, Orthoptera, Phasmatidae) (see Jackson and Groves (2015: 238). - Etymology: From the Latin "spectrum", meaning apparition or specter (see Palmer, 1904: 638). 1866. Eunycteris Gray, Proc. zool. Soc. Lond., 1866, I (v): 64. Publication date: May 1866. Comments: Type species: Pteropus phaiops Temminck, 1837 (=Pteropus melanopogon Peters, 1867). 1870. Pselaphon Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 110. - Comments: Type species: Pteropus pselaphon Layard, 1829, by monotypy. Not Pselaphus [= Pselaphon] Herbst, 1792 (Insecta, Coleoptera, Psephalidae) (Jackson and Groves, 2015: 238). - Etymology: From the Greek "ψηλαφάω", meaning to grope about (see Palmer, 1904: 588). 1899. Sericonycteris Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 6, 30. - Comments: Type species: Pteropus rubricollis E. Geoffroy Saint-Hilaire, 1810 (= Pteropus subniger Kerr, 1792), by original designation (according to Miller and Wilson, 1997: 1) or subsequent 54 ISSN 1990-6471 1907. 2011. designation (Palmer, 1904: 629; according to Jackson and Groves, 2015: 238). Originally described as a subgenus of Pteropus. Matschie dated his introduction on 14 May 1899. - Etymology: From the Greek "σηρικόν", meaning silk and "νυκτερίς", meaning bat (see Palmer, 1904: 627). Desmaplex Miller, Bull. U.S. natl. Mus., 57: vii, 60. Publication date: 29 June 1907. Comments: Type species: Pteropus leucopterus Temminck, 1853, by original designation (see Jackson and Groves, 2015: 238). Pterocarpus Mishra, Rout and Panda, Afr. J. Pharm. Pharmacol., 5 (1): 6. Publication date: January 2011. - Comments: Not of Jacquin, 1763, which is a pantropical genus of trees in the family Fabaceae (see http://en.wikipedia.org/wiki/Pterocarpus). (Lapsus) TAXONOMY: See Bergmans (1991) and Simmons (2005: 334). The International Commission on Zoological Nomenclature ruled in favor of rejecting Brisson (1762), but conserved several of the generic names including Pteropus (Opinion 1894; ICZN, 1998). Formerly included arquatus and leucotis which were transferred to Acerodon by Musser et al. (1982a). O'Brien et al. (2009) examined the western Indian Ocean member of the genus Pteropus, based on three mitochondrial genes – cytochrome b, 12S rRNA, and a portion of the control region. The combined data set indicates the lack of reciprocal monophyly in western Indian Ocean Pteropus. With the expetion of P. livingstonii and P. voeltzkowi, the other regional taxa demonstrate little genetic differentiation. The resulting poytomy indicates rapid and presumably recent divergence. Almeida et al. (2014: 83) studied the cytochrome b and the 12S rRNA mitochondrial genes sequences for all Pteropus species and some additional sequences of the nuclear RAG1, vWF, and BRCA1 genes for a subsample of taxa. Their conclusions were that the genus as a whole is monophyletic (originating in the Miocene), but also that the monophyly of the traditional speciesgroups could not be substantiated. They subdivided the genus in 13 species groups, where livingstonii and voeltzkowi are the only members of the "livingstonii" group. aldabrensis, niger, rodricensis, rufus, and seychellensis are part of the "vampyrus" group, whereas the extinct subniger could not be tested and is tentatively placed in an "incertae sedis" group. Currently (Simmons and Cirranello, 2020) recognized species of Pteropus: admiralitatum Thomas, 1894 – Solomon Isls; Admiralty Isls, New Britain, and Tabar Isl (Bismarck Arch.) (Simmons, 2005: 334); aldabrensis True, 1893; alecto Temminck, 1837 – Sulawesi, Saleyer Isl, Lombok, Bawean Isl, Kangean Isls, Sumba Isl, and Savu Isl (Indonesia); north and east Australia; south New Guinea (Simmons, 2005: 335); allenorum Helgen, Helgen and Wilson, 2009; anetianus (Gray, 1870) – Vanuatu including Banks Isls (Simmons, 2005: 335); aruensis Peters, 1867 – Aru Isls (Indonesia) (Simmons, 2005: 335); brunneus Dobson, 1878 – Percy Isl., Queensland (Australia) (Simmons, 2005: 335); caniceps Gray, 1870 – Halmahera (Indonesia) (Simmons, 2005: 335); capistratus Peters, 1876 – Bismarch Arch. (Papua New Guinea) (Simmons, 2005: 336); chrysoproctus Temminck, 1837 – Ambon, Buru, Seram, and small islands east of Seram (Indonesia) (Simmons, 2005: 336); cognatus K. Andersen, 1908 – Makira and Uki Ni Masi Isls (Solomon Isls) (Simmons, 2005: 336); conspicillatus Gould, 1850 – north Moluccas (Indonesia), New Guinea and West Papuan Isls (Raja Ampat Isl, off northwest coast of New Guinea), northeast Queensland (Australia) (Simmons, 2005: 336); coxi Helgen, Helgen and Wilson, 2009; dasymallus Temminck, 1825 – Taiwan, Ryuku Isls, and extreme southern Kyushu (Japan), Batan, Dalupiri, and Fuga Isls (Philippines) (Simmons, 2005: 337); ennisae Flannery and White, 1991 – New Ireland; faunulus Miller, 1902 – Nicobar Isls (India) (Simmons, 2005: 337); fundatus Felten and Kock, 1972 – Banks Isls (Vanuatu) (Simmons, 2005: 337); gilliardorum Van Deusen, 1969 – New Britain Isl. and New Ireland (Bismarck Arch.); griseus (E. Geoffroy Saint Hilaire, 1810) – Timor, Samao Isl, Dyampea Isl, Bonerato Isl, Saleyer Isl, Paternoster Isls, Pelang Isl, Sulawesi, and Banda Isls (Indonesia) (Simmons, 2005: 337); howensis Troughton, 1931 – Ontong Java Isl (Solomon Isls) (Simmons, 2005: 338); hypomelanus Temminck, 1853 – Andaman and Maldive Isls, New Guinea through Indonesia to Vietnam and Thailand, and adjacent islands, Philippines (Simmons, 2005: 338); intermedius K. Andersen, 1908 – southern Burma and western Thailand (Simmons, 2005: 338); keyensis Peters, 1867 – Kai Isls (Indonesia) (Simmons, 2005: 339); leucopterus Temminck, 1853 – Luzon, Catanduanes, and Dinagat Isls (Philippines) (Simmons, 2005: 339); livingstonii Gray, 1866; lombocensis Dobson, 1878 – Lombok, Sumbawa, Komodo, Flores, Lembata, Pantar, Alor and Timor Isls (Indonesia) (Simmons, 2005: 339); loochoensis Gray, 1870 – Okinawa Isl, Ryûkyû Isls (Japan) (Simmons, 2005: 339); lylei K. Andersen, 1908 – Thailand, Vietnam, Cambodia (Simmons, 2005: 339); macrotis Peters, 1867 – New Guinea, African Chiroptera Report 2020 Aru Isls (Indonesia), Boigu Isl (Australia) (Simmons, 2005: 340); mahaganus Sanborn, 1931 – Bougainville Isl (Papua New Guinea), Ysabel Isl and Choiseul Isl (Solomon Isls) (Simmons, 2005: 340); mariannus Desmarest, 1822; medius Temminck, 1825 – Maldive Islands, India (incl. Anadaman Islands), Sri Lanka, Pakistan, Bangladesh, Burma (=Myanmar), Nepal, China (incl. Tsinghai [=?Qinghai]): melanopogon Peters, 1867 – Amboina, Buru, Seram, Banda Isls, Yamdena (= Timor Laut), and adjacent islands (Indonesia) (Simmons, 2005: 340); melanotus Blyth, 1863 – Nicobar and Andaman Isls (India), Engano Isl and Nias Isl (Indonesia), Christmas Isl. (Simmons, 2005: 340); molossinus Temminck, 1853 – Pohnpei (= Ponape) and possibly Mortlock Isls (Caroline Isls, Micronesia) (Simmons, 2005: 341); neohibernicus Peters, 1876 – Bismarck Arch. and Admiralty Isls (Papua New Guinea), New Guinea, Misool and Gebi Isls, Gag Isl (Simmons, 2005: 341); niger (Kerr, 1792); nitendiensis Sanborn, 1930 – Nendö and Tömotu Neo (in the Santa Cruz Isls, Solomon Isls) (Simmons, 2005: 341); ocularis Peters, 1867 – Seram and Buru (Indonesia) (Simmons, 2005: 341); ornatus Gray, 1870 – New Caledonia and Loyalty Isls (Simmons, 2005: 342); pelagicus Kittlitz, 1836; pelewensis K. Andersen, 1908 – Pelew Isls (Micronesia) (Simmons, 2005: 342); personatus Temminck, 1825 – North Molucca Isls (Halmahera and Obi Isl groups), and Gag (Simmons, 2005: 342); pilosus K. Andersen, 1908 – Pelew Isls (Micronesia) (Simmons, 2005: 342); pohlei Stein, 1933 – Yapen, Biak-Spiori, Numfoor, and Rani Isls (off northwestern New Guinea) (Simmons, 2005: 342); poliocephalus Temminck, 1825 – Eastern Australia, from southern Queensland to Victoria (Simmons, 2005: 342); pselaphon Lay, 1829 – Bonin and Volcano Isls (Japan) (Simmons, 2005: 342); pumilus Miller, 1911 – Philippines (except Palawan region), Talaud Isls (Indonesia) (Simmons, 2005: 343); rayneri Gray, 1870 – Bougainville and Buka Isls (Papua New Guinea), Solomon Isls (Simmons, 2005: 343); rennelli Troughton, 1929 – Rennell Isl (Solomon Isls) (Simmons, 2005: 343); rodricensis Dobson, 1878; rufus E. Geoffroy Saint-Hilaire, 1803; samoensis Peale, 1848 – Fiji Isls, Samoan Isls (Simmons, 2005: 344); scapulatus Peters, 1862 – Australia, southern New Guinea, accidental on New Zealand (Simmons, 2005: 344); seychellensis A. Milne-Edwards, 1877; speciosus K. Andersen, 1908 – Philippines, Besar and Mata Siri (Java Sea), Talaud Isls (Simmons, 2005: 344); subniger (Kerr, 1792); temminckii 55 Peters, 1867 – Buru, Ambon, Seram (Indonesia), nearby small islands, perhaps Timor Isl (Indonesia) (Simmons, 2005: 345); tokudae Tate, 1934 – Guam (Mariana Isls, USA) (Simmons, 2005: 345); tonganus Quoy and Gaimard, 1830 – Karkar Isl (off northeastern New Guinea) and Rennell Isl (Solomon Isls), south to New Caledonia, east to Cook Isls (Simmons, 2005: 345); tuberculatus Peters, 1869 – Vanikoro Isl (Santa Cruz Isls, Solomon Isls) (Simmons, 2005: 345); ualanus Peters, 1883 – Kosrae (Micronesia) (Simmons, 2005: 345); vampyrus (Linnaeus, 1758) – Vietnam, Burum, Malay Peninsula, Borneo, Philippines, Sumatra, Java, and Lesser Sunda Isls, adjacent small islands including Anak Krakatau (Simmons, 2005: 346); vetulus Jouan, 1863 – New Caledonia (Simmons, 2005: 346); voeltzkowi Matschie, 1909; woodfordi Thomas, 1888 – New Georgia group, Russell and Florida Isls, Guadalcanal, Malaita (Solomon Isls) (Simmons, 2005: 346). COMMON NAMES: Czech: praví kaloni, upírové. Dutch: Kalongs. English: Flying-foxes, Flying Foxes. French: Renards volants, Roussettes géantes. German: Flugfüchse. GENERAL DISTRIBUTION: McNab (2009: 156) indicates that Pteropus "is primarily an island taxon, with 55 species (of the 57 species or 96.5 %) having some or all of their distribution on islands". Furthermore, McNab (2009: 158) states that the distributional pattern of Pteropus on islands is extremely variable, with some species being limited to small islands (e.g. P. voeltzkowi on Pemba, P. livingstonii on Anjouan and perhaps on Mohéli in the Comoros, P. rodricensis on Rodrigues, and P. niger on Mauritius, and other (Asian) species on islands that are hundreds or even thousands of kilometers apart. BIOGEOGRAPHY: Kingdon (1989) suggested that Pteropus doesn't occur in central parts of the continent since it might depend on seawater as source of salts. This is rejected by McNab and Armstrong (2001: 710) as this would not explain why there are no representatives of this genus in the African coastal areas. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Ethiopia, Madagascar, Mauritius, Réunion, Seychelles, Tanzania. Pteropus aldabrensis True, 1893 *1893. Pteropus aldabrensis True, Proc. U.S. Natl. Mus., 16 (948): 533. Publication date: 14 July 1893. Type locality: Seychelles: Aldabra Island. Syntype: USNM 20984: ♂, skin and 56 ISSN 1990-6471 1971. ? skull. Collected by: Dr. William Louis Abbott; collection date: 26 September 1892. Presented/Donated by: ?: Collector Unknown. Syntype: USNM 20985: ♂, skin and skull. Collected by: Dr. William Louis Abbott; collection date: 1892. Presented/Donated by: ?: Collector Unknown. - Comments: USNM 20984/36053 + 20985/36054: "cotypes" (syntypes); ad ♂♂, SS; coll. W.L. Abbott, 26 September 1892 and 5 October 1892: see Poole and Schantz (1942: 139), who also indicate "No type is specified in the original description, but these two specimens are mentioned by number and are therefore regarded as cotypes." Catalogued 30 June 1893: see Lyon and Osgood (1909: 256). Fisher and Ludwig (2015: 52) mention that Proc. U.S. Natl. Mus. 16: 1, (published on 14 July 1893) was a "Preprint" of Proc. U.S. Natl. Mus. 16: 533, which was published on 21 October 1893. (Current Combination) Pteropus seychellensis aldabrensis: Hill, Phil. Trans. Roy. Soc. Lond., B 260: 574. (Name Combination) [Pteropus seychellensis] aldabrabensis: Honacki, Kinman and Koeppl, Mammal species of the world: A taxonomic and geographic reference., 124. - Comments: Honacki et al. (1982: 124) mention "aldabrabensis" in the list of included taxa, and mention "P. aldabrensis (sic)." in the list with ISIS numbers. Bergmans (1991: 154) states it isn't clear to him why Honacki et al. (1982) changed the spelling. (Emendation) TAXONOMY: Bergmans (1991) showed that P. aldabrensis differs strongly from P. seychellensis, a view followed by Simmons (2005: 334). O'Brien et al. (2009) suggested that P. aldabrensis, P. rufus, P. niger and P. seychellensis (including comorensis) should be considered geographical forms of a single species P. giganteus (Brünnich, 1782) from the Indian subcontinent, Maldives and Andaman archipelagos. O'Brien (2011: 261) indicates that the last complete innundation of the Alabra atoll occurred some 125,000 years ago, and that the diversification of Pteropus in the area is therefore of recent age. Almeida et al. (2014: 83) places this in the 'vampyrus' group. COMMON NAMES: Castilian (Spain): Zorro volador de Aldabra. Czech: kaloň aldabránský. English: Aldabra Flying Fox, Aldabra Flying-fox. French: Renard volant d'Aldabra. German: Aldabra-Flugfuchs. Assessment History Global 2008: VU D2 ver 3.1 (2001) (Mickleburgh et al., 2008dh; IUCN, 2009). 2004: CR C2a(i) ver 3.1 (2001) (Mickleburgh et al., 2004cv; IUCN, 2004). Regional None known. Legal stutus CITES - Appendix II. MAJOR THREATS: The population is probably stable with no declines reported. However, because of the restricted nature of the species range it is considered to be especially vulnerable to threats such as tropical cyclones and rises in sea level (60% of the atoll is at or below 1 m asl.) (Mickleburgh et al., 1992; Hutson, 2004a; Justin Gerlach pers. comm., 2008; Mickleburgh et al., 2008dh; IUCN, 2009). CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU D2 ver 3.1 (2001)) because it is known from only a single location (the Aldabra Atoll), and it is plausible that this restricted range species could be threatened by tropical cyclones, similar stochastic events or a rising sea level (Mickleburgh et al., 2008dh; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008dh) [in IUCN (2009)] report that although the species is not directly protected (Tony Hutson pers. comm., 2008), the Aldabra Atoll is a highly protected UNESCO World Heritage site and a Special Reserve under the Seychelles National Parks and Nature Conservancy Act (Mickleburgh et al., 1992; Hutson, 2004a: 128). There is a distinct need for additional research into the total population number of this restricted range species and to monitor any changes in abundance (Hutson, 2004a: 128). Critically endangered - C2a(ii): see Jones et al. (2009b: 506). Conenna et al. (2017: Suppl.) mention its status as Vulnerable (VU). GENERAL DISTRIBUTION: Pteropus aldabrensis is endemic to the Aldabra Atoll (approximately 150 km 2) in the Seychelles. African Chiroptera Report 2020 Bats have been recorded from all main islands and have been observed flying between islands of the atoll (Hutson, 2004a; von Brandis, 2004). Native: Seychelles [Aldabra]. BIOGEOGRAPHY: See O'Brien et al. (2009) for further details. MOLECULAR BIOLOGY: DNA - See O'Brien et al. (2009) and Almeida et al. (2014). Karyotype - Unknown. Protein / allozyme - Unknown. ROOST: Racey and Nicoll (1984) [in O'Brien (2011: 262)] indicate that P. aldabrensis roosts in coconut palms, figs and Sideroxylon inerme. 57 with the largest roosting group on the small lagoon islet of Iles Michel that was probably well under 100 animals. Hutson (2004a) suggests that the total population was fewer than 250 animals in 1968, although he considered that the majority of the population may be present on the relatively little explored Middle Island (= Malabar Island) 26 km2 in size. von Brandis (2004) reports that large groups of these bats have been recorded in Casuarina trees, with the largest group of more than 100 animals reported at the eastern end of Middle Island (=Malabar Island) in 1995. Conrad Savy (pers. comm., 2008) indicates that the bats were commonly seen in good numbers during 2001 (Mickleburgh et al., 2008dh; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Seychelles. DIET: Hutson (2004a: 127) mentions that fruits of the following plants are eaten: Ficus lutea, F. rubra (as F. avi-avi) and F. reflexa, Calophyllum inophyllum, Terminalia catappa, Mystroxylon aethiopicum, and also flowers of Cocos nucifera and Agave sisalana. Roberts and Seabrook (1989) [in Courts (1998: 188)] report that P. seychellensis aldabrensis licked coccoids (Icerya seychellarum) from the leaves of infested Ficus trees, probably to supplement their diets. POPULATION: Structure and Density:- The population is reported to consist of only a few hundred animals (Cheke and Dahl, 1981; Carroll, 1985). Hutson (2004a) notes that no major colonies were located in 1968, Figure 7. Distribution of Pteropus aldabrensis Pteropus livingstonii Gray, 1866 *1866. Pteropus livingstonii Gray, Proc. zool. Soc. Lond., 1866, I: 66. Publication date: May 1866. Type locality: Comoros: Anjouan Island. Holotype: BMNH 1863.12.11.2: ad, skin and skull. Collected by: Dr. Livingstone. Presented/Donated by: Earl Russell. See Andersen (1912b: 250). (Current Combination) 1971. Pteropus livingstonei: Hayman and Hill, Mammals of Africa: Chiroptera, 10. (Lapsus) TAXONOMY: See Bergmans (1991), Simmons (2005: 339) and Smith and Leslie (2006: 1). P. livingstonii together with P. voeltzkowi form the most distinct and ancient lineage of the western Indian Ocean Pteropus spp. (O'Brien et al., 2009). The phylogenetic analyses performed by Chan et al. (2011: 7) suggest that both are sister species. This is supported by Almeida et al. (2014), who places this in the 'livingstoni' group. COMMON NAMES: Castilian (Spain): Zorro Volador De Livingston. Czech: kaloň komorský. Dutch: Zwarte Komorenkalong, Schoudervlekkalong. English: Comoro Black Flying Fox, Comoro Flying Fox, Black Flying- fox, Livingstone's Flying Fox, Comores Flying Fox. French: Renard volant des Comores, Renard volant noir, la roussette de Livingstone. German: Livingstones Flugfuchs. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: No fossils are known (Smith and Leslie, 2006: 2). 58 ISSN 1990-6471 CONSERVATION STATUS: Global Justification Sewall et al. (2016) provided the following justification for listing as Critically Endangered because of a serious population decline suspected from catastrophic habitat decline caused by cutting of trees for fuelwood and construction, and by conversion of all but the steepest upland areas to agricultural use; extensive declines in area of occupancy, extent of occurrence and quality of habitat. True population change in P. livingstonii is unknown and data on habitat change over time is limited. However, the species qualifies for CR under the A2c criterion because best estimates indicate habitat loss has exceeded 80% over the past three generations (estimated generation length: 8.1 years/generation), because remaining habitat is increasingly degraded and fragmented, and because declines in the extent and quality of habitat are continuing. P. livingstonii also meets the Endangered threshold under a second criterion, B1. The geographic range is small, such that the current extent of occurrence is estimated to be 1,856 km 2, less than the EN threshold of 5,000 km 2 under criterion B1. In addition, the habitat is severely fragmented, and there have been clear, continuing, observed declines in extent of occurrence, area of occupancy, habitat quality, and number of locations. Declines in number of mature individuals are also inferred from these declines in habitat. Therefore, this species would also be listed as EN under criterion B1ab(i, ii, iii, iv, v). Daniel et al. (2016: 742) supports the species listing as "Critically Endangered". Assessment History Global 2016: CE A2c ver 3.1 (2001) (Sewall et al., 2016). 2008: EN A2c; B1ab(iii) ver 3.1 (2001) (Mickleburgh et al., 2008t; IUCN, 2009). 2004: CR A4c ver 3.1 (2001) (Mickleburgh et al., 2004p; IUCN, 2004). 2002: CR A1c+2cd, B1+2c, C2a (Mickleburgh et al., 2002a: 22). 1998: EN A1c+2cd, B1+2c, C2a (Trewhella et al., 1998: 32). 1996: CR (Baillie and Groombridge, 1996). 1994: EN (Groombridge, 1994). 1993: EN (World Conservation Monitoring Centre, 1993). 1990: EN (IUCN, 1990). 1988: EN (IUCN Conservation Monitoring Centre, 1988). Regional None known. Legal Status CITES - Appendix II, see Smith and Leslie (2006: 4) for discussion. MAJOR THREATS: The species is threatened by the continuing degradation of its forest habitat by conversion of land to agricultural use, especially the use of lowland areas for export crops (such as cloves). In degraded forest the species is outcompeted by the bat Pteropus seychellensis (Mickleburgh et al., 2002a). Tree felling has additionally destroyed a number of roosts (Trewhella et al., 2005). There is also increased disturbance because of human population growth, however, the species is not hunted for food (Trewhella et al., 2005). The small remaining population is additionally potentially threatened in its restricted range by the effects of tropical cyclones (Mickleburgh et al., 2008t; IUCN, 2009). Since Anonymus (2000: 30) points out that this species generally roosts and feeds in native trees, the destruction of these forests can have a devastating effect, especially since P. livingstonii uses different tree species for food during different seasons. Sagot and Chaverri (2015: 1670) mention roost loss or disturbance, and habitat degradation or loss as major threats for this species. Daniel et al. (2016: 743) indicate that the estimated forest loss between 2000 and 2010 was 9.3 % per annum. Roost sites represent critical habitat for Pteropus livingstonii (Granek 2002, Sewall et al. 2007, Sewall et al. 2011b), yet tree felling has destroyed a number of roosts, leading to displacement of bats to less impacted areas, often at higher elevations (Trewhella et al., 2005, Sewall et al. 2011b). Of 23 roosts occupied in 2007 (Sewall et al. 2007), three roosts have been abandoned due to clear felling of forest in the past five years, and only one new but very small roost (15 bats) has been uncovered despite extensive searching effort (Daniel et al. in prep.). This new roost was found in a patch of degraded forest at 1050 m, the first to be found at such a high elevation, and is believed to have been established very recently. On Anjouan, forest clearance, underplanting or significant soil erosion following deforestation upslope of the roost was found within 50 m of all but one of the 16 occupied roosts (Daniel et al. in prep.). Further, the replacement of native forests with agricultural lands on Anjouan and Mohéli may render formerly inaccessible roost sites on Anjouan and Mohéli more accessible, possibly exposing roosts to increased levels of human disturbance. Finally, the loss of native forests has resulted in impermanence or complete drying of nearly all rivers on Anjouan and many on Mohéli African Chiroptera Report 2020 (Louette et al. 2004, Fernandez Astudillo 2012). As P. livingstonii roosts are often associated with rivers and other humid environments, a factor which may be related to their thermal sensitivity (Granek 2002), this loss of rivers may additionally affect the quality of roosting habitat. There is no evidence that the species is hunted for food (Trewhella et al., 2005), a key factor affecting other fruit bat species (Mickleburgh et al. 1992). This may be due in part to cultural taboos among the majority of the population that surround consumption of fruit bats. However, the apparent lack of hunting may also be due in part to P. livingstonii’s habit of roosting in inaccessible areas remote from towns; some limited hunting of the other two fruit bat species in the Comoros has been occasionally recorded (Sewall et al. 2003, B.J. Sewall, pers. obs.). It is unclear if the increased access to formerly inaccessible roost areas could expose the bats to any hunting pressure in the future, but any such hunting would likely first affect P. livingstonii congener, P. seychellensis comorensis, which sometimes roosts and forages in visible locations in or near towns. The mean surface temperature of the Comoros is thought to have increased about 1° C since 1900 and is expected to increase by 1-3° C more by 2100 due to global climate change (IPCC 2013). It is unclear whether or how such changes will affect P. livingstonii, but since thermal characteristics are important components of roost suitability for this and other Pteropus species (Pierson and Rainey 1992, Granek 2002) predicted temperature increases could affect availability of suitable roosting habitat. P. livingstonii small population is found within a restricted range solely on two small, adjacent islands. The species is therefore susceptible to single threatening processes, like cyclones, that could simultaneously or rapidly affect its entire range (Sewall et al. 2007). Further, the species may have naturally taken advantage of elevational differences in tree fruiting phenology to compensate for seasonal variation in fruit availability (Sewall 2002). Thus, the loss of large areas of foraging habitat including all the lower parts of its elevational range, could result in seasonally restricted food availability. CONSERVATION ACTIONS: Mickleburgh et al. (2008t) [in IUCN (2009)] report that a species action plan developed by the NGO Action Comoros is being implemented for this species (Sewall et al., 2003). A national environmental education programme has been implemented to raise awareness of this species 59 (Trewhella et al., 2005). National legislation to protect this species is being developed (Trewhella et al., 2005) and work has begun to establish a forest reserve for this bat on Moheli and on evaluating a site for a reserve on Anjouan (Trewhella et al., 2005). Sewall (2004: 10) suggests that the protection of specific "bat trees" can already provide a short-term solution. Since 1992, there is an active captive-breeding programme underway for this species initiated by the Durrell Wildlife Conservation Trust and the Bristol Zoo, where 35 animals are housed, of which 21 were born in captivity (Anonymus, 2000: 31). At the Jersey Wildlife Preservation Trust (JWPT), 6 bats (5 ♂♂ and 1 ♀) were imported from the Comores in 1992, and a second group with the same composition was imported in 1993, followed by an additional ♀ in 1995 (Carroll and Feistner, 1996: 334). Based on a study of mitochondrial and microsatellite markers, Ibouroi et al. (2018a: 1425) conclude that - for conservation purposes - the S. livingstonii populations from Anjouan and Mohéli should be considered as two separate management units, and that the focus should be targeted on the Anjouan population. A detailed conservation plan for the species, the Conservation Action Plan for Livingstone’s Flying Fox (Sewall et al. 2007) was developed by the non-governmental organizations Action Comores Anjouan and Action Comores International, and a consortium of other conservation groups, through a participatory process involving a broad range of stakeholders. The plan identifies a conservation strategy with several key elements, including habitat protection, forest management, environmental education, population monitoring, ecological research, ex-situ breeding, and conservation partnerships. This conservation plan was adopted by the government of the Union of the Comoros as the national conservation strategy for this species and has served as a guide for conservation action. A long-term comprehensive citizen science program involving Comorian villagers in population monitoring for this species was led by Action Comores Anjouan and Action Comores International between 1992 and 2006. Population surveys and habitat evaluation around roost sites was conducted by Engagement Communautaire pour le Développement Durable of the Bristol Zoological Society and Durrell Wildlife Conservation Trust between 2009 and 2012, and by Dahari more recently. A national environmental education programme was implemented by Action Comores Anjouan and Action Comores 60 ISSN 1990-6471 International, and it was successful in raising local and international awareness of this species and the threats it faces, in improving training of local personnel, and in increasing local participation in conservation and science programs (Trewhella et al. 2005). A small community-based ecotourism program for this species has been implemented on Mohéli by Projet Conservation de la Biodiversité et Développement Durable and local communities and is maintained by the local community in Ouallah 2 (Sewall et al. 2011b, Doulton et al. 2015). Dahari is implementing integrated landscape management around the southern forest block of Anjouan; this involves sustainable land management at the roosting and foraging habitat of this species and a pilot program of payment for ecosystem services both within the Moya Forest area (Doulton et al. 2015). Research by Dahari and partners into feeding ecology and the genetics of roost-site populations is also planned. There is an active captive-breeding programme underway for this species, initiated by Durrell Wildlife Conservation Trust in 1992. The program began with 17 founder individuals (7 females and 10 males) captured from the wild on Anjouan during 1992-1995 (Clark et al. 1997). The captive population of P. livingstonii subsequently expanded at a slower rate than other fruit bat species in captive breeding programs (e.g., P. rodricensis), at least initially (O’Brien 2011). However, by 2014, the P. livingstonii captive population had reached 59 individuals (22 females and 37 males) housed at four institutions (Durrell Wildlife Conservation Trust [Jersey, U.K.]; Bristol, Clifton, and West of England Zoological Society [Bristol, U.K.]; North of England Zoological Society [Chester, U.K.]; and Lisieux Cerza [Lisieux, France]) (Glenewar 2014). This species receives the highest level of legal protection available within the Union of the Comoros. It is listed as an ‘integrally-protected species’ (list 1 of RFIC 2001), which prohibits the capture or detention of P. livingstonii individuals without a permit. This law also expressly prohibits the killing of flying fox individuals; transport, purchase, sale, export or re-export of live or dead flying fox individuals or body parts; all disruption during the period of reproduction and raising of young; and the destruction of roosts (RFIC 2001). The Union of the Comoros also ratified the Convention on Biological Diversity in 1994, and in response has developed a National Biodiversity Conservation Strategy (Roby and Dossar 2000). This strategy highlights the importance of, threats to, and conservation recommendations for fruit bats of the Union of the Comoros (Sewall and Granek 2000). P. livingstonii is also listed on Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora, or CITES (UN 1973), which prevents international trade in specimens of this species without a permit. In practice, however, enforcement activities within the Comoros for these laws and treaties have been very limited in scope (Sewall et al. 2007). P. livingstonii also does not occur in any protected areas. However, critical roosting habitat at seven key roost sites that together harboured more than half the population was identified during the development of the conservation action plan (Sewall et al. 2007). Habitat conservation of these roost sites was identified as a key goal in this plan. As part of its implementation, Action Comores Anjouan and the Comoros Forest Reserves Project conducted a conservation assessment to identify the two highest-priority sites (roost sites Yiméré on Anjouan and Hassera-Ndrengé on Mohéli), where the establishment of protected areas for roost habitat would be both highly beneficial to biodiversity conservation and highly feasible (Sewall et al. 2011a). These two groups also completed conservation planning for protection of critical roosting habitat at all seven of these critical roost sites (Sewall et al. 2011b). Implementation efforts by Action Comores Anjouan and partners to conserve habitat around critical roosting habitat on Anjouan and Mohéli are under development. A project to establish larger reserves to protect rainforest and cloud forest on both Anjouan and Mohéli, which would include roosting and foraging habitat for this species, has been proposed by the United Nations Development Programme, the Global Environmental Facility, and the Comorian government. The success of habitat conservation and restoration efforts will be particularly critical to the long-term prospects for this species. GENERAL DISTRIBUTION: Pteropus livingstonii is endemic to two of the four Comoro Islands: Anjouan (formerly Johanna) and Moheli (Trewhella et al., 2001; Smith and Leslie, 2006: 1). On Anjouan, the species avoids lower parts of the island below 300 to 1100 m asl, and on Moheli between 150 to 700 m asl. Native: Comoros (Cheke and Dahl, 1981; Clark et al., 1997; Trewhella et al., 2001; Smith and Leslie, 2006: 1; Goodman et al., 2010c: 124). GEOGRAPHIC VARIATION: See Smith and Leslie (2006: 1). General description of exernal morphology: See Smith and Leslie (2006: 1). African Chiroptera Report 2020 DETAILED MORPHOLOGY: Baculum - Unknown For a description of the cranium, teeth, ear and tragus, and wingshape and aspect ratio see Smith and Leslie (2006: 1-2). Dickson et al. (2016: 963) and Killick et al. (2017: 1077) studied cardiomyopathies on a group of captive bats, where dilated cardiomyopathy may be a major cause of death (Drane et al., 2017: 89). SEXUAL DIMORPHISM: It is not clear if sexual dimorphism occurs in P. livingstonii due to insufficient numbers of adult females have been difficult to collect in the wild (Clark et al., 1997; Trewhella et al., 2005; Young et al., 1993; Smith and Leslie, 2006: 2). ECHOLOCATION: Not known to echolocate. MOLECULAR BIOLOGY: DNA - See O'Brien et al. (2009) and Almeida et al. (2014). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: Ibouroi et al. (2018b: 2395) reported that the species' habitat is restricted to steep, highelevation slopes of the Comoro's remaining natural forests. HABITS: See Smith and Leslie (2006). Based on wing morphology, Lindhe Norberg et al. (2000) classified P. livingstonii as a soaring species. ROOST: See Smith and Leslie (2006). Anonymus (2000: 30) indicates that P. livingstonii roosts in tall trees in the rain forest, at medium altitude. These nesting trees are, in general, local species. Trewhella et al. (2001) indicate that day-roost sites are also shared with P. s. comorensis. Daniel et al. (2016: 747) indicate that only four tree species were used as roost on Mohéli [M] and 12 on Anjouan [A]: Ficus exasperata Vahl (sandpaper tree, forest sandpaper fig, white fig, or sandpaper leaf tree) [M+A], Gyrostipula comoriensis J.-F. Leroy [A], Gambeya spp. [A], Ficus lutea Vahl (giant-leaved fig or Lagos rubbertree) [A], Nuxia pseudodentata Gilg [A], Albizia lebbeck (L.) Benth. (lebbeck, lebbek tree, flea tree, frywood, koko and woman's tongue tree) [M], Ocotea comoriensis Kosterm. [A], Weinmannia comoriensis Tul. [A], Brachylaena ramiflora (DC.) Humbert [A], Ficus 61 pirifolia Lamk. [A], Terminalia catappa L. (countryalmond, Indian-almond, Malabar-almond, seaalmond, tropical-almond or false kamani) [M], Antidesma hildebrandtii Pax & K. Hoffm. [A], Albizia glaberrima (Schumach. & Thonn.) [A], Khaya comorensis (Comorian mahogany) [M], Cryptocarya spp. [A]. Ibouroi et al. (2018b: Suppl.) reported the following roost trees being used on the Comoro Islands: Asteraceae: Brachylaena ramiflora; Combretaceae: Terminalia catappa; Cunoniaceae: Weinmannia comorensis; Gentianaceae: Anthocleista grandiflora; Lauraceae: Ocotea comorensis; Leeaceae: Leea guineensis; Loganiaceae: Nuxia pseudodenta; Meliaceae: Khaya comorensis; Mimosaceae: Albizia glaberrima, Albizia lebbeck; Moraceae: Ficus antandronarum, Ficus lutea, Ficus pirifolia; Myrtaceae: Eugenia comorensis; Sapotaceae: Gambeya boiviniana. Granek (2000a: 29 - 30) determined the following roost site characteristics: 1) between 500 and 1,100 m in elevation, 2) steeply sloped land, 3) some native forest cover was present, 4) presence of water in the same valley, 5) presence of an east- or south-facing slope 6) the valley was protected by mountains on three sides, forming a 'bowl' (protecting them from the wind and the mid-day sun), 7) presence of the following tree species: Gambeya sp. and Nuxia pseudodentata. However, Granek (2002: 95) indicates that there was no significant association between the bat roosts and characteristics 1, 2 and 3. MIGRATION: Intra-island movement among roosts is likely, but inter-island movement is unknown (Smith and Leslie, 2006: 3). DIET: See Smith and Leslie (2006: 3). Kock and Nader (1984: 30 - 31) and Sewall (2004:10) point out that this species almost completely feeds on native trees, and changes tree species according to the season. One of its favourite foods is the fruit of a giant-leaved fig tree (Ficus lutea) (Sewall, 2004: 9 - 10). Young et al. (1993) [in Carroll and Feistner (1996: 330)] indicate that R. obliviosus and P. livingstonii were captured at the same kapok and fig feeding sites on Anjouan, which would suggest that their feeding niches might overlap. However, they also indicate that there might be a vertical separation between the two species. Nectar and fruits of the Moraceae family form an important part of the diet, while pollen is not important Cotterill (1995). 62 ISSN 1990-6471 Trewhella et al. (2001: 141) reported the following food items (with known items in bold and suspected items in plain text): Aphloiaceae: Aphloia theaeformis [= Aphloia theiformis (Vahl) Benn.]; Asteraceae: Brachylaena ramiflora (DC.) Humbert; Clusiaceae: Rheedia anjouanensis H. Perrier [= Garcinia anjouanensis (H. Perrier) P.W. Sweeney & Z.S. Rogers; Cunoniaceae: Weinmannia comorensis Tul.; Gentianaceae: Anthocleista grandiflora L. (forest fever tree); Lauraceae: Ocotea comorensis ?; Malvaceae: Ceiba pentandra (L.) Gaertn. (kapok tree); Monimiaceae: Tambourissa comorensis Lorence; Moraceae: Artocarpus altilis (Parkinson) Fosberg (breadfruit), Artocarpus heterophyllus Lam. (jackfruit or jack tree), Ficus lutea Vahl (giant-leaved fig), Ficus pirifolia ?; Myrtaceae: Eugenia jambos L. [= Syzygium jambos L. (Alston)] (rose apple); Oxalidaceae: Averrhoa bilimbi L. (bilimbi, cucumber tree or tree sorrel); Sapotaceae: Gambeya sp.; Stilbaceae: Nuxia pseudodentata Gilg, 1895. Courts (1998: 189) reports that captive animals catch and eat insects flying by. Bell et al. (2019: 250) indicate that dominant male bats (in a captive colony) defend small feeding territories within the colony, and are therefore often more sedentary, which might result in these animals being heavier and running a higher risk of cardiomyopathy. PREDATORS: See Smith and Leslie (2006). POPULATION: Structure and Density:- In 2002 there were estimated to be 1,200 bats in 20 roosts, many of which have been recently located as a result of the implementation of a national environmental education programme (Trewhella et al., 2005). Sewall (2009: 3) indicates that the population might count 1,200 - 1,500 animals. Prior to the national environmental education programme there were estimated to be fewer than 200 bats. During an extensive survey in 2011 - 2012, Daniel et al. (2016: 742) estimated the population size to be about 1,260 animals, distributed over 21 roosts on Anjouan and Mohéli. Ibouroi et al. (2018b: 2408) observed 1,243 bats divided over 19 locations, with colony sizes ranging from 3 to 349 individuals (average: 65). Also see Smith and Leslie, 2006: 1). Repeated simultaneous surveys of all 23 known roosts during 1998-2006 typically recorded about 1,200 bats (Sewall et al. 2007); many of these roosts were located as a result of the implementation of a national monitoring program (Trewhella et al. 2005). Surveys have been conducted only sporadically since 2007, so it is unclear whether populations have changed recently. However, a survey conducted with sequential visits to all previously-known roosts in 2011 and 2012 estimated 1,300 bats across 22 roosts (Anjouan: 940 bats at 16 roosts, Mohéli: 360 bats at 6 roosts) (Daniel et al. in press.). Colony size typically ranges from 15 to 150 individuals (Sewall et al. 2007). The largest known roosts have sometimes reached about 250 individuals during the rainy season, but not all of these are mature individuals (Sewall et al. 2011 a,b). Generation length for the wild population of P. livingstonii is unknown, but several demographic parameters can be derived from the studbook for captive bats of this species (Glendewar 2014). Bats of this species can be long-lived in captivity (e.g., one founder captured as an adult in 1992 is still alive and has now lived 22 years in captivity). Females do not reproduce during the first few years of life in captivity (minimum dam age at first reproduction = 3.4 years, mean dam age at first reproduction = 5.9 years), and the average age of females giving birth is elevated (of all dams born in captivity that have since died or are >10 years of age, mean age when giving birth = 8.1 years). Thus, based on studbook data (Glendewar 2014), this last figure of 8.1 years represents our best estimate of the generation length of the captive population. This estimate may be conservative as most (75%) of these females are still alive and at least some presumably will reproduce again at a later age. It is unclear to what extent the generation length of the captive population correlates with the generation length of the wild population of P. livingstonii. However, the generation length of captive P. livingstonii correlates well with the few published generation length values for wild populations of other Pteropus species. In a wild population of P. poliocephalus, generation length was estimated conservatively as 7.4 years (Tidemann and Nelson 2011), though the true generation length could be much higher. Further, a generation length of 5.0 years was estimated for a rapidly declining wild population of P. conspicillatus (Fox et al. 2008); however, our calculations using these same data suggest that in a stable population, generation length for P. conspicillatus would be about 7.7 years. Thus, generation length values for the captive population of P. livingstonii correlate well with the only African Chiroptera Report 2020 published generation length values for wild Pteropus. Further, an estimate derived from taxonomicallyadjusted allometric equations for mammals suggests generation length for P. livingstonii is 8.1 years (Pacifici et al. 2013). Thus, the captive estimate appears in line with estimates for wild populations, and we conservatively estimate the generation length in the wild population of P. livingstonii to be 8.1 years. True changes in the population size of P. livingstonii are currently unknown, but are likely to be linked to changes in forest and underplanted forest habitat. Our estimates, derived from the little available data on forest loss rates (FAO 2010, UNDP 2013), suggest that habitat decline over the most recent threegeneration (or 24.3-year) period was 83%. Thus, our best estimate of loss of P. livingstonii habitat exceeds the threshold under the A2 criterion for CR status (=80% loss of population, as suspected by habitat change, over a three-generation period, where habitat loss is ongoing). 63 MATING: See Smith and Leslie (2006). Riccucci (2011: 141) reports on same sex sexual behaviour in captive held P. livingstonii. PARASITES: Fountain et al. (2019: 268) examined the captive population at the Jersey Zoo and found them to be infected by the following Staphylococcus species (in descending order): S. xylosus, S. aureus, S. saprophyticus, S. nepalensis, S. simulans, S. sciuri, S. kloosii, S. simiae, S. hominis, S. epidermidis, S. succinus, S. warneri, S. lentus, S. lugdunensis, S. cohnii, S. haemolyticus, S. equorum, S. capitis. UTILISATION: Is not hunted for food (Trewhella et al., 2005). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, Mayotte. Trend:- 2016: Decreasing (Sewall et al., 2016). 2008: Decreasing (Mickleburgh et al., 2008t; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 15 years. ACTIVITY AND BEHAVIOUR: See Smith and Leslie (2006: 1). Trewhella et al. (2001: 135) indicate that P. livingstonii is more diurnally active than the sympatric P. seychellensis comorensis, and is the dominant species during their encounters. REPRODUCTION AND ONTOGENY: See Smith and Leslie (2006: 2). Figure 8. Distribution of Pteropus livingstonii Szekely et al. (2015: Suppl.) mention a gestation period of 152 days. At birth, the young weights 137 g (adult weight: 696.5 g). Pteropus niger (Kerr, 1792) *1792. Vespertilio vampirus niger Kerr, in: Linnaeus, Animal Kingdom, 1 (1): xvii, 90. Publication date: February 1792. Type locality: Réunion: "Réunion island". - Comments: Andersen (1912b: viii) mentions that the type was once in the ancient Royal Cabinet in Paris, but is probably no longer in existence. 1797. Vespertilio caninus Blumenbach, Handb. Naturg, 5 ed.: 73. - Comments: Pavlinov et al. (1995) consider caninus to be a synonym of vampyrus, but Andersen (1912b: 220) indicates this is a synonym of niger Kerr, 1792. The name was originally proposed as a replacement for V. vampyrus L., but this includes vampyrus, rufus and niger. Andersen also indicates that Goldfuss' plate of "V. caninus" Blumenbach (1809), is a copy of Schreber's plate xiv, which is Pteropus niger. 64 ISSN 1990-6471 1803. 1804. 1808. 1810. 1870. 1895. ? Pteropus fuscus E. Geoffroy Saint-Hilaire, Catalogue des Mammifères du Muséum national d'Histoire naturelle, Paris, 46. Type locality: Réunion: Réunion and Madagascar [Goto Description]. - Comments: Andersen (1912b: 220) mentions that Geoffroy based himself on three specimens "viz., two mounted specimens (wings expanded), sent from Réunion by de la Nux (nos. 86 and 87), and a third 'envoyé de Madagascar, par le citoyen Macé, naturaliste' (no. 88)". Andersen also indicates that specimen 88 (from Madagascar) was probably obtained from a dealer. He was also unable to find any of these specimens. V[espertilio] mauritianus Hermann, Obs. Zool. , 19. Publication date: 31 August 1804. Type locality: Mauritius: "Mauritius" [Goto Description]. Pteropus rufus: Tiedemann, Zool, 1808: 535. Type locality: Madagascar: "Madagascar". - Comments: Not of E. Geoffroy Saint-Hilaire, 1803, but see). Pteropus vulgarisE. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 15: 92. Type locality: Réunion: Réunion and Madagascar: Restricted to Réunion by Andersen (1912b: 221), due to the type locality for Pteropus fuscus is Réunion. Holotype: MNHN A.1.C.18 - 745H - 154: ad ♂, mounted skin (skull not removed). - Comments: A redescription and renaming of Pteropus fuscus Geoffroy, 1803 (see Andersen, 1912b: 221). Spectrum vulgare Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 100. Type locality: : Mauritius, Réunion, ? Madagascar [Goto Description]. Pteropus pteropus Merriam, Science, N.S. 1 (14): 376. Publication date: 5 April 1895. Type locality: Réunion: Réunion and Madagascar. - Comments: Combination introduced into literature by Merriam on the mistaken supposition that Brisson's nomenclature was binomial (see Andersen, 1912b: 223). Pteropus niger: (Name Combination, Current Combination) TAXONOMY: See Bergmans (1991) and Simmons (2005: 341). O'Brien et al. (2009) suggested that P. aldabrensis, P. rufus, P. niger and P. seychellensis (including comorensis) should be considered geographical forms of a single species P. giganteus (Brünnich, 1782) from the Indian subcontinent, Maldives and Andaman archipelagos. Almeida et al. (2014: 85) place P. niger in the "vampyrus" group. COMMON NAMES: Castilian (Spain): Zorro volador negro de Mauricio. Czech: kaloň mauricijský. English: Greater Mascarene Flying Fox, Mascarene Flying-fox, Mauritian Flying Fox. French: Renard volant de l'île de Mascarene, Renard volant de Maurice, Renard volant des Mascaraignes, Roussette noire, grande Roussette des Mascareignes. German: Grosser Maskarenen-Flugfuchs. CONSERVATION STATUS: Global Justification Listed as Vulnerable based on a projected 30% population reduction in the next 21 years (three generations) due to the continuing decline in the extent and quality of a key habitat (forest, which is relied upon for roosting and at certain times of year for foraging); knowledge of impacts of past cyclones on this and other island fruit bats and the likelihood of this happening again on Mauritius within the next few years; and based on current pressures from hunting, which are likely to increase (Hutson and Racey, 2013).. Conenna et al. (2017: Suppl.) mention its status as Vulnerable (VU), but Anthony et al. (2018: 12075) indicated that the culling ot the bats in 2015-2016 resulted in upgrading the status to "Endangered" by the IUCN in 2018. Assessment History Global 2013: VU A3cd ver 3.1 (2001) (R). 2008: EN B1ab(iii,v) ver 3.1 (2001) (Jenkins et al., 2008k; IUCN, 2009). 1996: VU A1d+2cd ver 2.3 (1994) (Baillie and Groombridge, 1996). 1994: VU A1d+2cd ver 2.3 (1994) (Groombridge, 1994). 1993: VU (World Conservation Monitoring Centre, 1993). 1990: VU (IUCN, 1990). 1988: VU. 1986: Rare (IUCN Conservation Monitoring Centre, 1986). 1981: VU: A1d+2cd. Extinct on Réunion Island (see Cheke and Dahl, 1981). Regional None known. Legal Status CITES - Appendix II. MAJOR THREATS: Like other Pteropus species in the region, this species is threatened by cyclones, the loss of African Chiroptera Report 2020 65 roosting and feeding trees, and by persecution and/or hunting (Carroll and Feistner, 1996). Although no major cyclone has affected bats on Mauritius in recent years (Hutson and Racey, 2013), the risk remains high and climate change forecasts suggest that there will be an increase in the frequency and intensity of cyclones in this region. Cyclone damage can result in high levels of starvation and mortality, as well as loss of roost trees. Elsewhere, such cyclones have caused massive declines (up to 99%) in island fruit bat populations. In the previous 10 years six intense tropical cyclones had affected Mauritius (Hutson and Racey, 2013). Note that, for example, a cyclone in the 1970s reduced the population of the endemic fruit bat on the neighbouring island of Rodrigues to less than 1% of its current population (Powell and Wehnelt, 2003). Racey, 2013). Even if foraging area has increased following the wider introduction of fruit farms, especially for litchi, longan and mango, these are only available for a short period of the year and it is believed that there are times of year when the bats would not survive well without forest foraging. Forest habitat suitable for roosting and important for feeding, particularly at certain times of year, have seen major decline of what is already a habitat much depleted in both area and quality (1520% decline in the 10 years to 2005). While the significant holdings by government have shown little decline in recent years, private holdings have seen the majority of the declines, and that decline is likely to continue (Hutson and Racey, 2013). Hutson and Racey (2013) estimates the existing levels of take for sport or as measures for pest control are given at about 2,000 bats per year. This is between 4 and 10% of the recent population estimates. Any introduction of a cull that would significantly reduce fruit bat damage to fruit (mainly litchi, longan and mango) must similarly impact on the bat population. Since the species is currently listed by IUCN as Vulnerable and is protected under Mauritian law (Wildlife and National Park Act 1993, currently under revision), the introduction of a cull is likely to result in the species being removed from the protected list and would be aimed at making a significant impact on the population, but is also very likely to increase the exploitation for sport and for unmanaged control. Hence, the moment a cull were introduced, it would immediately trigger the need for a reevaluation of the status of the species. While no full analysis of sustainable take is available for such fruit bats it is widely considered that a take of 10% per year will result in significant population decline (e.g.Pierson and Rainey, 1992). Page and D’Argent G. (1997) demonstrated that less than 2% of Mauritius’ native forest remained, and this was recognized in the Ministry of Environment and Sustainable Development’s 2006 Pocketbook of Environmental Statistics. Forest areas currently occupy about 25% of the total land area, but many of these areas are heavily invaded by exotic vegetation. The report on Economic and Social Indicators, Environment statistics 2009, from the Central Statistics Office (quoted in National Economic and Social Council Report 17, 2011, Maintaining the Green Cover of Mauritius) identifies a loss of forest area amounting to a change of 5.1% of the total land area over the period 1995 to 2005. From the figures given, currently half the forest area is privately owned. The report identified loss to state owned forest of 1.5%, but a loss of 27.6% of privately owned forest. Very little change was identified for the four following years to 2009. The same report quotes from Digest of Environment Statistics 2009, CSO, giving a loss of forest, shrubs and grazing land of 10,000 ha (17.2%) between 1995 and 2005. The data suggests that the government is maintaining its own holdings, indeed Appendix 3 to this report identified a number of government initiatives for the maintenance, restoration and creation of forest areas, but at present these are mainly new and small initiatives and in no way compensate for the losses to privately owned forest (Hutson and Although protected (Nyhagen, 2004), there is currently a level of control and sport hunting that is likely to be sustainable (Robinson et al., 2010). However, there is increased pressure to reduce populations, through conflict with fruit growers at certain times of year. Any significant increase in the take of fruit bats is likely to have serious impact on the populations and combined with the other major threats described above could threaten the survival of this endemic species. In October 2006 the government endorsed a culling programme for 2,000 bats because of their perceived role in inflicting serious economic damage on lychees. This is obviously in conflict with the fact that the species is protected under the Wildlife and National Parks Act 1993. Only six bats were officially culled, despite illegal hunting parties being known to kill up to a few hundred bats in one night. The law is currently being revised and it is unknown whether further culling will be supported, although most likely it will be (Jenkins et al., 2008k; IUCN, 2009). Anthony et al. (2018: 12075) reported that in November-December 2015 30,938 bats were culled and in December 2016 yet another 7,380 bats. Florens and Baider (2018: 60) furthermore 66 ISSN 1990-6471 reported that 12,500 bats were planned to be culled in 2018. these trees, and that protecting trees with nylon nets could yield 1/3 extra fruits. Olival (2015: 6) reviews why culling is not an effective means to mitigate conflicts between fruit growers and bats, nor to reduce the likelihood of zoonotic disease risk. GENERAL DISTRIBUTION: Historically, Pteropus niger is distributed within the Mascarene Islands (Reunion Island, Mauritius Island, subfossil on Rodriquez Island). Since becoming extirpated from La Réunion in the early eighteenth century, this species has been restricted to the western Indian Ocean island of Mauritius. In 2007 a small colony of P. niger (up to 6 individuals) was located on La Réunion, though a few individuals have been observed on previous occasions over the past decade, most likely blown over from Mauritius by annual cyclones. Breeding within this small population on La Réunion is suspected but not confirmed. Caceres (2010: 9) mentions that the orginal population disappeared from La Réunion between 1772 and 1801. Florens (2015: 1325; 2016: 33) indicated that the official population estimates are between 90,000 and 1,000,000 bats. The government set a cull target of 20 %, which resulted in the death of 20,000 bats. However, as the real number of bats accounted for about 50,000 animals, this meant that about half of the population was eradicated. Hutson and Racey (2013) report that there is no data on specific threats to specific roost sites, but the threats to bats at roost sites in Mauritius (principally deforestation, hunting and cyclones) can impact a number of roosts at one time and in no particular pattern. In the case of a larger cyclone a very large number of roosts can be damaged at the same time and which roosts are damaged will depend on details of the movement of the cyclone and its strength. These cyclones also affect the foraging of the bats, both around the roost site and elsewhere and there are many examples of where cyclone damage has resulted in high levels of starvation and mortality, in addition to damage to roost trees, in island-dwelling fruit bat species. CONSERVATION ACTIONS: Hutson and Racey (2013) repeats what is stateed by Jenkins et al. (2008k) [in IUCN (2009)] who report that P. niger became a protected species in 1993 on Mauritius (Nyhagen et al., 2005) and is reported to occur in protected areas, including the Black River Gorges National Park (Nyhagen, 2004), nature reserves and mountain reserves. There is a clear need to assess the population size, population dynamics, response to cyclones, assess relative fruit predation, roosting habitat requirements and impact of culling on this species and effectiveness of fruit protection to prevent foraging. Oleksy et al. (2018: "5") also suggest to harvest the fruits at an earlier stage to reduce their attractiveness to bats as bats prefer to eat fruits that are to ripe to be sold. Additionally, they found that 10 % of the lychee losses and up to 50 % (or more) of the mango losses could be attributed to natural falling of fruit, which could also be reduced by early harvesting. Their study also indicated (p."6") that netting of trees could reduce damage caused by bats 12 to 23 fold. Another study concentrating on backyard lychee trees, Tollington et al. (2019: 1) suspected that fruit bats were responible for about 42 % of the total damage to Native: Mauritius; Réunion. BIOGEOGRAPHY: See O'Brien et al. (2009) for further details. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: The adult's weight is on average 456.3 g (see Szekely et al., 2015: Suppl.), but Oleksy et al. (2019: 5) reported an average weight of 617 (± 76) g for females and 630 (± 96) g for males. MOLECULAR BIOLOGY: DNA - See O'Brien et al. (2009). Karyotype - Unknown. Protein / allozyme - Unknown. HABITS: P. niger is primarily nocturnal, but Nyhagen et al. (2005: 492) observed some specimens foraging during the day. Using solar-powered GSM-tracking, Oleksy et al. (2019: 5) found that on average males traveled 9.38 km each night and females 6.06 km. The maximum distance traveled during one night was 92.92 km for a male and 79.19 km for a female. They also found that home ranges of males averaged 74,633 ha, whereas these for females only covered 31,072 ha. The core foraging areas averaged 2,222 ha for males, 1,364 ha for females. ROOST: On La Réunion, Caceres (2010: 14) reported that Yellow jade orchid tree (Michelia champaca, Chinese fan palm (Livistona chinensis), Longan (Dimocarpus longan), soft bollygum (Litsea glutinosa) and Wampee (Clausena lansium) are used as roosting trees. These roost trees can be African Chiroptera Report 2020 used for several years, although during the year several roosts on different places of the island can be used. DIET: Nyhagen et al. (2005: 493) reported that on Mauritius, P. niger ate 22 plant species, 20 of which were visited for fruit, two for floral resources and one for leaves (of Diospyros tessellaria both fruits and flowers were eaten). Nyhagen (2004: 120) described the characteristics, which fruit eaten by P. niger posses: easily accessible (95 %), medium size (47 %), medium size seed (68 %), vivid colours (53 %), sweet smell (58 %), sweet flavour (74 %), and not juicy, i.e. low water content (58 %). Four of the plant species (Artocarpus heterophyllus, Diospyros tessellaria, Mangifera indica and Mimusops petiolaris) possess five of these seven characteristics. Nyhagen (2004: 122) also mentions that this bat is the only native vertebrate to feed on fruits of Labourdonnaisia glauca. Seltzer et al. (2013: table S2) provide an overview of the plant species found beneath bat feeding roosts. For P. niger these include Psidium guajava L. (apple guava - Myrtaceae) and Syzygium jambos L. (Alston) (Malabar Plum Myrtaceae). Florens et al. (2017: 89) provided the following list of fruits eaten by P. niger: Aphoiaceae: Aphloia theiformis (Vahl) Benn.; Apocynaecae: Tabernaemontana persicariifolia Jacq.; Araliaceae: Polyscias maraisiana (Marais) Lowry & G.M. Plunkett (Ox tree); Arecaceae: Dictyosperma album (Bory) H. Wendl. & Drude exScheff. (princess palm, hurricane palm), Hyophorbe lagenicaulis (L.H. Bailey) H.E. Moore (bottle palm, palmiste gargoulette), Latania loddigesii Martius (Latanier de l'île Ronde, Latanier de Maurice); Burseraceae: Protium obtusifolium (Lam.) Marchand; Calophyllaceae: Calophyllum tacamahaca Willd., 1811; Celastraceae: Cassine orientalis (Jacq.) Kuntze (bois d'olive); Chrysobalanaceae: Grangeria borbonica Lam., 1789; Combretaceae: Terminalia bentzoe (L.) L. f.; Ebenaceae: Diospyros egrettarum I. Richardson, Diospyros tessellaria Poir. (black ebony, Mauritian ebony), Diospyros melanida Poir.; Elaeocarpaceae: Elaeocarpus integrifolius Lam.; Lecythidaceae: Foetidia mauritiana Lam.; Loganiaceae: Geniostoma borbonicum (Lam.) Spreng.; Melastomataceae: Warneckea trinervis (DC.) Jacq.-Fél.; Moraceae: Ficus reflexa Thunb., Ficus rubra Vahl, 1805, Ficus mauritiana Lam.; Myrtaceae: Eugenia alletiana Baider & V. Florens, Eugenia elliptica Lam., 1789, Eugenia pollicina J. Guého & A.J. Scott, Eugenia lucida Lam., Eugenia tinifolia Lam., Syzygium glomeratum (Lam.) DC., 67 Syzygium populifolium (Baker) J. Guého & A.J. Scott, 1980; Pandanaceae: Pandanus eydouxia Balf. f., Pandanus utilis Bory; Pittosporaceae: Pittosporum senacia Putt.; Rubiaceae: Pyrostria borbonica (J.F. Gmelin) Razafim. & B. Bremer; Salicaceae: Ludia mauritiana J.F. Gmelin, 1791; Sapindaceae: Stadmania oppositifolia Lam.; Sapotaceae: Labourdonnaisia calophylloides Bojer (Bois de natte à petites feuilles), Labourdonnaisia glauca Bojer (Bois de natte à grandes feuilles), Labourdonnaisia revoluta Bojer, Mimusops maxima (Poir.) R.E. Vaughan (Grand natte), Mimusops petiolaris Dubard, 1915 (macaque, makak, bois makak), Sideroxylon boutonianum A. DC, Sideroxylon cinereum Lam., Sideroxylon puberulum C. DC. (Manglier rouge), Sideroxylon grandiflorum A. DC. (tambalacoque, dodo tree). POPULATION: Structure and Density The present population on Mauritius appears to be stable; 14 roosting sites were reported by Nyhagen (2004), but Robin, 2007) identified 57 roosts. This latter study only looked at 24 of the 57 roosts and calculated a minimum population size of 12,000 - 16,000. It can therefore be assumed that the minimum island wide population is over 25,000 (V. Tatayah pers. comm. [in Jenkins et al. (2008k) in IUCN, 2009]). There has been a range of population estimates with the first estimate of 10,000 individuals in 1974 (Cheke and Dahl, 1981). Cheke (pers. comm. 2012 in Hutson and Racey (2013)) admits that this was a very crude estimate based on counts of a few colonies and talks with hunters and other anecdotal sources. The Mauritian government states that it has carried out counts since 2003, and data are available for 2006 and 2010. These give figures of 22-25,000 for 2006 and 49-56,000 for 2010 (Sookhareea, 2011) using a range of techniques depending on circumstances. The population figures given are for all animals: the numbers of mature individuals is estimated to be ca. 65% of the total population, hence 33,000 of an estimated population of 50,000, and 16,000 for a population of 25,000 (based on a 50% mortality in the first year, and a 30% mortality in subsequent years, with a period of at least two years to maturity and a generation time of seven years). Robin (2007) for 2007 suggests 12,000-16,000 using evening dispersal counts. The counts over the last 10 years have not been carried out in a consistent fashion and may not be directly comparable. Nevertheless, it is likely that there has been an increase in population in that time, although the scale of that increase is uncertain. An increase is likely since there has been no 68 ISSN 1990-6471 significant cyclone in that period, and major cyclones will result in heavy population losses, which will be slow to recover. The number of known roost sites has also increased from c.14 in 2003 (Nyhagen, 2004) to c. 57 (Robin, 2007; Sookhareea, 2011). A methodology has now been established for counting all known roost sites and it is hoped that regular surveys will provide data to track the population trend. In 2007 a small population (10-20 animals) was identified on Réunion (where the species formerly occurred, but has been extinct for well over 100 years). These animals can be assumed to have blown over from Mauritius (as suggested for other stray individual records since 2000). Limited erratic breeding has been demonstrated on Réunion, and so this population has not been considered for the purposes of this assessment. VIRUSES: Rhabdoviridae Of the 36 bats from Mauritius tested by Mélade et al. (2016a: 6) against Lagos bat lyssavirus and Duvenhage lyssavirus, two and 13 respectively, displayed a sero reaction. Also one out of 21 specimens tested for European bat lyssavirus 1 showed this reaction. UTILISATION: This species is locally used for sport hunting and as a source of food (R). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar, Mauritius, Réunion. Anthony et al. (2018: 12074) referred to a survey held in October 2016, which yielded a population estimate of about 62,000 individuals. Trend 2013: Stable (Hutson and Racey, 2013). 2008: Decreasing (Jenkins et al., 2008k; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) report a maximum longevity of 19.4 years. PARASITES: Hirst (1923: 980) mentions that the mite Ancystropus mulleri Kolenati was collected from P. vulgaris [= P. niger]. Figure 9. Distribution of Pteropus niger Pteropus rodricensis Dobson, 1878 *1878. Pteropus rodricensis Dobson, Catalogue of the Chiroptera of the collection of the British Museum, xxxvii, 20, 36, pl. 3, fig. 3. Publication date: June 1878. Type locality: "Africa": Mascarene Islands: Rodrigues Island [Goto Description]. Holotype: BMNH 1876.3.11.1: ad ♂, skull and alcoholic. Collected by: J. Gulliver. Presented/Donated by: The Royal Society. See Andersen (1912b: 275). Paratype: SMF 44785: juv ♀, skull and alcoholic. Collected by: J. Gulliver. Presented/Donated by: ?: Collector Unknown. (Current Combination) 1907. Pteropus mascarinus Mason, Ann. Mag. nat. Hist., ser. 7, 20 (117): 221. Publication date: 1 September 1907. Type locality: Mauritius: 15 mi (=22 km) NE Mauritius: Round island (La Ronde) [Goto Description]. Holotype: BMNH 1925.8.10.1:. TAXONOMY: See Bergmans (1991) and Simmons (2005: 343). Almeida et al. (2014: 85) place P. rodricensis in the "vampyrus" group. COMMON NAMES: Castilian (Spain): Zorro volador de la isla Rodrigues. Czech: kaloň rodrigueský, kaloň zlatý. Dutch: Rodriguez vliegende hond. English: Rodriguez Flying Fox, Rodrigues Flying Fox. French: Renard volant de Rodrigues, Renard Volant de l'île de Rodriguez. German: Rodriguez-Flughund, Rodrigues-Flugfuchs. CONSERVATION STATUS: Global Justification African Chiroptera Report 2020 Listed as Endangered (B1ac(iv)+2ac(iv) ver 3.1 (2001)) because the global population is restricted to a single location (the island of Rodrigues), and has a extent of occurrence (EOO) and area of occupancy (AOO) that are both less than 500 km 2. The population undergoes extreme fluctuations due to severe tropical cyclones, which can cause mortalities of over 50% but the population subsequently recovers at a rate of about 12-15% a year provided there are no further severe cyclones. This fruitbat exploits both native habitats and those dominated by introduced species. Native habitats are highly fragmented. Assessment History Global 2016: EN B1ac(iv)+2ac(iv) ver 3.1 (2001) (Tatayah et al., 2017). 2008: CR B1ab(iii)c(iv) ver 3.1 (2001) (Mickleburgh et al., 2008u; IUCN, 2009). 2004: CR B1ac(iv) ver 3.1 (2001) (Mickleburgh et al., 2004q; IUCN, 2004). 2002: CR: B1+3d (Mickleburgh et al., 2002a: 22), extinct on Round Island. 1996: CR (Baillie and Groombridge, 1996). 1994: EN (Groombridge, 1994). 1993: EN (World Conservation Monitoring Centre, 1993). 1990: EN (IUCN, 1990). 1988: EN (IUCN Conservation Monitoring Centre, 1988). 1986: EN (IUCN Conservation Monitoring Centre, 1986). Regional None known. Legal Status U.S. ESA - Endangered. CITES - Appendix II. MAJOR THREATS: Tatayah et al. (2017) report that deforestation has been a serious threat to the species, especially where mature fruit trees and important roost trees were felled. Because of the deforestation of this forest buffer, any remaining patches of forest (and their roosting bats), were more susceptible to tropical cyclones ((Trewhella et al., 2005; Mickleburgh et al., 2008u). These cyclones can cause significant fluctuations in bat population size and, along with shortage of food and dehydration, are now the major current threat to the species (Powell and Wehnelt, 2003). Tatayah et al. (2017) also report that deforestation is no longer occurring since at least 2010. In the past the species was also hunted for food, however, this is now rare (Trewhella et al., 2005; Mickleburgh et al., 2008u; IUCN, 2009). Due to its habit of raiding fruit trees (mangoes, lychees, papaya etc), there is a human-wildlife conflict (Barnes, 2013; Price, 2013). A new law has been passed for the Republic of Mauritius: the Native Terrestrial Biodiversity and National Parks Act 69 (2015). Whilst the Rodrigues Fruit Bat is still protected under this law, it has provisions for declaring any species as a ‘pest’ and may allow culling of the species (in spite of it being a threatened species). An official cull of nearly 31,000 Mauritius Fruit Bat Pteropus niger was sanctioned in 2015 (Tatayah et al., 2017). Whilst the law still protects the Rodrigues Fruit Bat, there are now provisions and a precedent for culling. However, the Rodrigues Regional Assembly is an autonomous administration under the Republic of Mauritius and has the right under the Rodrigues Regional Assembly Act (2001) to pass its own laws and regulations, and it is unlikely to sanction culling of bats in the near future (Tatayah et al., 2017). CONSERVATION ACTIONS: Mickleburgh et al. (2008u) [in IUCN (2009)] report a successful captive breeding programme for this species was initiated by the Durrell Wildlife Conservation Trust. In 1995, the International Studbook contained 542 individuals distributed over 18 institutions (Carroll and Feistner, 1996: 334). With breeding populations of this bat now maintained at 46 zoos around the world (David White, studbook keeper, 2016, pers.comm in Tatayah et al. (2017)). In-situ conservation efforts have concentrated on restoration of the natural habitat, watershed protection and awareness raising among the local people through environmental education programmes (Powell and Wehnelt, 2003; Trewhella et al., 2005; Tatayah et al., 2017). GENERAL DISTRIBUTION: Pteropus rodricensis is confined to the western Indian Ocean island of Rodrigues (Mauritius). It was historically present on the island of Mauritius and Round Island (Mauritius), but is now extinct on these islands. It appears to range up to around 200 m asl. Native: Mauritius [Mauritius (main island) Regionally Extinct, Rodrigues]. DETAILED MORPHOLOGY: Baculum - Unknown Gardner et al. (2007: 196) studied the heart of several P. rodricensis specimens kept in captivity, and found it to have a normal cardiac silhouette, comparable to domestic small mammals. Carter and Adams (2013: 33) postulated that tracheal and primary bronchi calcification will occur in echolocating Artibeus jamaicensis but not in non-echolocating Syconycteris australis and Pteropus rodricensis. The calcification would be 70 ISSN 1990-6471 needed to counteract the compressive forces executed by the flight muscles on the respiratory tract, as there is a sychronisation between wingbeat and respiration, and the latter is linked with the production of echolocation calls. However, to their surprise, they found that the tracheae and primary bronchi of P. rodricensis were also heavily calcified. MOLECULAR BIOLOGY: O'Brien and Hayden (2004: 144) report that, notwithstanding the reduction of the population to less than 70 individuals in the 1970's, the species has still retained some genetic diversity. DIET: Courts (1998: 189) reports that captive animals catch insects flying by, and that these were readily eaten. The bats also ate mealworms (Tenebrio molitor) and waxmoth larvae (Galeria mellonella). Blood glucose - Heard and Whittier (1997) found blood glucose concentrations within the normal mammalian range. POPULATION: Structure and Density:- Until around 1916, the species was reported to be abundant on Rodrigues, and even in 1955 large numbers (about 500) still roosted in tamarinds (Mickleburgh et al., 2002a). In 1965 there were fewer bats, but the species was still common. There was a marked decline in the 1970's, and following Cyclone Celine II in 1979, the population was reduced to around 70 bats. By 1980 the population had recovered to between 200 and 250 animals (Carroll and Mace, 1988), and at the end of February 1990, the population was estimated to be greater than 1,000 bats (Mickleburgh et al., 2002a). The population had recovered to around 5,076 bats (Powell and Wehnelt, 2003). However, the impact of cyclone Kalunde in March 2003 appears to have reduced the population to around 4,000 animals (Anonymus, 2006), although Caceres (2010: 19) reports that the current population is estimated at about 5,500 individuals. Price (2013) report that Island-wide bat counts are conducted three times a year at the nine major (‘permanent’, older) roosts and up to nine smaller (‘temporary’, more recent) roosts. In 2016, the population had grown to ca. 20,000 individuals. It is notable that some of the more recent roosts have significantly more bats than the traditional roosts, and this may be an indication of bats recolonizing habitat where they have been extirpated in the past or where vegetation has recovered. Trend:- 2016: Increasing (Tatayah et al., 2017). 2008: Increasing (Mickleburgh et al., 2008u; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 28 years. ACTIVITY AND BEHAVIOUR: For a selection of sounds, check http://macaulaylibrary.org/search?taxon=Pteropus rodricensis&taxon_id=11036948&taxon_rank_id= 67&tab=audio Riccucci (2016: 94) observed several different behaviours: "play chase" by immature bats flying to one location and rapidly leaving; "play wrestle", which involves close belly contact between individuals, with restrained biting on the neck. He also noticed that pairs or groups of bats (mostly females) wrestle together in long play sessions. REPRODUCTION AND ONTOGENY: Kunz et al. (1994: 691) report on allomaternal care, or epimeletic (care-giving) behaviour, exhibited by one female toward another, before, during. and after giving birth. Szekely et al. (2015: Suppl.) report a gestation period of 198 days. The young then weights 45 g (adult: 350 g). MATING: McCracken and Wilkinson (2000: 339) refer to Carroll and Mace (1988), who indicated that P. rodricensis forms harem groups consisting of one male and up to eight females. Riccucci (2011 141) suggests that male-male mounting behaviour is a way to assert dominance. PARASITES: BACTERIA Gram-positive bacterium - Heard et al. (1979) recorded the following gram-positive bacteria from Pteropus rodricensis - Actinomycetis, Bacillus spp., Corynebacterium spp., Enterococcis spp., Lactobacillus spp., Staphylococcus aureus, Staphylococcus spp., Streptococcus spp. hemolytic, Streptococus sp Group D and Streptococcus sp. Gram-negative bacterium - Heard et al. (1979) recorded Bacteroides spp., Escherichia coli, Klebsiella oxytoca, Klebsiella pneumonia, Morganella morganii, Proteus mirabilis, and Proteus spp. Helmick et al. (2004: 88) reported a Pasteurella from one captive bat. FUNGI Heard et al. (1979) recorded an unspecified yeast. African Chiroptera Report 2020 71 UTILISATION: In the past the species was also hunted for food, however, this is now rare (Trewhella et al., 2005). Mahomoodally et al. (2019: 6) indicate that fat from P. rodricensis has an ethno-pharmacological use: it is melted and applied on the scalp against hair loss. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Mauritius. Figure 10. Distribution of Pteropus rodricensis Pteropus rufus E. Geoffroy St.-Hilaire, 1803 *1803. Pteropus rufus E. Geoffroy Saint-Hilaire, Catalogue des Mammifères du Muséum national d'Histoire naturelle, Paris, 47. Type locality: Madagascar: North and Central Madagascar [Goto Description]. - Comments: Type locality originally "Madagascar", restricted to North and Central Madagascar by Andersen (1908b: 367) (See Peterson et al., 1995: 14). Andersen (1912b: viii) indicates that the type was once in the Paris Museum, but is probably no longer in existence. On page 206, he also mentioned that the specimen was sent by "le citoyen Macé, naturaliste" (nr 90 of Geoffroy's catalog). (Current Combination) 1810. Pteropus Edwardsii E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 15: 92. Type locality: Madagascar: "Madagascar". - Comments: Renaming of P. rufus E. Geoffroy SaintHilaire, 1803 (see Andersen, 1906c: 206; Allen, 1939a: 60). Mentioned as edwardii by Smith (1833: 52) and as edwarsi by Koopman (1993a). Description is based on the same specimen as Pteropus rufus Geoffroy, 1803. 1816. Pteropus madagascariensis Oken, Lehrbuch Naturgeschichte, Jena, 3 (2): 936. Type locality: Madagascar: "Madagascar". - Comments: Renaming of P. edwardsii E. Geoffroy Saint-Hilaire, 1810 (see Andersen, 1912b: 207; Allen, 1939a: 60). 1825. Pteropus phaiops Temminck, Monogr. Mamm, 1: 178. Type locality: Madagascar: "Madagascar". - Comments: Syntypes: RMNH Jentink Cat. Syst., p. 146 [under Pt. edwardsi] c (skull not removed), d (skull removed); Cat. Ost., p. 259, c. 1838. Pteropus phæops Oken, Allg. Naturgesch, 7 (2): 990. Type locality: Madagascar. 1908. Pteropus rufus princeps K. Andersen, Ann. Mag. nat. Hist., ser. 8, 2 (10): 367. Publication date: 1 October 1908. Type locality: Madagascar: Tuléar province: Fort Dauphin [25 02 S 47 00 E] [Goto Description]. Holotype: BMNH 1891.11.30.10: ad ♂, skull and alcoholic. Collected by: M. Cloisel. See Andersen (1912b: 14), Peterson et al. (1995: 14). ? Pteropus rufus rufus: (Name Combination) TAXONOMY: See Bergmans (1991) and Simmons (2005: 343 344). Commonly cited from "E. Geoffroy, 1803. Cat. Mamm. Mus. Nat. Hist. Nat. Paris, p. 47", but this work was never published, which would make this a nomen nudum. However, Opinion 2005 (Case 3022) fixed the authorship and date of this name to E. Geoffroy St.-Hilaire (1803) by (see Anonymus, 2002b: 154). O'Brien et al. (2009) suggested that P. aldabrensis, P. rufus, P. niger and P. seychellensis (including comorensis) should be considered geographical forms of a single species P. giganteus (Brünnich, 1782) from the Indian subcontinent, Maldives and Andaman archipelagos. Almeida et al. (2014: 85) place P. rufus in the "vampyrus" group. 72 ISSN 1990-6471 COMMON NAMES: Castilian (Spain): Zorro volador de Madagascar. Czech: kaloň madagaskarský, upír černolícní. English: Madagascan Flying Fox, Madagascan Flying-fox, Red Flying Fox, Madagascar Fruit Bat, Madagascar Flying Fox. French: Renard volant de Madagascar, Renard volant roux, Roussette rougeâtre, Renard volant malgache. German: Madagaskar-Flugfuchs. Malagasy: Fanihy. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Gunnell et al. (2014: 3) refer to Goodman and Junkers (2013), who report on fossil material from the Andrahomana cave in Madagascar. CONSERVATION STATUS: Global Justification This species is listed as Vulnerable (VU A2acd ver 3.1 (2001)) based on the loss of over 30% in the last three generations (18 years; Pacifici et al., 2013). The species can be legally hunted and the reduction can be attributed to the increased level of hunting particularly with the introduction of shotguns and continued loss of its preferred roosting habitats (Andriafidison et al., 2008a; IUCN, 2009; Racey, 2016). Assessment History Global 2016: VU A2acd ver 3.1 (2001) (Racey, 2016). 2008: VU A2acd ver 3.1 (2001) (Andriafidison et al., 2008a; IUCN, 2009). 2004: VU A2b+3d ver 3.1 (2001) (Participants of CBSG CAMP Madagascar Mammal Worksh, 2004; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). 1993: LR/lc (World Conservation Monitoring Centre, 1993). Regional None known. Legal Status CITES - Appendix II. MAJOR THREATS: There are numerous threats to this species. It is listed as a game species under Malagasy law and can be legally hunted between the first of May and the first of September (Anonymus, 2007b: 6; Durbin, 2007; Randrianandrianina et al., 2010: 412). Only the few roosts that are located in protected areas therefore receive some protection (Racey et al., 2009). Pteropus rufus is hunted for food across Madagascar (MacKinnon et al., 2003; Rakotondravony, 2006; Rakotonandrasana and Goodman, 2007; Racey et al., 2009; Randrianandrianina et al., 2010) where it is an important subsistence food, but also of commercial importance. Although quantitative data on the supply of P. rufus for food and the impact of this harvest on colonies/populations is lacking, there is some evidence that the offtake is locally unsustainable (Racey et al., 2009). Hunting occurs at roost sites and where the bats forage, especially when they feed on trees in villages (Jenkins and Racey, 2008). There is evidence that P. rufus from Ile Sainte Marie is hunted commercially and frozen shipments are sent to the mainland port of Toamasina (Rakotonandrasana and Goodman, 2007). The conversion of the forest used by roosting bats into agriculture is another threat to P. rufus and results in the permanent loss of suitable trees which in many cases have been used for decades. Pteropus rufus feeds on a number of cultivated fruits that have a high economic value in rural Madagascar and is often subjected to persecution. Randrianandrianina et al. (2010: 413) observed that a minimum of five P. rufus specimens were caught each night, when they were feeding on kapok (Ceiba pentandra) nectar in July. For one roost in Andranomena, Oleksy et al. (2015b: 4) reported that six hunters camped for three nights per week for approximately eight months per year and captured about 20 bats per night. Oleksy et al. (2015b: 4) and Cardiff and Jenkins (2016: 97) mention that roost sites are also threatened by fire and agricultural expansion. These roost sites are particularly vulnerable as they are located in large trees located in forest fragments in savanna or scrub habitats. CONSERVATION ACTIONS: Racey (2016) who support Andriafidison et al. (2008a) [in IUCN (2009)] who report that it is listed on CITES Appendix II and is a game species under Malagasy law (Durbin, 2007) but neither of these provide any practical in situ conservation measures (Racey et al., 2009). There are a few roosts in protected areas, notably Parc National Kirindy-Mité, Parc National de Masoala, Parc National de Mananara-nord (MacKinnon et al., 2003) and Berenty Private Reserve (Long, 2002), but many of the existing parks and reserves appear to be without roosting colonies (e.g. Goodman, 1996; 1999; Alonso et al., 2002; Goodman et al., 2005a; Schmid and Alonso, 2005). The ongoing process to triple the surface of protected areas in Madagascar is providing an unprecedented opportunity to include traditional roosts in the new conservation sites. There is also significant scope for local institutions to conserve roosts and this is already occurring in some parts of Madagascar where the bats use sacred forests (Jenkins et al., 2007b) or where communities have created social contracts to protect the bats (Jenkins et al., 2007a). African Chiroptera Report 2020 GENERAL DISTRIBUTION: Pteropus rufus is endemic to Madagascar. It is one of the most widespread bat species on the island and appears to only be absent from the highly populated central highlands (MacKinnon et al., 2003). The highest density of roost sites is in coastal regions, especially from Morombe in the south-west to Antsiranana in the north (MacKinnon et al., 2003). It has also been recorded from the islands of Nosy Be, Nosy Komba, and Ile SainteMarie (Rakotonandrasana and Goodman, 2007: 6). Native: Madagascar (Bollen and Van Elsacker, 2002; MacKinnon et al., 2003; Simmons, 2005: 344; Rakotonandrasana and Goodman, 2007: 6; Long and Racey, 2007; Jenkins and Racey, 2008). BIOGEOGRAPHY: See O'Brien et al. (2009) for further details. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 54) described the baculum having a distal tip forming an arch-shaped structure and a basal portion which is lobed with a distinct central prolongation; length: 3.22, 3.86 mm, width: 3.53, 4.15 mm. In subadults, the bones are notably reduced in length and width. MOLECULAR BIOLOGY: DNA - See O'Brien et al. (2009). Karyotype - 2n = 38 and FN = 68, containing 11 pairs of metacentrics, 6 submetacentrics, 1 pair of acrocentrics; X: large submetacentric, Y: small acrocentric (see Richards et al., 2016d: 187, 189). Protein / allozyme - Unknown. HABITAT: MacKinnon et al. (2003) failed to locate any roost sites within the endemic spiny forests of southwest Madagascar. Long and Racey (2007) did not find any evidence to show that P. rufus uses the endemic plants of the spiny forest in the Mandrare region of Madagascar. Jenkins et al. (2007b: 212) indicate that P. rufus is generally found in forest fragments or linear patches of vegetation along water. HABITS: Jenkins and Racey (2008) indicate that day roosts and night feeding areas are often spatially distinct, and are over 40 km apart. ROOST: P. rufus roost in foliage (Jenkins and Racey, 2008). 73 In their 2005 survey, Jenkins et al. (2007b: 213) have found six roosting sites, with colony sizes ranging from 40 to over 1,000 animals. Three of these were protected by local taboes or ancestral traditions. Ravelomanantsoa et al. (2019: 110) indicate that P. rufus uses large trees (e.g. Cocos nucifera Arecaceae) as roost. DIET: Bollen and Van Elsacker (2002: 37) found that P. rufus at Ste. Luce, ate the fruits of 40 endemic species, and that typically one seed species was found in a dropping. Oleksy and Jones (2013: 113) reported that 110 plant species were found in its diet, of which 59 (55 %) were endemic. However, Long (2002) and Long and Racey (2007) found P. rufus at Berenty in south-east Madagascar to have a narrow dietary breadth, consisting of only 14 plant species. The single most important food source for Petropus rufus in south-east Madagascar, is the pollen of Agave sisalana a commercial introduced species, while they also eat other locally cultivated and introduced fruits (Mangifera indica, Psidium cf. cattleianum, Sclerocarya caffra, Cordia sinensis and Hylocereus sp.) and native and endemic forest species (Tamarindus indica (leaves), Celtis philippensis, Ficus megapoda, F. grevei, F. pachyclada and Grewia sp.) (Long, 2002; Long and Racey, 2007). Jenkins et al. (2007b: 212) indicate that the most important food plants include Ficus guatteriifolia, Syzigium sp., Terminalia fatrea, Uapaca thouarsii and U. littoralis. Raheriarisena (2005: 255) mentions that this species’ diet is composed of fruits (59.0 - 65.1%), flowers (17.3 - 35.0%), and leaves (6.0 - 18.0%), but also that the percentages of these elements vary with the seasonal variations of plant phenology and the species’ activity, especially its reproduction status. Bollen et al. (2004a: 603) found that P. rufus preferred to consume multi-seeded fruits. Bollen (2007: 140) lists the following food plants consumed by P. rufus: Annonaceae: Polyalthia madagascariensis Cavaco & Keraudren, 1957; Araliaceae: Polyscias sp.; Areceae: Dypsis nodifera Mart., 1849, Dypsis prestoniana Beentje, 1995; Bignoniaceae: Ophiocolea delphinensis H. Perrier, 1938; Canellaceae: Cinnamosma madagascariensis var. namoronensis H. Perrier, 1948; Combretaceae: Terminalia fatraea (Poir.) DC., 1828; Ericaceae: Vaccinium emirnense Hook, 1837; Euphorbiaceae: Uapaca ferruginea Baillon, 1858, Uapaca littoralis Denis, 1927, Uapaca thouarsii Baillon, 1858; Flacourtiaceae: 74 ISSN 1990-6471 Ludia antanosarum Capuron & Sleumer, 1972, Scolopia orientalis Sleumer, 1972; Grossulariaceae: Brexia sp.; Lauraceae: Beilschmiedia madagascariensis (Baillon) Kostermans, 1952, Ocotea sp.; Liliaceae: Dracaena reflexa var. nervosa H. Perrier, 1936; Loganiaceae: Anthocleista longifolia (Lam.) Boiteau, 1973, Anthocleista madagascariensis, Baker, 1882, Bakerella ambongoensis Balle, 1964, Bakerella sp.; Monimiaceae: Tambourissa purpurea (Tul.) A. DC, 1868; Moraceae: Ficus baroni Baker, 1883, Ficus guatteriifolia Baker, 1890, Ficus pyrifolia Burm. f., 1768; Myrtaceae: Syzygium sp.2; Rubiaceae: Canthium variistipula (Arènes ex Cavaco), Ixora sp., Mapouria aegialodes Bremekamp, 1963, Mapouria sp., Hyperacanthus mandenensis Rakotonas. & A.P. Davis, 2004, Tricalysia cf. cryptocalyx Baker, 1882, Vepris elliotii (Radlk.) I. Verdoorn, 1926; Sapotaceae: Sideroxylon beguei var. saboureani, Sarcolaena multiflora Thouars, 1805. Ravelomanantsoa et al. (2019: 110) reported that fruits of Ziziphus jujuba Mill.(Jujube, Red dade, chinese dade - Rhamnaceae) are consumed. Oleksy et al. (2015a: 678) followed 15 radiotracked animals (nine ♂♂ and six ♀♀ females) between July and September 2012 and found that females had larger home ranges and core foraging areas, which might be explained by the higher energy requirements of the females during the gestation period. The bats showed a strong preference for overgrown sisal (Agave sisalana) plantations, and also ate large quantities of figs (Ficus grevei). Oleksy et al. (2017: 1) found seeds of Ficus polita, F. grevei and F. lutea in bat faeces. They found that the seeds that passed through the bat's digestive track had an increased germination success. Based on stable isotope analyses, Reuter et al. (2016c) found that Pteropus rufus, Eidolon dupreanum and Rousettus madagascariensis have broadly overlapping diets, although there are differences between P. rufus and E. dupreanum. Furthermore, they found that the diet shifted when these two species occurred sympatrically. Raharimihaja et al. (2016: 9512) tested a number of methods to deter P. rufus from feeding on commercial fruits (Litchi chinensis). They found that an organic product made from dried blood and vegetable oil (Plantskydd®) with a taste and odour aimed at deterring mammal feeding, gave the best results. PREDATORS: Mikula et al. (2016: Supplemental data) mention the Madagascar harrier-hawk (Polyboroides radiates (Scopoli, 1786)) as diurnal avian predator. POPULATION: Structure and Density:- Roosts are conspicuous and noisy but the bats are often tightly clustered making abundance estimates at the larger sites difficult. MacKinnon et al. (2003) reported that colony size ranged from 10 to 5,000 animals with a median of 400. They found that nationally 17.5% of 154 roosts had been abandoned in the last ten years and desertion rates were much higher (70%) in the central highlands. The estimated national population size in 2000 was 300,000 (MacKinnon et al., 2003). In December 2005, Rahaingodrahety et al. (2008) recorded colony sizes of 900 at Berenty Private Reserve, 412 at Amborabao and 54 at Sainte Luce. Based on tooth-derived estimated of adult survival, Brook et al. (2019a: 165) showed that even with a perfect juvenile annual survival (100 %) stable population trajectories for P. rufus are not possible. Trend:- 2016: Decreasing (Racey, 2016). 2008: Decreasing (Andriafidison et al., 2008a; IUCN, 2009). ACTIVITY AND BEHAVIOUR: For a selection of sounds, check http://macaulaylibrary.org/search?taxon=Pteropus rufus&taxon_id=11067862&taxon_rank_id=67&ta b=audio REPRODUCTION AND ONTOGENY: Kaudern (1914: 11 - 12) reported that he was unable to shoot any pregnant female or females with young during his travels on Madagascar, which took place in the dry season (May to September), so he assumed they had their reproductive season in Spring or Summer [as P. edwardsi]. Rand (1935) [in Fox et al. (2009: 272)] found two large foetuses in each of the three female specimens that were available to him. PARASITES: BACTERIA Gram-positive bacterium - Reviewed in McCoy (1974), who listed Enterococcus spp. and Staphylococcus spp. Gram-negative bacterium - Mayoux et al. (1971: 29) and McCoy (1974) listed Citrobacter sp., Enterobacter sp., Escherichia coli, Salmonella typhi, S. typhimurium, Salmonella spp., and Shigella flexneri. African Chiroptera Report 2020 Gomard et al. (2016: 5) added Leptospira to this list (see also Dietrich et al., 2018a: 3). PROTOZOA Ranaivoson et al. (2019: 1) were the first to report Babesia sp. (order Piroplasmida; family Babesiidae), an erythrocythic protozoan) from Madagascan bats (and the first Pteropodid infection). They examined 203 P. rufus, of which nine (all males) were positive for babesial infection, whereas none of the 203 bats carried any ectoparasites. VIRUSES: Coulanges et al. (1974) [in Fontenille et al. (1989: 239)] found antibodies against West-Nile virus in 11.4 % of the 94 bats from the Anjiro region they examined. Coronaviridae - Coronaviruses Thirteen novel strains of betacoronaviruses were identified in Madagascar in 2010 (Razanajatovo et al., 2015). Filoviridae Ebola virus - Brook et al. (2019b: 1004) reported the presence of reliable reactive antibodies to tested antigens in serum from P. rufus. However, they also mention (p. 1010) that no filoviruses (nor henipaviruses) have been identified from Madagascan bats (either via live viruses or RNA). Paramyxoviridae 3 out of 20 Madagascan specimens tested by Mélade et al. (2016b: 4) were positive for paramyxoviruses. Henipavirus Nipah (NiV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) found an overall contamination of 2.3%. Hendra (HeV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) found an overall contamination of 2.3%. Brook et al. (2019b: 1004) reported the presence of reliable reactive antibodies to tested antigens for HeV in serum. Tioman (TiV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) found an overall contamination of 2.3%. Rhabdoviridae – Rabies like viruses Lyssavirus - Rabies related viruses Andriamandimby et al. (2013: 3) reported on one P. rufus from Madagascar, which tested positive to antibodies for European Bat Lyssavirus type 1 (EBLV-1). Of the 11 bats tested by Mélade et al. (2016a: 6) for Duvenhage lyssavirus, five showed a positive reaction. 75 UTILISATION: Pteropus rufus being the largest bat is the main source of bat bushmeat in Madagascar (Jenkins and Racey, 2008). In many regions nets are erected inside or on the periphery of roosts to intercept flying bats (Jenkins et al., 2007b; Jenkins and Racey, 2008). In southern Madagascar P. rufus is netted when it feeds on sisal (Agave sisalana) plants at night and at least some of the bats are sold locally (Rahaingodrahety, 2007). Firearms (shotguns) are also used to hunt P. rufus in many areas (Jenkins and Racey, 2008). In eastern Madagascar individual trees with roosting bats are felled by hunters who club the fallen animals with sticks (MacKinnon et al., 2003). Pteropus rufus is hunted in the Daraina area in northeastern Madagascar (Rakotondravony, 2006) where roosts of over 1,000 individuals have been reported from littoral forest (Jenkins and Racey, 2008). Although the people living around the forests refrain from eating the bats, the colonies are reportedly subject to frequent hunting by local immigrants (Rakotondravony, 2006). In Makira, P. rufus is eaten widely by local communities and at $1.5 US/kg was the most expensive meat (Golden, 2005). In the Mahavavy-Kinkony area of western Madagascar, regular hunting at a P. rufus roost was reported by Rakotoarivelo and Randrianandrianina (2007); hunting teams of up to eight men visited the roost on a regular basis and caught up to 100 bats on each occasion. These were sold around villages and to local restaurants for $0.5 US each. On Nosy Boraha, P. rufus is regularly exploited from at least one roost and is served seasonally in restaurants in the main town on the island (Rakotonandrasana and Goodman, 2007). One hunter also reported that frozen shipments of P. rufus are sent to the mainland port of Toamasina (Rakotonandrasana and Goodman, 2007). At a small restaurant near Mahajanga in 2000, at least 30 P. rufus were reportedly sold daily, although it is not clear whether this figure is monthly average estimate or reflects a seasonal peak in availability (Racey et al., 2009). In the sisal plantations surrounding Berenty Private Reserve, hunters can expect to catch 8 - 12 individuals per week between September and May and 25 - 30 between June and August, where they are sold discretely in villages around the reserve for $ 0.7 - 1.0 US (Rahaingodrahety, 2007) therefore a single hunter could catch between 164 and 192 bats per year. Pteropus rufus and Eidolon dupreanum appear to be the only species sold in restaurants, where they are either served individually, complete with head and wings, jointed, or are diced up into small pieces and accompanied by rice (Jenkins and Racey, 2008). Rakotondravony (2006) considered hunting of P. rufus a minor threat in northern Madagascar, while 76 ISSN 1990-6471 Golden (2005) reported large annual harvests in the Makira forest and Maroantsetra were unsustainable. See also Goodman et al. (2008d). Golden and Comaroff (2015: 5) indicate that flying foxes and other fruit bats (including P. rufus) are frequently the subject of food taboos, which they link to the fact that these bats are carriers of virulent zoonotic diseases. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Comoros, Madagascar. Figure 11. Distribution of Pteropus rufus Pteropus seychellensis A. Milne-Edwards, 1877 *1877. Pteropus seychellensis A. Milne-Edwards, Bull. Sci. Soc. Philom. Paris, sér. 7, 2: 221. Type locality: Seychelles: Mahe Island. Lectotype: MNHN ZM-MO-1878-1110 - 35A-157: ad ♂, mounted skin and skull. Collected by: M. Lantz. Paralectotype: MNHN ZM-MO1878-1106: Collected by: M. Lantz. Rode (1941: 230): nr 157b. Paralectotype: MNHN ZM-MO-1878-1107 - 35B: ♂, mounted skin (skull not removed). Collected by: M. Lantz. Rode (1941: 230): nr 157a. Paralectotype: MNHN ZM-MO-1878-1108: Collected by: M. Lantz. Rode (1941: 230): nr 157c. Paralectotype: MNHN ZM-MO-1878-1109 - 35C: Collected by: M. Lantz. Rode (1941: 230): nr 157d. Paralectotype: MNHN ZM-MO1878-1111: ad ♀, skin and skull. Collected by: M. Lantz. Paralectotype: MNHN ZM-MO1878-1112: ad ♀, skin and skull. Collected by: M. Lantz. - Comments: Bergmans (1991: 151) indicated that the lectotype (MNHN CG 1878-1110), as well as most of the other specimens on which Milne-Edwards based his description, were in fact collected by Lantz on Marianne Island, and not on Mahé Island. (Current Combination) 1880. Pteropus comorensis Wallace, Island Life, 400. Type locality: Comoros: "Comoro Islands". - Comments: Allen (1939a: 60) and Simmons (2005) mention J.M. Nicoll, 1908 as author, since the original description by Wallace (1880) is actually a nomen nudum. 1908. Pteropus comorensis Nicoll, Three voyages of a naturalist being an account of many littleknown islands in three oceans visited by the "Valhalla" R.Y.S. - Witherby & Co. London, 87. Type locality: Mayotte: Buzi Island [0 - 600 ft] [Goto Description]. Paratype: SMF 44784: ♂, skull and alcoholic. Collected by: Michael John Nicoll; collection date: 1906. Presented/Donated by: ?: Collector Unknown. Syntype: BMNH 1906.6.3.14: ad ♂, skin and skull. Collected by: Michael John Nicoll; collection date: 2 March 1906. Presented/Donated by: Earl of Crawford. Syntype: BMNH 1906.6.3.15: ad ♂, skin and skull. Collected by: Michael John Nicoll; collection date: 2 March 1906. Presented/Donated by: Earl of Crawford. Syntype: BMNH 1906.6.3.16: ad ♂, skin and skull. Collected by: Michael John Nicoll; collection date: 25 February 1906. Presented/Donated by: Earl of Crawford. - Comments: Nicoll (1908: 88) mentions that the bats were shot in the evening when returning to the ship, and before (p. 87) he mentioned that every evening the bats were flying from the little island (= "Buzi Island") to the main island (= Mayotte). 2016. Pteropus seychellensis comoroensis: Amador, Moyers Arévalo, Almeida, Catalano and Giannini, J. mamm. Evolut., 25 (1): Suppl. (for 2018). Publication date: 24 November 2016. (Lapsus) ? Pteropus seychellensis comorensis: (Name Combination) ? Pteropus seychellensis seychellensis: (Name Combination) African Chiroptera Report 2020 TAXONOMY: Hill (1971a: 574) included aldabrensis, but we follow Bergmans (1991) and Simmons (2005: 344) who considers this to be a separate species. It seems probable that the two recognized subspecies Pteropus seychellensis seychellensis (from the Seychelles) and Pteropus seychellensis comorensis (from the Comoros Islands and Mafia Island) represent distinct taxa at the species level. In addition, the population of Pteropus seychellensis comorensis from Mafia island (Tanzania) might represent a species distinct from the populations of this bat recorded from the Comoros Islands (Mickleburgh et al., 2008di; IUCN, 2009). O'Brien et al. (2009) suggested that P. aldabrensis, P. rufus, P. niger and P. seychellensis (including comorensis) should be considered geographical forms of a single species P. giganteus (Brünnich, 1782) from the Indian subcontinent, Maldives and Andaman archipelagos. O'Brien et al. (2009) show that P. s. seychellensis is more closely related to P. niger from Mauritius, than to P. s. comorensis. Further, based on cytochrome b and 12S results, there appears to be no notable genetic differentiation in populations of P. s. comorensis from Grande Comore and Mayotte, suggesting genetic exchange. Chan et al. (2011: 7) found that P. s. comorensis and P. rufus alleles differed by as few as two basepairs. Based on this mitochondrial haplotype network, P. s. seychellensis was more closely related to P. niger and these two were more closely related to P. rufus than to P. s. comorensis. Almeida et al. (2014: 85) place P. seychellensis in the "vampyrus" group. COMMON NAMES: Castilian (Spain): Zorro volador de las Seychelles. Comorian: ndema, ndrema. Czech: kaloň seychelský. English: Seychelles Flying Fox, Seychelles Fruit Bat. French: Renard volant des Seychelles, Roussette des Seychelles. German: Seychellen-Flugfuchs. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008di; IUCN, 2009; Goodman, 2017k). 77 Assessment History Global 2016: LC ver 3.1 (2001) (Bergmans et al., 2017b). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008di; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cs; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). 1993: LR/lc (World Conservation Monitoring Centre, 1993); VU [assessed as P. s. aldabrensis] (World Conservation Monitoring Centre, 1993). 1990: VU [assessed as P. s. aldabrensis] (IUCN, 1990). 1988: VU [assessed as P. s. aldabrensis] (IUCN Conservation Monitoring Centre, 1988). Regional None known. Legal Status CITES - Appendix II. MAJOR THREATS: Overall, there currently appear to be no major threats to the species. There is some limited hunting for food on a few islands in the species range (e.g. Mahé and Praslin in the Seychelles) however, this does not appear to be significantly impacting the population at present. Additional localised threats include mortality through collision with power lines, and possible persecution as a pest of fruit crops (Mickleburgh et al., 2008di; IUCN, 2009; Bergmans et al., 2017b). However, Sagot and Chaverri (2015: 1670) mention the following threats: Roost loss or disturbance, habitat degradation or loss, introduced predators. Due to the numerous amount of bats around Mutsamudu (Anjouan, Coromo islands), Middleton and Hume (2010: 132) report that this bat is not a part of the people's diet. CONSERVATION ACTIONS: Mickleburgh et al. (2008di) [in IUCN (2009)] report that a large population (2,000 to 3,000 bats) of this species is present in the Morne Seychellois National Park on Mahé, Seychelles (Nicoll and Racey, 1981). There is a general need to continue monitoring activities for this species in order to detect any possible declines and to determine if there are any movements between populations. Studies are underway to better determine the taxonomic status of the population of bats from Mafia Island (Pteropus seychellensis comorensis) largely to understand if this population should be recognised at the species level distinct from P. seychellensis. The population of Mafia Island may be threatened, and an assessment of the situation on this island and of the possibilities for the species’ protection there is highly desirable (Mickleburgh et al., 2002a). 78 ISSN 1990-6471 Based on their study of mitochondrial and microsatellite markers, Ibouroi et al. (2018a: 1425) conclude that the populations of P. seychellensis occurring on the Comoro Islands can be treated as one management unit for conservation purposes (contrary to what they found for P. livingstonii). GENERAL DISTRIBUTION: Pteropus seychellensis has been recorded from the Seychelles, the Union of the Comoro and Tanzania. In the Seychelles it is present on a number of islands including Mahé, Praslin, La Digue, and Silhouette. Gerlach (2004) mentions that it has recently been recorded roosting on several islands where its presence was previously unconfirmed. In the Comoros it is known from the islands of Anjouan, Grand Comore and Moheli. In Tanzania, it is restricted to Mafia island, possibly Zanzibar, although Simmons (2005) considers this record extremely dubious. Ibouroi et al. (2018b: 2395) indicated that its distribution is negatively correlated to natural forest across the entire area on the Comoro Islands, but that its foraging areas were positively correlated with natural forests and high-elevation areas. Cheke (2011: 59) argues this species (as P. comorensis) was introduced on Mafia island by Comorians, rather than having arrived on the island by itself, which he supports by indicating that the bat was not noticed (perhaps not present) until the 1930s, and that no bones were found in middens (p. 65). Known across an elevational range on Grand Comore from sea level to 1,000 m, on Anjouan from sea level to 760 m, on Mohéli from sealevel to 500 m, and Mayotte from sea level to 250 m (Cheke and Dahl, 1981; Mickleburgh et al., 1992; Trewhella et al., 2001; Goodman et al., 2010c: 123). Native: Comoros (Goodman et al., 2010c: 123 as P. s. comorensis); Mayotte; Seychelles; Tanzania. BIOGEOGRAPHY: See O'Brien et al. (2009) and Goodman et al. (2010c: 123) for further details. MOLECULAR BIOLOGY: DNA - See O'Brien et al. (2009). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: Goodman et al. (2010c: 123) indicate that P. s. comorensis is adapted to the anthropgenic transformation of the former natural habitats of the different islands, including the widespread plantations of fruit trees. They occur in a variety of habitats from urban settings, agricultural zones, to remnant forest patches. HABITS: See Cheke and Dahl (1981); Trewhella et al. (1995); Trewhella et al. (2001) and Louette (2004). Gerlach (2003: 79) reports on P. seychellensis specimens flying low over the sea surface, and dipping their belly fur in the water. He postulates that this is possibly a strategy to remove parasites. The risk of falling in sea was minimised as the bats only dipped in exceptionally calm weather. Based on wing morphology, Lindhe Norberg et al. (2000) classified this species (as P. seychellensis comorensis) as "non-soaring". ROOST: Racey and Nicoll (1984) [in O'Brien (2011: 262)] indicate that P. seychellensis roosts in tall trees (Casuarina or Albizzia). Ibouroi et al. (2018b: Suppl.) reported the following roost trees being used by P.seychellensis comorensis on the Comoro Islands: Anacardiaceae: Mangifera indica, Brachylaena ramiflora; Bombacaceae: Adansonia digitata, Ceiba pentandra; Caesalpiniaceae: Tamarindus indica Combretaceae: Terminalia catappa Fabaceae: Pterocarpus indicus; Lauraceae: Ocotea comorensis; Leeaceae: Leea guineensis; Loganiaceae: Nuxia pseudodenta; Mimosaceae: Albizia glaberrima, Albizia lebbeck; Moraceae: Artocarpus heterophyllus, Ficus antandronarum, Ficus lutea, Ficus pirifolia; Myrtaceae: Eucalyptus sp., Eugenia jambolana; Sapindaceae: Litchi chinensis. DIET: Pteropus seychellensis eats 27 species of plant, including 17 fruits (Racey and Nicoll, 1984). Trewhella et al. (2001: 141) reported the following food items (with known items in bold and suspected in plain text): Anacardiaceae: Anacardium occidentale L. (cashew), Mangifera indica L. (mango); Annonaceae: Annona muricata L. (soursop), Annona reticulata L. (custard apple), Annona squamosa L. (sugar apple or sweetops); Aphloiaceae: Aphloia theaeformis [= Aphloia theiformis (Vahl) Benn.]; Arecaceae: Cocos nucifera L. (coconut tree); Cannabaceae: Trema orientalis (L.) Blume (charcoal-tree, Indian charcoal-tree,pigeon wood or Oriental trema); Caricaceae: Carica papaya L. (papaya); Combretaceae: Terminalia catappa L. (Indianalmond); Fabaceae: Erythrina sp. (coral trees); African Chiroptera Report 2020 Gentianaceae: Anthocleista grandiflora L. (forest fever tree); Lamiaceae: Vitex sp. (chaste tree); Loganiaceae: Strychnos spinosa Lam. (Natal orange, spiny orange or green monkey orange); Malvaceae: Adansonia digitata L. (baobab), Adansonia madagascariensis Baill. (baobab), Ceiba pentandra (L.) Gaertn. (kapok tree); Monimiaceae: Tambourissa anjouanensis, Tambourissa leptophylla (Tul.) A. DC., Tambourissa mohelienses Lorence, Tambourissa paradoxa Perkins; Moraceae: Artocarpus altilis (Parkinson) Fosberg (breadfruit), Artocarpus heterophyllus Lam. (jackfruit or jack tree), Ficus sp.; Musaceae: Musa sapientum L. (banana or plantane); Myrtaceae: Eugenia jambos L. [= Syzygium jambos L. (Alston)] (rose apple), Psidium cattleyanum Sabine (Cattley guava, strawberry guava or cherry guava), Psidium guajava L. (common guava), Syzygium cuminii (L.) Skeels (Java plum); Oxalidaceae: Averrhoa bilimbi L. (bilimbi, cucumber tree or tree sorrel); Rubiaceae: Psychotria aff. aledjoensis De Wild. [= Psychotria mannii Hiern, 1877]; Rutaceae: Citrus x aurantifolia (Christm.) Swingle (key lime), Citrus x aurantium L. (bitter orange, seville orange, sour orange, bigarade orange, or marmalade orange), Citrus reticulata Blanco, 1837 (mandarine orange), Citrus x sinensis (L.) Osbeck (orange). POPULATION: Structure O'Brien et al. (2009), based on cytochrome b and 12S results, indicated that there appeared to be no notable genetic differentiation in populations of P. s. comorensis from Grande Comore and Mayotte, suggesting genetic exchange. Density This species is widespread throughout the Seychelles, with some colonies consisting of up to 300 individuals. It is very common in villages and towns (Goodman, 2007). The population was estimated to be around 10,000 individuals on the Seychelles in 1979, and was believed to be close to this level in 2004 although adequate censuses have not been completed for all islands (Gerlach, 2004). 79 and play-chasing. He indicated that this behaviour was not limited to young bats, but saw this also among adults. Trewhella et al. (2001: 135) indicate that the sympatric P. livingstonii is more diurnally active (leaving their roosts at 11 a.m.) than P. seychellensis comorensis (leaving its roosts not before 3 p.m.), and is the dominant species during their encounters. REPRODUCTION AND ONTOGENY: See Trewhella et al. (1995) and Reinhardt and Jacobs (2006). MATING: McCracken and Wilkinson (2000: 339) indicate that Cheke and Dahl (1981) reported that P. seychellensis forms harem groups. PARASITES: BACTERIA Dietrich et al. (2018a: 3) reported the presence of bacteria of the genus Leptospira. VIRUSES: Astroviridae Hoarau et al. (2018: 1) tested 21 bats from the Comoros, but none of them tested positive for this group of viruses. Rhabdoviridae Four of 40 tested bats from Mayé showed a seroreaction against Lagos bat lyssavirus (LBV) (Mélade et al., 2016a: 6). Six against Duvenhage lyssavirus (DUVV), and one out of 17 tested, against European bat lyssavirus 1 (EBL-1). For the 19 specimens tested from Mayotte, two were positive for LBV and one for DUVV. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Comoros, Mayotte, Seychelles, Tanzania. On the Comoro Islands, Ibouroi et al. (2018b: 2406) observed 11,898 bats at 59 different sites, with colony sizes varying between 11 and 742 individuals (average: 184). Trend 2016: Stable (Bergmans et al., 2017b). 2008: Stable (Mickleburgh et al., 2008di; IUCN, 2009). ACTIVITY AND BEHAVIOUR: Riccucci (2016: 93) reported on play behaviour on Praslin Island, which included play-fighting, tussle, Figure 12. Distribution of Pteropus seychellensis 80 ISSN 1990-6471 Pteropus subniger (Kerr, 1792) *1792. Vespertilio vampyrus subniger Kerr, in: Linnaeus, Animal Kingdom, 1 (1): xvii, 91. Publication date: February 1792. Type locality: Réunion: "Réunion island". - Comments: Andersen (1912b: viii) mentions that the type was originally in the Cabinet Réaumur, but is probably no longer in existence. 1802. Spectrum rubidum Daudin, in: Buffon, Histoire Naturelle des Quadrupèdes, Didot ed, 14: 188. Type locality: Mauritius: Reunion. - Comments: Type locality fixed by Andersen (1912b: 169) since the name was proposed for Buffon's Rougette, which was based on a specimen sent by de la Nux from Reunion. The year of publication might not be correct (see Allen, 1939a: 61). 1803. Pteropus fuscus: Desmarest, Nouveau Dictionnaire d'Histoire Naturelle, 19: 544. Type locality: Réunion: "Réunion Island". - Comments: Not of E. Geoffroy Saint-Hillaire, 1803, see Allen (1939a: 61). 1803. Pteropus ruber E. Geoffroy Saint-Hilaire, Catalogue des Mammifères du Muséum national d'Histoire naturelle, Paris, 48. Type locality: Mauritius: Réunion [Goto Description]. Comments: Originally "Madagascar", but corrected to Réunion by Andersen (1912b: 169). Andersen also indicates that the type specimen is not in the Paris Museum. 1810. Pteropus rubricollis E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 15: 93. Type locality: Réunion: "Réunion Island". - Comments: One specimen sent by Mr. de la Nux: see Geoffroy Saint-Hilaire (1810a: 94). Andersen (1912b: 169) indicates that this is the same specimen as used by Brisson (1756) for his description of "Pteropus collo rubro" and by Buffon (1763) for his "Rougette". 1814. Pteropus torquatus G. Fischer, Zoognosia, 3: 553. - Comments: This is a renaming and redescription of Brisson's "Pt. collo rubro" (see Andersen, 1912b: 169). 1815. Pteropus collarisIlliger, Abh. K. Preuss. Akad. Wiss. Berlin, 78, 84. Type locality: Mauritius: Réunion: Type locality originally "E. African Islands", but restricted to Réunion by Andersen (1912b: 169). - Comments: "According to Andersen (1912b: 169), it looks like the publication was part of series dated 1804-1811, but actually appeared in 1815 in the section on E. African Islands." Simmons (pers. comm.). 1837. Pteropus vulgaris Temminck, Monogr. Mamm, 2: 74, pl. 38. ? Pteropus subniger: (Name Combination, Current Combination) TAXONOMY: See Bergmans (1991) and Simmons (2005: 344). Almeida et al. (2014: 83) indicate that P. subniger could not be tested, but tentatively placed in an "incertae sedis" group. COMMON NAMES: Castilian (Spain): Zorro volador oscuro de Mauricio. Czech: kaloň maskarénský, upír obecný, upír čerwenokrký, kaloň obojkový. English: Dark Flying Fox, Lesser Mascarene Flying Fox, Lesser Mascarene Flying-fox. French: Petit Renard volant de Mascareignes, Roussette foncée, Rougette, Petit renard volant de Mascarene, Roussette à collet rouge. German: Kleiner Maskarenen-Flugfuchs. CONSERVATION STATUS: Global Justification The last authentic record of this species on Mauritius was in 1859, but it is believed to have died out between 1864 and 1873. On Réunion, no new records appeared after 1862 and it seems probable that it became extinct in the 1860s (Mickleburgh et al., 2008de; IUCN, 2009). Assessment History Global 2008: Ex ver 3.1 (2001) (Mickleburgh et al., 2008de; IUCN, 2009). 2004: EX ver 3.1 (2001) (Mickleburgh et al., 2004cu; IUCN, 2004). 1996: Ex (Baillie and Groombridge, 1996). 1994: Ex (Groombridge, 1994). 1993: Ex (World Conservation Monitoring Centre, 1993). 1990: Ex (IUCN. 1990). 1988: Ex (IUCN Conservation Monitoring Centre, 1988). 1981: probably extinct (Cheke and Dahl, 1981). Legal Status CITES - Appendix II. MAJOR THREATS: Both deforestation and local hunting are thought to have contributed to the extinction of this species. African Chiroptera Report 2020 It was thought to have lived in hollow trees (Mickleburgh et al., 2008de; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008de) [in IUCN (2009)] report that this is no longer applicable. 81 UTILISATION: Caceres (2010: 9) indicates that in the 1730s its population (on La Réunion) was large enough to trade its fat. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Mauritius, Réunion. GENERAL DISTRIBUTION: Pteropus subniger was previously known from the Mascarenes Islands (Mauritius and Réunion). It was restricted to elevations of between 1,200 and 1,600 m asl. O'Brien (2011: 286) reports it from the Tanzanian island of Mafia. Regionally extinct: Mauritius; Réunion. POPULATION: Structure and Density:- This species is now extinct. It is believed to have become extinct on Mauritius between 1864 and 1873. It is likely to have disappeared from Réunion in the 1860s. It may have lasted a little longer but is now certain to be extinct. In the 1730s it was common enough to consider for the bat oil trade (Mickleburgh et al., 2008de; IUCN, 2009). Figure 13. Distribution of Pteropus subniger Pteropus voeltzkowi Matschie, 1909 *1909. Pteropus (Spectrum) voeltzkowi Matschie, Sber. Ges. naturf. Freunde Berlin, 486. Publication date: October 1909. Type locality: Tanzania: Pemba Island: Fufuni [ca. 07 31 S 38 25 E]. Holotype: ZMB 88677: ad ♂, skin and skull. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 13 March 1903. Paratype: ZMB ad ♀. See Turni and Kock (2008). Paratype: ZMB 88678: ad ♀. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 13 March 1903. See Turni and Kock (2008). Paratype: ZMB 88679: ad ♀. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 12 March 1903. See Turni and Kock (2008). Paratype: ZMB 88680: ad ♀. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 13 March 1903. See Turni and Kock (2008). Paratype: ZMB 88681: ad ♀. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 13 March 1903. See Turni and Kock (2008). 2010. P[teropus] voletzkowi: Goodman, Weyeneth, Ibrahim, Saïd and Ruedi, Acta Chiropt., 12 (1): 133. (Lapsus) ? Pteropus voeltzkowi: (Name Combination, Current Combination) TAXONOMY: See Bergmans (1991) and Simmons (2005: 346). P. voeltzkowi together with P. livingstonii form the most distinct and ancient lineage of the western Indian Ocean Pteropus spp. (O'Brien et al., 2009). The phylogenetic analyses performed by Chan et al. (2011: 7) suggest that both are sister species. This is supported by Almeida et al. (2014: 85), who place this species in the "livingstonii" group. COMMON NAMES: Castilian (Spain): Zorro volador de Voeltzkow. Czech: kaloň pembánský, kaloň pembský. Dutch: Pemba vliegende hond. English: Pemba Flying-fox, Pemba Flying Fox. French: Renard volant de Pemba. German: Pemba-Flugfuchs. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU D2 ver 3.1 (2001)) because it is known from only a single location (Pemba Island), where ongoing active conservation activities have significantly reduced 82 ISSN 1990-6471 the plausibility of future declines of this species (Mickleburgh et al., 2008df; IUCN, 2009; Entwistle and Juma, 2016). Entwistle and Juma (2016) report that hunting has been reduced but not stopped, human disturbance at roost sites and conflict with local fruit growers are plausible threats increasing that could drive the species to a higher threatened status. Assessment History Global 2016: VU D2 ver 3.1 (2001) (Entwistle and Juma, 2016). 2008: VU D2 ver 3.1 (2001) (Mickleburgh et al., 2008df; IUCN, 2009). 2004: VU D2 ver 3.1 (2001) (Mickleburgh et al., 2004cw; IUCN, 2004). 2002: CR C2a (Mickleburgh et al., 2002a: 22). 1996: CR (Baillie and Groombridge, 1996). 1994: EN (Groombridge, 1994). 1993: EN (World Conservation Monitoring Centre, 1993). 1990: EN (IUCN, 1990). 1988: VU (IUCN Conservation Monitoring Centre, 1988). Regional None known. Legal Status CITES - Appendix II. MAJOR THREATS: Much of the natural forest habitat of this bat has been cleared or severely fragmented. The species has been hunted for food with the use of shotguns replacing traditional methods, resulting in an unsustainable use (Seehausen, 1991; Entwistle and Corp, 1997a). As of 2005, hunting had been reduced but not stopped on Pemba (Trewhella et al., 2005). An additional threat is posed by the collision of bats with overhead electric cables (Mickleburgh et al., 2008df; IUCN, 2009). In addition human disturbance at roost sites appears to have a significant impact on colony sizes (Robinson et al., 2010). A potential emerging threat may include increasing conflict with local fruit growers, given the species’ growing population and tendency to eat cultivated fruit (Robinson et al., 2010). CONSERVATION ACTIONS: Entwistle and Juma (2016) supports Mickleburgh et al. (2008df) [in IUCN (2009)] who report that ongoing awareness raising on the importance and uniqueness of the endemic fruit bat, and the need for sustainable hunting, has been undertaken through environmental education programmes (Trewhella et al., 2005; Juma, 2007). Income for the local community is being generated through bat related ecotourism activities (Juma, 2007). It has been reported from the recently gazetted Ngezi-Vumawimbi Nature Forest Reserve and Msitu Kuu Forest (Pakenham, 1984; Juma, 2007). Illegal logging and the invasive umbrella tree (Maesopsis eminii), that degrades the habitat of this bat, are being controlled within the Nature Forest Reserve (Juma, 2007). A captive breeding program was set up in 1994, starting with 18 specimens, of which 12 were housed in the Phoenix Zoo (USA), but only 4 ♂♂ and 1 ♀ survived, and Carroll and Feistner (1996: 334) were not yet able to provide information about the program's success. GENERAL DISTRIBUTION: Pteropus voeltzkowi is endemic to the island of Pemba in Tanzania, where it occurs at elevations from sea level to 45 m asl. Native: Tanzania. MOLECULAR BIOLOGY: DNA - See O'Brien et al. (2009). Karyotype - Unknown. Protein / allozyme - Unknown. ROOST: Entwistle and Corp (1997a: 137) reported roosts to be found in primary forest (n = 5), secondary forest (overgrown clove plantations) (n = 6), traditional graveyards (n = 4), mangroves (n = 1), and isolated trees (n = 3). The tree species included: Uapaca guineensis, Mangifera indica, Ficus natalensis, Ficus lutea, Syzygium jambolanum, Terminalia catappa, Adansonia digitata, Antiaris toxicaria, Erythrophleum suaveolens, Afzelia quanzensis, Parinari curatellifolia, and Blighia unijugata. DIET: Seltzer et al. (2013: table S2) provide an overview of the plant species found beneath bat feeding roosts. For P. voeltzkowi this includes Ficus sur Forssk. (Broom cluster fig - Moraceae). Entwistle and Corp (1997b: 357) analysed faecal pellets, ejecta and dropped fruits, as well as information obtained from villagers and students, and provided the following overview of the bat's diet: Mangifera indica L. (mango - Anacardiaceae [fruit]), Artocarpus altilis (Parkinson) Fosberg (breadfruit - Moraceae [fruit]), Ficus spp. (figs Moraceae [fruit]), Carica papaya L. (papaya Caricaceae [fruit]), Ceiba pentandra (L.) Gaertn. (kapok - Malvaceae [flowers/pollen]), Anacardium occidentale L. (cashew apple - Anacardiaceae [fruit]), Musa spp. (banana or plantain - Musaceae [fruit and flowers/pollen]), Psidium guajava L. (guava - Myrtaceae [fruit]), Terminalia catappa L. ('kungu' or Indian-almond - Combretaceae [fruit]), African Chiroptera Report 2020 83 Parinari curatellaefolia Planch. ex Benth. (mupundu or mobola plum - Chrysobalanaceae), Cocus nucifera L. (coconut - Arecaceae [leaves]), Eugenia aromatica (L.) Baill. [= Syzygium aromaticum (L.) Merrill & Perry (clove - Myrtaceae [fruit]), Calophyllum inophyllum L. (Alexandrian laurel - Callophyllaceae), Syzygium jambolanum Lam. [= Syzygium cumini] (L.) Skeels. (jambolana or Java plum - Myrtaceae [flowers/pollen]). They also indicated (p. 359) that cultivated species are prominently present in the diet during the months of June and July. individuals (Robinson et al., 2010) distributed across 44 active roost sites, with up to 87% of the population found at just four roost sites. Roosts ranged from solitary individuals to colonies of up to 5,040 bats. An unpublished report indicates a more recent population count of over 28,700 in 2011, with evidence of year-on-year increases over the intervening period (DCCFF, 2013). POPULATION: Structure and Density:- In the early 1990s the population of this species appears to have been reduced to a few hundred animals (Seehausen, 1991; Mickleburgh et al., 1992), with an estimate of 2,800 to 3,600 individuals in 1992, the majority being found at just two roost sites (O. Seehausen, unpubl.). In 1995 Entwistle and Corp (1997a) estimated a population of of 4,600 to 5,500 individuals, with 94% of bats found at 10 roosts (of 41 in total). Over the subsequent years surveys conducted by local teams on the island reported increasing population counts (Entwistle, 2001; Trewhella et al., 2005; Carter, 2005; Juma, 2007). Entwistle (2001: 356) reported that more recent local participation in survey work had helped to give a more accurate population estimate of ca. 6,900 bats (Trewhella et al., 2005). The population of bats has continued to increase and by the end of 2006 there were ca. 19,000 animals (Juma, 2007). A full survey in 2008 provided a population estimate between 18,200 and 22,100 DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Tanzania. Trend: 2016: Increasing (Entwistle and Juma, 2016). 2008: Increasing (Mickleburgh et al., 2008df; IUCN, 2009). Figure 14. Distribution of Pteropus voeltzkowi Subfamily Rousettinae Andersen, 1912 1912. 1912. 1970. Epomophorinae K. Andersen, Cat. Chiroptera Brit. Mus. Publication date: 23 March 1912. - Comments: Considered as a valid subfamily by Bergmans (1997: 69) who includes the tribes Epomophorini Gray, 1866 (Epomophorus, Micropteropus, Hypsignathus, Epomops, Nanonycteris), Myonycterini Lawrence and Novick, 1963 (Myonycteris, Lissonycteris, Megaloglossus), Scotonycterini Bergmans, 1997 [new tribe] (Scotonycteris, Casinycteris), Plerotini Bergmans, 1997 [new tribe] (Plerotes)? However, Almeida et al. (2016: 83) included the Epomophorine genera in the Rousettinae. Rousettinae K. Andersen, Cat. Chiroptera Brit. Mus. Publication date: 23 March 1912. Comments: Bergmans (1997: 69) recognizes the Rousettinae as a separate subfamily in which he included the tribes Rousettini Andersen, 1912 (Rousettus, Eonycteris, Eidolon) and Dobsoniini Andersen, 1912 (Dobsonia, Aproteles). Almeida et al. (2016: 83), however, provided an entirely different arrangement. Rousettina Koopman and J.K. Jones Jr., Classification of Bats. - Comments: Not included in Simmons (2005) since it was not proposed as family name, but as a subdivision within the Pteropodidiae (Simmons, pers. comm.). TAXONOMY: The subfamily was already recognized by Bergmans (1997: 69) and confirmed by the phylogenetic study by Almeida et al. (2016: 83), although their content differs. Bergmans (1997: 69) considered the Epomophorinae to be a valid separate subfamily, whereas Almeida et al. (2016: 83) included them in the Rousettinae. 84 ISSN 1990-6471 Currently recognized tribes and genera of Rousettinae (Almeida et al., 2020: 11): Rousettini Andersen, 1912 (Rousettus Gray, 1821), Eonycterini Almeida et al., 2016 (Eonycteris Dobson, 1873), Epomophorini Gray, 1866 (Epomophorus Bennett, 1836, Epomops Gray, 1870, Hypsignathus H. Allen, 1861, Nanonycteris Matschie, 1899), Myonycterini Lawrence and Novick, 1963 (Megaloglossus Pagenstecher, 1885, Myonycteris Matschie, 1899), Stenonycterini Nesi et al., 2013 (Stenonycteris Gray, 1870), Scotonycterini Bergmans, 1997 (Casinycteris Thomas, 1910, Scotonycteris Matschie, 1894), Plerotini Bergmans, 1997 (Plerotes K. Andersen, 1910). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burundi, Cameroon, Congo (Democratic Republic of the), Kenya, Liberia. TRIBE Epomophorini Gray, 1866 *1866. Epomophorini Gray, Proc. zool. Soc. Lond., 1866, I (v): 65. Publication date: May 1866. TAXONOMY: Includes the genera Epomophorus Bennett, 1836, Epomops Gray, 1870, Hypsignathus H. Allen, 1861, and Nanonycteris Matschie, 1899 (Almeida et al., 2020: 11).. Genus Epomophorus Bennett, 1836 1835. Pteropus: Ogilby, Proc. zool. Soc. Lond., 1835, III (xxxi): 100. Publication date: 9 October 1835. - Comments: Preoccupied by Pteropus Brisson, 1762. *1836. Epomophorus Bennett, Proc. zool. Soc. Lond., 1835, III (xxxiv): 149. Publication date: 12 February 1836. - Comments: Type species: Pteropus whitei var. epomophorus Bennett, 1836 (= Pteropus gambianus Ogilby, 1835). - Etymology: From the Greek "έπί", meaning upon, "ωμος", meaning shoulder, and "φόρος", meaning bearing, refering to the conspicuous tufts of hair on the shoulders of the males (see Palmer, 1904: 268; Boulay and Robbins, 1989: 4). (Current Combination) 1853. Pachysoma Temminck, Esq. zool. Côte de Guiné, 64. - Comments: Preoccupied by Pachysoma MacLeay, 1821, a genus of Coleoptera: see Boulay and Robbins (1989: 1). Not Pachysoma I. Geoffroy Saint-Hilaire, 1828. Andersen (1912b: 514) indicates that Pachysoma Temminck includes both Epomophorus and Cynopterus. 1975. Epomorphus: Höhne, Loose and Seeliger, Ann. Microbiol. (Inst. Pasteur), 126 A: 503. (Lapsus) 1978. Epomophevus: Arata and Johnson, in: Pattijn, Ebola Virus Haemorrhagic Fever, 135. (Lapsus) 2014. Epimorphosis: Walldorf and Mehlhorn, Parasitology Research Monographs, 5 - Springer Verlag - Berlin Heidelberg: 13.. (Lapsus) 2016. Epomorphorus: Pancer, Gut and Litwinska, Post. Mikrobiol., 55 (2): 208. (Lapsus) 2017. Epomophorous: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 5. Publication date: 6 April 2017. (Lapsus) ? Epomophorus sp.: TAXONOMY: Revised by Bergmans (1988), who transferred Micropteropus grandis to this genus. A key to this genus is presented in Boulay and Robbins (1989) and Claessen and De Vree (1991). See Simmons (2005: 322). Almeida et al. (2016: 83) found Epomophorus to be paraphyletic not only because they transferred dobsonii from the genus Epomops, but also because Micropteropus pusillus seems highly related. However, they suggest further investigation of rapidly evolving nuclear loci and additional species of both genera to decide whether Micropteropus and Epomophorus are synonyms. Included in the Epomophorinii tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Hassanin et al. (2020: 5) include it in the Epomophorina subtribe of the Epomophorini tribe. Almeida et al. (2020: 16) considered Micropteropus Matschie, 1899 to be a synonym of African Chiroptera Report 2020 Epomophorus. They also provisionally considered minimus Claessen and De Vree, 1991 to be a synonym of E. minor Dobson, 1879. Currently recognized species of Epomophorus (Almeida et al., 2020: 16): angolensis Gray, 1870; anselli Bergmans and Van Strien, 2004; crypturus Peters, 1852; dobsonii Bocage, 1889; gambianus (Ogilby, 1835); grandis (Sanborn, 1950); intermedius (Hayman, 1963) labiatus (Temminck, 1837); minor Dobson, 1880; pusillus Peters, 1867; wahlbergi (Sundevall, 1846). COMMON NAMES: Czech: výložkoví kaloni. English: Epauletted Fruit-bats. French: Epomophores, Chauvessouris à épaulettes. German: Epaulettenflughund, Epaulettenflughunde. Italian: Epomòfori. Kiluba (DRC): Mulima. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Fenton (1992: 49) suggests that the white ear spots present disruptive patterns, which contribute to the cryptic appearance of these bats when resting in foliage. DETAILED MORPHOLOGY: The region of the shoulder pouches was morphologically, anatomically and histologically investigated by Püscher (1972), who found special muscles to move the dermal pouch, but no special cutaneous glands. PREDATORS: Steyn (1964: 257) reported an Epauletted Fruit Bat of 6 inches being eaten by a small Python of 2 feet 7 inches. Mikula et al. (2016: Supplemental data) mention Epomophorus sp. to be preyed upon by African goshawk (Accipiter tachiro (Daudin, 1800)) [possibly either E. crypturus or E. wahlbergi], Bat hawk (Macheiramphus alcinus Bonaparte, 1850). PARASITES: BACTERIA Bartonellae Kamani et al. (2014: 628) tested 30 Nigerian Epomophorus sp., and found at least 16 of them testing positive for Bartonella DNA. HAEMOSPORIDA Perkins and Schaer (2016: Suppl.) mention the presence of Hepatocystis epomophori Rodhain, 85 1926 in Epomophorus sp. bats. Of the 138 South Sudanese Epomophorus sp. bats examined by Schaer et al. (2017: 2), 130 (94 %) were infected by Hepatocystis. In the Amurum forest reserve (Nigeria), Atama et al. (2019: 1550) found Hepatocystis parasites in 25 % of the examined Epomophorus bats. HEMIPTERA Cimicidae: Haeselbarth et al. (1966: 8) recorded Afrocimex constrictus Ferris & Usinger 1957 associated with Epomorphorus sp. from Kiambu, near Nairobi, Kenya. DIPTERA Nycteribiidae: Cyclopodia greeffi Karsch 1884 found in a belt across central Africa, between lat. 10o south and north, while in West Africa it occurs further north up to lat. 20o (Haeselbarth et al., 1966: 117). VIRUSES: Coronaviridae Joffrin et al. (2020: 7) reported Beta-D coronaviruses from Epomophorus sp. from Tanzania. Flaviviridae Andral et al. (1968: 855) reported on the isolation of yellow fever virus in Ethiopia from either Epomophorus anurus, "Epomophorus monstruosus" or "Epomophorus limbatus". Paramyxoviridae Orthorubulavirus Krüger et al. (2018: 318) state that broad geographic distribution of the genus Epomophorus in Sub-Saharan Africa makes it conceivable that Bat Mumpsvirus (batMuV) is widely dispersed in the region. This virus is closely related to the human Mumpsvirus of which in 2016 100,576 cases were reported in Africa, and which is serologically not distinguishable from the human virus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Botswana, Burundi, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Gabon, Kenya, Liberia, Mozambique, Namibia, Nigeria, Rwanda, Senegal, South Africa, South Sudan, Tanzania, Uganda, Zambia. Epomophorus angolensis Gray, 1870 *1870. [Epomophorus macrocephalus] var. angolensis Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 125. Type locality: Angola: Benguela [12 34 S 13 24 E, 160 m] [Goto Description]. Holotype: BMNH 86 ISSN 1990-6471 ? 1864.1.9.4: juv ♂. Presented/Donated by: Mr. J.J. Monteiro Esq. - Etymology: Named from a specimen taken at Benguela, Angola (see Smithers, 1983: 56; Taylor, 2005). Epomophorus angolensis: (Name Combination, Current Combination) TAXONOMY: Member of the gambianus species group Simmons (2005: 322). See also Bergmans (1988). COMMON NAMES: Afrikaans: Angola-witkolvrugtevlermuis. Czech: kaloň angolský. English: Angolan Epauletted Fruit-bat, Angolan Epauletted Fruit Bat. French: Epomophore d'Angola, Roussette à épaulettes d'Angola. German: Angola-Epaulettenflughund. CONSERVATION STATUS: Global Justification Listed as Near Threatened. Almost qualifies as threatened under criterion A2c because it is suspected to be experiencing a significant decline at a rate of 20-25% over three generations (12 years; Pacifici et al., 2013) because of the loss of riverine roosting and fruit trees. (Mickleburgh et al., 2008d; IUCN, 2009; Mildenstein, 2016a). This is a precautionary listing, as it is possible that the species has a less restricted habitat, and as such should be listed as Least Concern (Mickleburgh et al., 2008d; IUCN, 2009; Mildenstein, 2016a). Assessment History Global 2016: NT ver 3.1 (2001) (Mildenstein, 2016a). 2008: NT ver 3.1 (2001) (Mickleburgh et al., 2008d; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004b; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). 1994: Rare (Groombridge, 1994). Regional None known. MAJOR THREATS: This species is threatened by the removal of roost trees, often for use as fuel. Griffin (1998) indicates that the removal of riverine trees is a particular threat to species of Epomophorus in Namibia, with relatively little suitable habitat remaining; presumably this is similar problem in Angola. It may have been threatened by the availability of firearms during the long civil war in Angola, although this requires confirmation (Mildenstein, 2016a). CONSERVATION ACTIONS: Current conservation efforts Although there appear to be no direct conservation measures in place for this species it has been recorded from Mupa National Park and Bikuar National Park in Angola (Mickleburgh et al., 2008d; IUCN, 2009; Mildenstein, 2016a). Conservation needs/priorities There is a need to conserve remaining areas of suitable roosting and feeding sites for this species (Mickleburgh et al., 2008d; IUCN, 2009). Studies are needed on the species’ population sizes, distribution, and extent of occurrence throughout its range. Monitoring of population sizes and locations over time are also important to establish whether these are stable or experiencing trends of decline (Mildenstein, 2016a). The threats to these bats are poorly understood. Studies are needed on the species’ habitat requirements and on the effects of forest loss and degradation on the species’ population sizes/distribution. Research is also needed on the amount of hunting and the level of bushmeat trade, and the effects of that hunting on population sizes and persistence (Mildenstein, 2016a). Effective roost site protection efforts are needed to minimize disturbance and protect colonies. Similar to most threatened flying foxes, local capacity building for conservation managers and education and awareness within local communities are greatly needed to support conservation efforts (Mildenstein, 2016a). GENERAL DISTRIBUTION: Epomophorus angolensis is present in Angola and adjacent parts of northern Namibia. It is largely a lowland species, but may range into more montane areas. In Angola it is present west and south of the Mosaic of Guineo-Congolian lowland rain forest and secondary grassland (Bergmans, 1988). In eastern Angola the range seems to be halted by the extensive mosaic region of Brachystegia bakerana thicket and edaphic grassland (Bergmans, 1988). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 339,737 km2. Native: Angola (Bergmans, 1988; CrawfordCabral, 1989; Monadjem et al., 2010d: 551; Taylor et al., 2018b: 60); Namibia (Monadjem et al., 2010d: 551). African Chiroptera Report 2020 HABITAT: Bergmans (1988; 1999) reports that the species is present in wetter Zambezian miombo woodland, north Zambezian woodland; Colophospermum mopane woodland and scrub woodland. Skinner and Chimimba (2005) suggest that the species may be largely confined to riverine forest and other evergreen forest with fruit-bearing trees. Arumoogum et al. (2019: 188) simulated the role of biotic and a-biotic factors in view of climate change and found that the current suitable habitat for E. angolensis is primarily mediated by abiotic variables associated with productivity, but under extreme future climate change scenarios, the most significant factor would be the distribution of freestanding fig trees. 87 Trend:- 2016: Decreasing (Mildenstein, 2016a). 2008: Decreasing (Mickleburgh et al., 2008d; IUCN, 2009). UTILISATION: The species may be hunted for the bushmeat trade (Mildenstein, 2016a). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Namibia. HABITS: Shortridge (1934) reported on animals hanging singly from bare branches of large Acacia trees near the Cunene River, however, loose colonies of up to 200 have also been found. POPULATION: Structure and Density:- The abundance of this species is poorly known, with very little recent information (Mickleburgh et al., 2008d; IUCN, 2009; Mildenstein, 2016a). Figure 15. Distribution of Epomophorus angolensis Epomophorus anselli Bergmans and Van Strien, 2004 *2004. Epomophorus anselli Bergmans and Van Strien, Acta Chiropt., 6 (2): 258. Type locality: Malawi: Kasungu National Park: Lisanthu [13 00 S 33 10 E, 1 000 m]. Holotype: ZMA MAM.21693b: ad ♂, skull and alcoholic. Collected by: Hugo Jachmann; collection date: 19 March 1982; original number: #25. See Bergmans and Van Strien (2004: 258 - 259). Paratype: ZMA MAM.26105: sad ♀, skin and skull. Collected by: Nico[laas] Jan Van Strien; collection date: 19 May 1988; original number: #162. Locality: Lifupa Camp 13.05 S 3005E, 1050 m; see Bergmans and Van Strien (2004: 258 - 259). - Comments: Publication date: The volume is for "September 2004", but the paper has been accepted on 16 October 2004. - Etymology: In honour of Mr. William Frank Harding Ansell for his important contributions to the mammalogy of Malawi and other African countries (see Bergmans and Van Strien, 2004: 262). (Current Combination) 2020. Epomophorus sp1: Hassanin, Bonillo, Tshikung, Pongombo Shongo, Pourrut, Kadjo, Nakouné, Tu, Prié and Goodman, J. Zool. Syst. Evol. Res., 4. Publication date: 20 February 2020. (Name Combination) TAXONOMY: See Bergmans and Van Strien (2004). COMMON NAMES: Czech: kaloň Ansellův. French: Epomophore d'Ansell. German: Ansells Epaulettenflughund. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since it has only recently been described, and there is still very little information on its extent of occurrence, status, threats and ecological requirements (Bergmans and van Strien, 2008; IUCN, 2009; Mildenstein, 2016b). Assessment History Global 2016: DD ver 3.1 (2001) (Mildenstein, 2016b). 2008: DD ver 3.1 (2001) (Bergmans and van Strien, 2008; IUCN, 2009). 88 ISSN 1990-6471 Regional None known. MAJOR THREATS: Bergmans and Van Strien (2004) note that they "have no reason to believe that the species is in any danger but at the same time the available data are few and between 43 and 16 years old". CONSERVATION ACTIONS: Current conservation efforts Bergmans and van Strien (2008) [in IUCN (2009)] reported that this species has been recorded from the Kasungu National Park in Malawi, and further field surveys are needed to better understand the distribution range, threats and conservation status of this species. Conservation needs/priorities Studies are needed on the species’ population sizes, distribution, and extent of occurrence throughout its range. Monitoring of population sizes and locations over time are also important to establish whether these are stable or experiencing trends of decline (Mildenstein, 2016b). The threats to these bats are poorly understood. Studies are needed on the species’ habitat requirements and on the effects of forest loss and degradation on the species’ population sizes/distribution. Research is also needed on the amount of hunting and the level of bushmeat trade, and the effects of that hunting on population sizes and persistence (Mildenstein, 2016b). Native: Malawi (Bergmans and Van Strien, 2004; Monadjem et al., 2010d: 551). HABITAT: Bergmans and Van Strien (2004) record that 'the type specimen was mist netted in Miombo woodland at an elevation of 1,000 m, over a stream between two water berry trees, Syzygium cordatum Hochst., in fruit (Bergmans and Jachmann, 1983).' Furthermore, they state that 'the vegetation of Lifupa Camp, the locality where the female paratype was collected (exact elevation unknown, but between 1,000 and 1,100 m) is Miombo woodland with Brachystegia and Julbernardia (Bergmans and Jachmann, 1983)'. POPULATION: Structure and Density:- It is only known from a few specimens (Bergmans and van Strien, 2008; IUCN, 2009; Mildenstein, 2016b). Trend:- 2016: Unknown (Mildenstein, 2016b). 2008: Unknown (Bergmans and van Strien, 2008; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Malawi. Effective roost site protection efforts are needed to minimize disturbance and protect colonies. Similar to most threatened flying foxes, local capacity building for conservation managers and education and awareness within local communities are greatly needed to support conservation efforts (Mildenstein, 2016b). GENERAL DISTRIBUTION: Epomophorus anselli is currently only known from Malawi, where it has been collected in the Kasungu National Park (collected in 1982 and 1988) and (most probably) the Karonga area (collected in 1961) (Bergmans and Van Strien, 2004). Figure 16. Distribution of Epomophorus anselli Epomophorus crypturus Peters, 1852 *1852. Epomophorus crypturus Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 26, pl. 5, pl. 13, fig. 1 - 6. Type locality: Mozambique: Lower Zambezi River: Tete [16 10 S 33 35 E, 230 m] [Goto Description]. Lectotype: ZMB 10080: ad ♀, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. Lectotype designated by Andersen (1912b: 537); formerly ZMB 553; see Turni and Kock (2008). Paralectotype: ZMB 10018: ♀, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between African Chiroptera Report 2020 1855. 1960. 1986. 1988. 2020. ? ? ? 89 1843 and 1847. See Andersen (1912b) and Turni and Kock (2008: 8). Paralectotype: ZMB 10019: ♀, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Andersen (1912b) and Turni and Kock (2008: 8). Paralectotype: ZMB 10020: ♂, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Andersen (1912b) and Turni and Kock (2008: 8). Paralectotype: ZMB 10078: skull only. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Andersen (1912b) and Turni and Kock (2008: 8). Paralectotype: ZMB 10081: skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. Formerly ZMB 355. See Andersen (1912b) and Turni and Kock (2008: 8). Andersen (1912b: 537) also mentions that most of the paratypes are immature specimens, and that Peters' adult male is in fact an immature. Pt[eropus] crypturus: Giebel, Die Säugethiere., 1002. (Name Combination) Epomophorus gambianus parvus Ansell, Rev. Zool. Bot. afr., 61: 160. Type locality: Zambia: Zambezi district: Chingi [13 01 S 22 44 E]. Holotype: BMNH 1959.610: ad ♂. Comments: Type locality originally Balovale [=Zambezi] district, restricted to Chingi by Ansell (1978: 17). Epomophorus crypturus parvus: Meester, Rautenbach, Dippenaar and Baker, Transv. Mus. Monogr., 5: 27. (Name Combination) Epomophorus gambianus crypturus: Bergmans, Beaufortia, 38 (5): ???. (Name Combination) Epomophorus sp2: Hassanin, Bonillo, Tshikung, Pongombo Shongo, Pourrut, Kadjo, Nakouné, Tu, Prié and Goodman, J. Zool. Syst. Evol. Res., 4. Publication date: 20 February 2020. (Name Combination) Epomophorus angolensis: - Comments: Not of Gray, 1870. Epomophorus crypturus crypturus: (Name Combination) Epomophorus gambianus: - Comments: Not of Ogilby, 1835. TAXONOMY: Bronner et al. (2003) state that Epomophorous crypturus was previously afforded full species status (Corbet and Hill, 1980; Swanepoel et al., 1980; Koopman, 1982; Meester et al., 1986; Claessen and De Vree, 1990, 1991; 2005). Meester et al. (1986) states that Koopman (1966) considers the possibility that the extralimital E. anurus Heuglin, 1864, may be conspecific with crypturus. This would extend the range of crypturus to Uganda, Rwanda, southern Sudan, Ethiopia, Eritrea and westwards to Nigeria. However, Kock (1969a) and Koopman (1975) regard anurus instead as a synonym of the extralimital E. labiatus (Temminck, 1837), while Hayman and Hill (1971) retain it as a separate species, so that a close relationship with crypturus does not seem to be indicated. Further uncertainty surrounds the status of E. gambianus parvus. Ansell (1978), while treating parvus as a subspecies of gambianus, expresses doubt whether parvus can be distinguished at species level from crypturus, and suggests that crypturus and gambianus are conspecific, with parvus as an intermediate subspecies or a stage in a cline running between them. Koopman (1982) retains gambianus as a separate species, but includes parvus in crypturus, without specifying whether or not he regards it as subspecifically distinct. While acknowledging that this problem remains to be solved we shall provisionally treat parvus as a taxon of crypturus. COMMON NAMES: Afrikaans: Peters se witkolvrugtevlermuis, Peterswitkolvrugtevlermuis, Klein Vrugtevlermuis. Czech: kaloň mosambický. English: Peters's Epauletted Fruit Bat, Smaller Epauletted fruit bat. French: Epomophore de Peters, Roussette à épaulettes de Peters. German: Peters' Epaulettenflughund. Nyanja (Malawi): Mleme. Portuguese: Morcego frugivoro de Peters. ETYMOLOGY OF COMMON NAME: In honour of Wilhelm Carl Hartwig Peters who, in 1852, published a monumental work on zoological material collected mainly in the Tete District of Mozambique (Smithers, 1983: 58). SIMILAR SPECIES: Taylor and Monadjem (2008: 24) provided data which enabled them to separate E. crypturus from E. wahlbergi. In the first species, the maxillary width is always smaller than in the latter (♂♂: < 14 mm; ♀♀: < 13 mm). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in a number of protected 90 ISSN 1990-6471 areas, has a tolerance of a degree of habitat modification, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008e; IUCN, 2009; Taylor, 2016c). Assessment History Global 2016: LC ver 3.1 (2001) (Taylor, 2016c). 2008: LC ver 3.1 (2001) assessed as E. crypturus (Mickleburgh et al., 2008e; IUCN, 2009). 2004: LC ver 3.1 (2001) assessed as E. crypturus (Mickleburgh et al., 2004c; IUCN, 2004). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016x). 2004: DD ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: There appear to be no major threats to this species as a whole. Some populations may be impacted by general persecution as crop pests (Mickleburgh et al., 2008e; IUCN, 2009; Taylor, 2016c). CONSERVATION ACTIONS: Taylor (2016c) agrees with Mickleburgh et al. (2008e) [in IUCN (2009)] who state that this species has been recorded from many protected areas, and no direct conservation measures are needed for this widespread and adaptable species as a whole. GENERAL DISTRIBUTION: E. crypturus is a widespread southern African species distributed from southern parts of the Democratic Republic of the Congo and southern Tanzania, to the eastern coastline of South Africa. It ranges from eastern Angola and northern Botswana to the southeastern African coastline. It has been recorded at elevations of up to 2,185 m asl, although it has mostly been collected between 500 and 1,500 m asl (Mickleburgh et al., 2008e; IUCN, 2009). For South Africa, Babiker Salata (2012: 49) found that its distribution is stongly associated with the mean temperature of the coldest quarter. Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,213,597 km2. Native: Angola (Bergmans, 1988; Monadjem et al., 2010d: 551); Botswana (Bergmans, 1988; Monadjem et al., 2010d: 551); Congo (The Democratic Republic of the) (Schouteden, 1944; Hayman et al., 1966; Monadjem et al., 2010d: 551); Malawi (Happold et al., 1988; Bergmans, 1988; Happold and Happold, 1997b: 813; Monadjem et al., 2010d: 551); Mozambique (Smithers and Lobão Tello, 1976; Bergmans, 1988; Monadjem et al., 2010d: 551; 2010c: 376); Namibia (Monadjem et al., 2010d: 552); South Africa (Monadjem et al., 2010d: 552); Swaziland (Monadjem et al., 2010d: 552); Tanzania; Zambia (Ansell, 1978; Monadjem et al., 2010d: 552); Zimbabwe (Monadjem et al., 2010d). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - In Zimbabwe, Peterson and Nagorsen (1975: 4) recorded the following: 2n = 35 for a male, and 2n = 36 for a female (XO/XX; see also Nesi et al., 2011: 552). HABITAT: This species is generally associated with dry savanna and riverine forest with fruit bearing trees (Mickleburgh et al., 2008e). ROOST: Colonies often roost under the canopy of trees, in thick foliage (Mickleburgh et al., 2008e). MIGRATION: Populations exhibit considerable movements in search of food, and may come into towns and feed on crops and fruit trees (Mickleburgh et al., 2008e). DIET: Bonaccorso et al. (2014: 41) studied the foraging movements of E. crypturus in the Kruger National Park (RSA) in relation to the distribution of sycamore figs Ficus sycomorus. They also performed germination trials on 20 intact seeds that had passed through the gastro-entero tract of the bats and found 100 % germination. They also found (p. 48) that during a very dry period, the bats flew between 4 and 14 km from their roosting site to fruiting F. sycomorus. In normal circumstances, they only flew 400 - 2,000 m. The bats visited up to four fig trees per night. POPULATION: Structure and Density:- It is a quite common species, sometimes forming loose colonies of hundreds of bats (Mickleburgh et al., 2008e; IUCN, 2009; Taylor, 2016c). It is a quite common species, sometimes forming loose colonies of hundreds of bats. Trend:- 2016: Unknown (Taylor, 2016c). 2008: Unknown (Mickleburgh et al., 2008e; IUCN, 2009). REPRODUCTION AND ONTOGENY: In the Kruger Park, Pienaar (1964: 8) reported single young attached to their mothers in December and January. African Chiroptera Report 2020 In Malawi, Happold and Happold (1990b: 564) found lactating females in November, March and May, suggesting aseasonal reproduction. MATING: McCracken and Wilkinson (2000: 349) suggest that E. crypturus uses nocturnal display sites where females visit males for mating. Males call from roosts in trees that are spaced about 50 m apart along rivers. 91 DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Botswana, Central African Republic, Congo (Democratic Republic of the), Eswatini, Malawi, Mozambique, Namibia, South Africa, Tanzania, Zambia, Zimbabwe. At Shingwedzi (Kruger National Park), Adams and Snode (2015: 4) found the following values for the male mating calls: mean fundamental frequency: 15.71 ± 0.07 kHz (first harmonic), only in one case followed by multiple harmonics at 16, 48, 64, 96, and 112 kHz. VIRUSES: Coronaviridae - Coronaviruses SARS-CoV: Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in four individuals from Limpopo Province, none tested positive (0/4) and six individuals from Mpumalanga Province, South Africa, none tested positive (0/6). Figure 17. Distribution of Epomophorus crypturus Epomophorus dobsonii Bocage, 1889 *1889. Epomophorus Dobsonii Bocage, J. Sci. mat. phys. nat., ser. 2, 1 (1): 1, Fig 1. Publication date: March 1889. Type locality: Angola: Benguela district: Quindumbo [Goto Description]. Neotype: AMNH 88068: ♂, skin and skull. Collected by: the PhippsBradley Expedition; collection date: 27 February 1933; original number: 902. At Chitau, 4930 ft. Designated by Bergmans (1989) (see AMNH website). - Comments: Andersen (1912b: 501) mentions that the original type was an adult male in the Lisbon museum. Frequently mentioned as dobsoni. - Etymology: Named after George Edward Dobson who, among his many other papers on bats, published a Catalogue of the Chiroptera in the collection of the British Museum in 1878 (see Taylor, 2005). 1899. Epomophorus (Epomops) dobsoni: Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 57. (Name Combination) 1912. Epomops dobsoni: K. Andersen, Catalogue of the Chiroptera in the British Museum. Second Edition. Volume I: Megachiroptera: viii, lvi, 500. (Name Combination, Lapsus) 1939. Epomops dobsonii: G.M. Allen, Bull. Mus. comp. Zool., 83: 57. Publication date: February 1939. (Name Combination) 2016. E. dobosonii: Calderon, Guzman, Salazar-Bravo, Figueiredo, Mattar and Arrieta, Manter, Suppl. 6: 7. Publication date: 30 September 2016. (Lapsus) ? Epomophorus dobsoni: (Alternate Spelling) ? Epomophorus dobsonii: TAXONOMY: See Bergmans (1979b; 1989) and Simmons (2005: 323). The phylogenetic analyses by Almeida et al. (2016: 83) suggest that dobsonii is closely related to Epomophorus wahlbergi and they transfered the species to the latter genus. COMMON NAMES: Afrikaans: Dobson se vrugtevlermuis, Dobsonvrugtevlermuis. Czech: kaloň Dobsonův. English: Dobson's Fruit Bat, Dobson's Epauletted Fruit Bat, Dobson's Epauletted Bat, Singing Fruit Bat. French: Epomophore de Dobson, Roussette de Dobson. German: Dobsons Epaulettenflughund. 92 ISSN 1990-6471 CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bx; IUCN, 2009). Assessment History Global 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bx; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cj; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There appear to be no major threats to this species (Mickleburgh et al., 2008bx; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008bx) [in IUCN (2009)] report that it has been recorded from Kasungu National Park in Malawi and from Parc National de l'Upemba in the Democratic Republic of the Congo. There is a need for additional studies into the natural history of this species and into any possible future threats to the species (Mickleburgh et al., 1992). GENERAL DISTRIBUTION: Epomophorus dobsonii is distributed in East Africa, Central Africa and southern Africa. It has been recorded from Angola, Namibia, Zambia, southern Democratic Republic of the Congo, Malawi and Mozambique, being found as far north as Tanzania and possibly Rwanda. It has been recorded at elevations of up to 1,500 m asl (in Malawi) and at 1,890 m asl at Kibwele, Tanzania. Tanzania; Zambia (Ansell, 1973; Bergmans, 1989; Monadjem et al., 2010d: 554); Zimbabwe. Presence uncertain: Namibia (see Cotterill, 2004a: 260); Rwanda. POPULATION: Structure and Density:- It does not appear to be an especially rare species in Malawi. There is little information on the abundance of the species over the rest of its range, but it is presumed to be reasonably common (Mickleburgh et al., 2008bx; IUCN, 2009). Trend:- 2008: Stable (Mickleburgh et al., 2008bx; IUCN, 2009). REPRODUCTION AND ONTOGENY: Hill (1941) [in Fox et al. (2009: 272) reported that "E. dobsoni" gave birth to two young, but Fox et al. (2009: 272) indicate that this statement was based on translated field notes, and it is not clear whether they refer to twins or to two birthing seasons each year. VIRUSES: Rhabdoviridae Lyssavirus - Rabies related viruses de Jong et al. (2011: 9) and Willoughby et al. (2017: Suppl.) report Lagos Bat Virus from E. dobsonii. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Botswana, Congo (Democratic Republic of the), Malawi, Rwanda, Tanzania, Zambia, Zimbabwe. For southern Africa (south of 8° S), CooperBohannon et al. (2016: Table S2) calculated a potential distribution area of 503,585 km 2. Native: Angola (Crawford-Cabral, 1989; Bergmans, 1989; Monadjem et al., 2010d: 553; Taylor et al., 2018b: 60); Botswana (Monadjem et al., 2010d: 554); Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 554); Malawi (Bergmans, 1989; Monadjem et al., 2010d: 554); Mozambique; Namibia; Figure 18. Distribution of Epomophorus dobsonii Epomophorus gambianus (Ogilby, 1835) *1835. Pteropus Gambianus Ogilby, Proc. zool. Soc. Lond., 1835, III (xxxi): 100. Publication date: 9 October 1835. Type locality: The Gambia: Banjul [Goto Description]. Holotype: African Chiroptera Report 2020 1898. 2017. 2019. ? 93 BMNH 1907.1.1.233: ad ♀, skin and skull. Collected by: Dr. Percy Rendall. Formerly in Tomes collection. Paratype: BMNH 1874.a-c: 2 sad FF, 1 imm F, skin and skull. Collected by Liet. Rendall. Paratype: BMNH 1874.a = 124b: imm ♀. Presented/Donated by: ?: Collector Unknown. Paratype: BMNH 1874.b = 124c: ad ♀. Collected by: Dr. Percy Rendall. Presented/Donated by: ?: Collector Unknown. Paratype: BMNH 1874.c = 124d: imm ♀. Presented/Donated by: ?: Collector Unknown. Paratype: BMNH 1907.1.1.231: sad ♀, skin and skull. Collected by: Dr. Percy Rendall. Formerly in Tomes collection. - Comments: Type locality restricted by Kock et al. (2002: 80). - Etymology: Named as the species was described from a specimen from Gambia in West Africa (see Taylor, 2005). Epomophorus guineensis Bocage, J. Sci. mat. phys. nat., ser. 2, 5 (19): 136, fig. 3. Publication date: June 1898. Type locality: Guinea-Bissau: Bolama [11 35 N 15 30 W] [Goto Description]. - Comments: Andersen (1912b: 541) mentions the type specimen as being a mounted ad ?, skull extracted, coll. Sr. Damasceno da Costa, and in the Lisbon Museum. Epomorphus giambianus: Banyard and Fooks, Microbiol. Aust., 38 (1): 18. Publication date: March 2017. (Lapsus) E[pomophorus] gambiansus: Mbu'u, Mbacham, Gontao, Kamdem, Nlôga, Groschup, Wade, Fischer and Balkema-Buschmann, Vector-Borne Zoon. Dis., xxx: "6". Publication date: 13 April 2019. (Lapsus) Epomophorus gambianus: (Name Combination, Current Combination) TAXONOMY: See Boulay and Robbins (1989), Claessen and De Vree (1990) and Simmons (2005: 322). Two subspecies are currently recognized: E. g. gambianus (Ogilby, 1835) and E. g. pousarguesi Trouessart, 1904. COMMON NAMES: Afrikaans: Gambiaanse witkolvrugtevlermuis. Czech: kaloň výložkový, kaloň větší. English: Gambian Epauletted Fruit bat, Gambian Epauleted Bat. French: Epomophore de Gambie, Chauve-souris à épaulettes de Gambie. German: Gambia Epaulettenflughund. Tanoboase (Ghana): sreso ampane [="original or normal bat'], ampane kronfo [="thief or criminal bat"] (see Ohemeng et al., 2017: 184). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001) assessed as full species in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008f; IUCN, 2009; Tanshi and Fahr, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Tanshi and Fahr, 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008f; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004I; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There are no major threats to this species as a whole. In parts of its range it is locally threatened by overhunting for food and habitat loss through logging and conversion of land to agricultural use (Mickleburgh et al., 2008f; IUCN, 2009) and climate change (Tanshi and Fahr, 2016). CONSERVATION ACTIONS: Mickleburgh et al. (2008f) [in IUCN (2009)] state that the species has a wide range, and is presumably present in many protected areas. GENERAL DISTRIBUTION: Senegal, Gambia over Guinea (Fahr and Ebigbo, 2003: 128), Mali (Meinig, 2000: 104), Benin (CapoChichi et al., 2004: 161), Cameroon (Bakwo Fils et al., 2014: 3), Central African Republic (Morvan et al., 1999: 1195) to W Ethiopia, south to South Sudan. Simmons (2005: 322) includes Malawi and Botswana, these records may be associated with crypturus and need to be verified. E. g. gambianus - west Africa. E. g. pousarguesi - east Africa. Native: Benin (Capo-Chichi et al., 2004: 161), Burkina Faso (Kangoyé et al., 2015a: 605), Cameroon (Bakwo Fils et al., 2014: 3), Central African Republic (Morvan et al., 1999: 1195), Chad, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Gabon, Ghana, Guinea (Fahr 94 ISSN 1990-6471 and Ebigbo, 2003: 128; Decher et al., 2016: 261), Guinea-Bissau (Bocage, 1889a:2; Seabra, 1898b; Monard, 1939; Veiga-Ferreira, 1948; Bergmans, 1988; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 22), Liberia, Mali (Meinig, 2000: 104), Niger, Nigeria, Senegal, Sierra Leone, South Sudan, Sudan, The Gambia, Togo. SEXUAL DIMORPHISM: Dobson (1873: 247) quotes Tomes' description of the male as: "The conspicuous shoulder-tufts of E. macrocephalus are here very fully developed. They consist of a very slight warty excrescence clothed with fur, which differs from that which surrounds it only in being of a dirty white colour." MOLECULAR BIOLOGY: Riesle-Sbarbaro et al. (2016: 447) reported on the complete mitochondrial genome of this species, which consists of 16,702 bases. Riesle-Sbarbaro et al. (2018: "13") examined mitochondrial and nuclear DNA and found that connectivity and free mixing occurred between the colonies sampled over most of its distribution area. Their data lso indicated that male dispersal occurs more often and/or over longer distances than females. The Ethiopian colonies seem to form an exception as the genetic differences indicate that they apparently diverged about 1.6 Mya from the bats from the more western part of the distribution area. Also the populations from southern Ghana (Accra and Ve-Golokwati) show some genetic differentiation. Riesle-Sbarbaro et al. (2018: "14") suggest that the geographical barriers (e.g. mountains and water bodies such as rivers and lakes) and other environmental developments (e.g., urbanization of megacities) are the likely drivers for the regional divergences. HABITAT: This species is present in a variety of tropical moist and dry forest, savanna, bushland and mosaic habitats, and has additionally been reported from mangroves and swamp forests (Mickleburgh et al., 2008f). It is able to persist in areas of partially degraded forest. HABITS: Anonymus (2002a: 103) mentions that in some areas of Benin, E. gambianus might be causing large trees to die, as they swarm in large numbers on a restricted surface. ROOST: Roosting usually takes place as individuals or small groups in dense vegetation (Happold, 1987). E. gambianus forms roosting colonies in crowns of trees, containing a few to 50 - 100 individuals (Kunz, 1996: 44). Ayensu (1974: 716) indicates that in Ghana, Azadirachta indica or the Neem tree is a favorite roosting haven. Lawson et al. (2018: 110) found them also roosting in mango (Mangifera sp.) and fig (Ficus sp.) trees around schools, churches, market places and people's homes. MIGRATION: Isotopic data collected by Omatsu et al. (2008: 21 - 22) suggest that E. gambianus has an East-West migratory behaviour inside its distribution range, which is even more pronounced than the migration displayed by Eidolon helvum. Six specimens that were captured in February in Mali were most probably originating from some 3,000 km to the east. DIET: Ayensu (1974) reports that in West Africa, E. gambianus visits Parkia clappertoniana (Dawadawa), Ceiba pentandra (Silk-Cotton tree or Kapop tree), Mangifera indica (Mango tree), Anacardium occidentale (Cashew tree), Psidium guajava (Guava tree), Carica papaya (Papaya or pawpaw), and Azadirachta indica (Neem tree) when foraging. Bachirou et al. (2011: 2) indicate that E. gambianus is the most important seed disperser for the Neem tree in Cameroon. Kangoyé et al. (2010: 291) show a picture of E. gambianus pollinating Ceiba pentandra flowers, and also indicate that this species was found transporting Shea fruits (Vitellaria paradoxa) or figs (Ficus sp.), whereas others were found in cashew fields (Anacardium occidentale). Seltzer et al. (2013: table S2) provide an overview of the plant species found beneath bat feeding roosts. For E. gambianus these include Rauvolfia caffra Sond. (Quinine Tree - Apocynaceae) and Psidium guajava L. (Apple guava - Myrtaceae). Based on faecal pellets and ejecta, AmponsahMensah et al. (2019: 5687) found the following fruits (FR) and flowers (FL) in the diet of bats from Ghana: Anacardiaceae: Anacardium occidentale L. (Cashew ) FR), Mangifera indica L. (Mango FR), Spondias mombin L. (Yellow mombin - FR); Annonaceae: Annona muricata L. (Soursop - FR), Polyalthia longifolia Sonn. (Indian mast tree, false ashoka - FR); Bignoniaceae: Spathodea campanulata P. Beauv. (African tulip tree - FL); Bombacaceae: Bombax buonopozense P. Beauv. (Gold Coast bombax, Red-flowered silk-cotton FL), Ceiba pentandra (L.) Gaertn. (Kapok, Silk Cotton - FL); Caricaceae: Carica papaya L. (Papaya, Pawpaw - FR); Combretaceae: Terminalia catappa L. (Indian almond - FR); Cucurbitaceae: Melothria sp. (Mouse melon - FR); Fabaceae: Daniellia oliveri (Rolfe) Hutch. & Dalziel African Chiroptera Report 2020 (African copaiba balsam - FL), Parkia biglobosa (Jacq.) R.Br. ex G.Don (African locust bean - FL); Gentianaceae: Anthocleista vogelii Planch. (Cabbage tree - FR, FL); Malvaceae: Adansonia digitata L. (Baobab - FL), Sterculia rhinopetala K. Schum. (Brown sterculia, Red sterculia - FR); Meliaceae: Azadirachta indica A.Juss (Neem - FR, FL); Moraceae: Antiaris toxicaria Lesch. (False iroko, Antiaris - FR), Ficus spp. (five species Figs - FR), Milicia excelsa (Welw.) C.C. Berg (Iroko, Odum - FR); Musaceae: Musa sp. (Banana - FR, FL); Myrtaceae: Psidium guajava L. (Guava - FR), Syzygium sp. (woodland waterberry - FR); Solanaceae: Solanum sp. (Potato tree - FR); Verbenaceae: Vitex doniana Sweet (Black plum FR). Ficus sp. was present in over a third of all the samples and were eaten in all months. Other fruits that played a vital role, and were prefered over other available fruits, include Vitex doniana and Anthocleista vogelii. The main reason was probably the higher protein content of these fruits. They were also highly available in the period of late pregnancy (March and September) and parturition (April/May and October/November) (p. 5689). Amponsah-Mensah et al. (2019: 5690) could not confirm whether E. gambianus were actively eating pollen, nectar, or both, even when they observed bats visiting flowering trees and being covered in pollen. PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator, as well as the Pied crow (Corvus albus Statius Muller, 1776). Ayivor et al. (2017: 6) indicate that the latter targets young, just weaned E. gambianus. POPULATION: Structure and Density:- This is a common species in the western parts of its range. It is gregarious and form colonies of several hundreds individuals or less (Mickleburgh et al., 2008f; Tanshi and Fahr, 2016) . Trend:- 2016: Unknown (Tanshi and Fahr, 2016). 2008: Unknown (Mickleburgh et al., 2008f; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 7.8 years. REPRODUCTION AND ONTOGENY: Szekely et al. (2015: Suppl.) report that the gestation period lasts for 150 days, and that the young's weight at birth is 10 g (the adult weights 80 g). 95 MATING: Nesi et al. (2011: 552) indicate that the male's courting call has a frequency of 1,750 Hz. PARASITES: BACTERIA Höhne et al. (1975: 505) found Listeria monocytogenes bacteria in one out of two E. gambianus specimens from Lomé, Togo. HAEMOSPORIDA Schaer et al. (2013a: 17416) report the presence of hemosporidian parasites of the genus Hepatocystis in one investigated bat. Miltgen et al. (1977: 595) pointed out that the first reports of gametocytes from this parasite were made by the Léger(s) from Niger in 1914. VIRUSES: Willoughby et al. (2017: Suppl.) report the following viruses: Lagos bat lyssavirus, Nipah henipavirus, Reston ebolavirus, Zaire ebolavirus. Coronaviridae Joffrin et al. (2020: 7) reported Beta-D coronaviruses from bats from Cameroon. Filoviridae - Filoviruses Ebolavirus Hayman et al. (2012d: 1208) examined 37 bats from the Greater Accra region, Ghana, and found 14 to be positive for EBOV antibodies. Paramyxoviridae Paramyxoviruses Hayman et al. (2008a: 926; 2008b: 2) detected antibodies to Nipah virus in one bat collected in Ghana in 2007. Drexler et al. (2012a: Suppl. Table S1) found three bats out of 54 (5.4 %) testing positive for Henipavirus in the Democratic Republic of the Congo and Ghana. Rhabdoviridae Lyssavirus - Rabies related viruses 2006 in Nigeria, Dzikwi et al. (2010: 269) tested 12 brains by direct fluorescent antibody (DFA) and mouse inoculation test (MIT) which tested negative for lyssavirus antigens. Lagos bat virus (LBV): Hayman et al. (2008a) detected antibodies in Ghana. In 2006, 12 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), where 3 tested positive (25%) for neutralizing anitibodies (Dzikwi et al., 2010: 269). Mokola virus (MOKV): In 2006, 12 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), none tested positive for neutralizing anitibodies (Dzikwi et al., 2010: 269). 96 ISSN 1990-6471 In their overview table, Maganga et al. (2014a: 8) reported that the following viruses were already found in E. gambianus: Lagos bat virus (LBV), Nipah virus (NPHV), Zaire Ebola virus (ZEBOV), Reston Ebola virus. Niger, Nigeria, Senegal, Sierra Leone, South Sudan, Sudan, Tanzania, The Gambia, Togo, Zambia, Zimbabwe. UTILISATION: In some localities in Benin, specimens of this species are eated, but always in small numbers (Anonymus, 2002a: 102). Dougnon et al. (2012: 85) reported that the carcasses have an average weight of 40.52 ± 5.08 g (about 43 % of the body mass). They also evaluated the quality of the meat (for 100 g): total mineral matter: 14.1 g, lipids: 13.36 g, protein: 58.4 g, Calcium: 44 mg, Sodium: 48 mg, Phosphore: 80 mg and Potassium: 88 mg. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Cameroon, Central African Republic, Chad, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Liberia, Mali, Figure 19. Distribution of Epomophorus gambianus Epomophorus gambianus gambianus (Ogilby, 1835) *1835. Pteropus Gambianus Ogilby, Proc. zool. Soc. Lond., 1835, III (xxxi): 100. Publication date: 9 October 1835. Type locality: The Gambia: Banjul [Goto Description]. Holotype: BMNH 1907.1.1.233: ad ♀, skin and skull. Collected by: Dr. Percy Rendall. Formerly in Tomes collection. Paratype: BMNH 1874.a-c: 2 sad FF, 1 imm F, skin and skull. Collected by Liet. Rendall. Paratype: BMNH 1874.a = 124b: imm ♀. Presented/Donated by: ?: Collector Unknown. Paratype: BMNH 1874.b = 124c: ad ♀. Collected by: Dr. Percy Rendall. Presented/Donated by: ?: Collector Unknown. Paratype: BMNH 1874.c = 124d: imm ♀. Presented/Donated by: ?: Collector Unknown. Paratype: BMNH 1907.1.1.231: sad ♀, skin and skull. Collected by: Dr. Percy Rendall. Formerly in Tomes collection. - Comments: Type locality restricted by Kock et al. (2002: 80). - Etymology: Named as the species was described from a specimen from Gambia in West Africa (see Taylor, 2005). 1835. Pteropus macrocephalus Ogilby, Proc. zool. Soc. Lond., 1835, III (xxxi): 101. Publication date: 9 October 1835. Type locality: The Gambia: "Gambia" [Goto Description]. Comments: collected by Rendall. 1835. Pteropus megacephalus Swainson, Natural History and Classification of Quadrupeds, 91, 92, 356, figs 31, 154. Type locality: The Gambia. - Comments: Type locality originally as West Africa, but Andersen (1912b: 539) mentions "Gambia" as type locality. 1836. Pteropus epomophorus Bennett, Proc. zool. Soc. Lond., 1835, III (xxxiv): 149. Publication date: 12 February 1836. Type locality: The Gambia: "Gambia". Holotype: BMNH 1907.1.1.232: ad ♂, skin only. Collected by: Dr. Percy Rendall. (Type for both epomophorus and whitei). Ex Tomes collection: see Andersen (1912b: 542). Holotype: NRM [unknown]: ad ♀, alcoholic (skull not removed). Collected by: Bror Yngve Sjöstedt; collection date: 4 May 1905. Kilimandjaro-Meru Expedition. See Lönnberg (1908b: 7) and Andersen (1912b: 810). 1836. Pteropus Whitei Bennett, Trans. Linn. Soc. Lond., 2 (1): 37, pl. 6, pl. 7. Publication date: 2 October 1836. Type locality: The Gambia: "Gambia" [Goto Description]. Holotype: BMNH 1907.1.1.232: ad ♂, skin only. Collected by: Dr. Percy Rendall. (Type for both epomophorus and whitei). Ex Tomes collection: see Andersen (1912b: 542). 1838. Epomophorus whitii: Gray, Mag. Zool. Bot., Edinburgh, 2 (12): 504. Publication date: 1 February 1838. - Comments: Also incorrect subsequent spelling, see Boulay and Robbins (1989: 1). (Name Combination) 1898. Epomophorus guineensis Bocage, J. Sci. mat. phys. nat., ser. 2, 5 (19): 136, fig. 3. Publication date: June 1898. Type locality: Guinea-Bissau: Bolama [11 35 N 15 30 W] African Chiroptera Report 2020 1899. 1950. ? ? ? 97 [Goto Description]. - Comments: Andersen (1912b: 541) mentions the type specimen as being a mounted ad ?, skull extracted, coll. Sr. Damasceno da Costa, and in the Lisbon Museum. Ep[omophorus (Epomophorus)] zechi Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 46, pl 10, fig. 3 a-c. Type locality: Benin: Togoland: Kunjuruma [Goto Description]. Lectotype: ZMB 10170/17171: ad ♂, skin and skull. Collected by: Baumann; collection date: 1894. Lectotype designated by Andersen (1912b: 541); see Turni and Kock (2008) [skin: ZMB 10170 - skull: ZMB 17171]. Paralectotype: ZMB 10009: ad ♂, skin and skull. Collected by: Baumann; collection date: 1894. Paralectotype: ZMB 10011: juv ♂, skull only. Collected by: Bloess. Gross-Popo. Paralectotype: ZMB 10065: ad ♀, skull and alcoholic. Collected by: Anton Reichenow. Accra, Ghana. Paralectotype: ZMB 3413: ad ♂, skull and alcoholic. Lagos, Nigeria, acquired from Salmin. Paralectotype: ZMB 356: ad ♂, skin and skull. Collection date: 1855. West Africa acquired from Salmin. Paralectotype: ZMB 3654: ad ♂, skull and alcoholic. Collected by: Unger. Accra, Ghana. Paralectotype: ZMB 4784: ad ♂, skull and alcoholic. Collected by: Anton Reichenow. Accra, Ghana. - Comments: Kunjuruma is the location from lectotype: see Allen (1939a: 56). Boulay and Robbins (1989: 1) mention "Accra, Goldkuste [=Ghana], Gross-Popo, Misahoho in Togoland, Lagos [Nigeria], restricted to Kunjuruma, Togoland [=Benin] by Andersen (1912b)". Turni and Kock (2008: 10) mention "Matschie (1899b: 46) gave the locality of the specimens collected by Baumann as 'Misahöhe, Togoland' (= Missahoe, Togo); however, according to the entry book the locality is Kunjuruma." Preface by Matschie is dated 14 May 1899. Epomophorus reii Aellen, Rev. suisse Zool., 57 (3): 559. Type locality: Cameroon: Rei Bouba [ca. 08 37 N 14 09 E] [Goto Description]. Holotype: MHNC 32-8-970: ad ♀. Collected by: MSS Cameroun; collection date: 13 September 1947. Presented/Donated by: ?: Collector Unknown. Paratype: MHNC 32-8-969: imm ♀. Collected by: MSS Cameroun; collection date: 13 September 1947. Presented/Donated by: ?: Collector Unknown. - Comments: in part?. Epomophorus anurus: - Comments: Not of Heuglin, 1864. Epomophorus gambianus gambianus: (Name Combination, Current Combination) Epomophorus sp.: TAXONOMY: Simmons (2005:322) does not include crypturus. GENERAL DISTRIBUTION: This subspecies ranges from Senegal in the west, through much of West Africa and parts of Central Africa. It is typically a lowland species occurring below 500 m asl. Native: Benin; Burkina Faso; Cameroon; Central African Republic; Chad; Congo (The Democratic Republic of the); Côte d'Ivoire; Gambia; Ghana; Guinea; Guinea-Bissau (Bocage, 1889a:2; Seabra, 1898b; Monard, 1939; Veiga-Ferreira, 1948; Bergmans, 1988; Lopes and CrawfordCabral, 1992; Rainho and Ranco, 2001: 22); Liberia; Mali; Nigeria; Senegal; Sierra Leone; Togo. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Cameroon, Chad, Côte d'Ivoire, Ethiopia, Ghana, Guinea, Liberia, Mali, Niger, Nigeria, Senegal, Sierra Leone, Tanzania, The Gambia, Zimbabwe. Epomophorus gambianus pousarguesi Trouessart, 1904 *1904. Epomophorus pousarguesi Trouessart, Catalogus Mammalium tam viventium quam fossilium, supplement, 55. Type locality: Central African Republic: Grande Brousse between Yabanda and Mpoko. Holotype: MNHN ZM-MO-1892-1020: Collected by: Jean Dybowski; collection date: 1-11 December 1891. Bergmans (1978a: 682, 683) mentions two different numbers for this specimen. In the table on pag 682, he mentions 1899-1020 and on page 683 he mentions 1892-1020. - Comments: Boulay and Robbins (1989: 1) mention "Africa Occ.-Centrali, a fl. Chari (Inter Yabanda et Mpoko), reg. Lac. Chad. Central African Republic as type locality; restricted to 'along the track between Mpoko (near Makorou) and Yabanda' by Bergmans (1978b)". The type locality is sometimes given as "Eastern Congo, Shari region". Description based on E. macrocephalus Pousargues (not Ogilby) see Allen (1939a: 56). 98 ISSN 1990-6471 ? ? Epomophorus gambianus pousarguesi: (Name Combination, Current Combination) Epomophorus gambianus: - Comments: Not of Ogilby, 1835. TAXONOMY: Recognized as a subspecies (Simmons, 2005: 322). GENERAL DISTRIBUTION: In Ethiopia populations may be found up to 2,000 m asl. Native: Ethiopia, South Sudan, Sudan. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Central African Republic. Epomophorus grandis (Sanborn, 1950) *1950. Micropteropus grandis Sanborn, Publ. Cult. Comp. Diamantes Angola, Lisboa, 10: 55, figs 2 - 3. Publication date: 29 November 1950. Type locality: Angola: Lunda: Dundo [07 22 S 20 50 E]. Holotype: FMNH 66433: ad ♀, skull and alcoholic. Collected by: A. de Barros Machados; collection date: September 1948; original number: 2015. 1988. Epomophorus grandis: Bergmans, Beaufortia, 38 (5): ???. (Name Combination, Current Combination) TAXONOMY: Transferred from Micropteropus to Epomophorus by Bergmans (1988). See Simmons (2005: 322). COMMON NAMES: Czech: kaloň Sanbornův. English: Sanborn's Epauletted Fruit-bat, Lesser Angolan Epauletted Fruit Bat. French: Epomophore de Sanborn, Petit Epomophore frugivore d'Angola. German: Sanborns Epaulettenflughund. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of recent information on its extent of occurrence, ecological requirements, threats and conservation status (Mickleburgh et al., 2008h; IUCN, 2009, Fahr and Mildenstein, 2016). Assessment History Global 2016: DD ver 3.1 (2001) (Fahr and Mildenstein, 2016). 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008h; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004e; IUCN, 2004). 2000: DD (Hilton-Taylor, 2000). 1996: EN (Baillie and Groombridge, 1996). 1994: Rare (Groombridge, 1994). Regional None known. MAJOR THREATS: The threats to this species are not known. There is deforestation taking place in parts of the species range. However, it is not known how adaptable this species is to habitat modification (Mickleburgh et al., 2008h; IUCN, 2009; Fahr and Mildenstein, 2016). CONSERVATION ACTIONS: Current conservation efforts It is not known if the species is present in any protected areas. The area where it occurs has been inaccessible for many years ((Mickleburgh et al., 2008h) in IUCN (2009, Fahr and Mildenstein, 2016) . Conservation needs/priorities Studies are needed on the species’ population sizes, distribution, and extent of occurrence throughout its range ((Mickleburgh et al., 2008h) in IUCN (2009, Fahr and Mildenstein, 2016). Monitoring of population sizes and locations over time are also important to establish whether these are stable or experiencing trends of decline. The threats to these bats are poorly understood. Studies are needed on the species’ natural history and habitat requirements and on the effects of forest loss and degradation on the species’ population sizes/distribution. Research is also needed whether the species is hunted, and if so, on the amount of hunting and the level of bushmeat trade, and the effects of that hunting on population sizes and persistence (Fahr and Mildenstein, 2016). Effective roost site protection efforts are needed to minimize hunting mortality and disturbance to nontarget individuals. Similar to most threatened flying foxes, local capacity building for conservation managers and education and awareness within local communities are greatly needed to support conservation efforts. African Chiroptera Report 2020 GENERAL DISTRIBUTION: Epomophorus grandis is a poorly known as it has only been recorded at the type locality of Dundo in northeast Angola (Sanborn, 1950), and from Pointe Noire on the coast of Congo (Bergmans, 1988). 99 UTILISATION: It is not known if this species is hunted for the bushmeat trade (Fahr and Mildenstein, 2016). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Congo. Native: Angola (Bergmans, 1988; Sanborn, 1950; Monadjem et al., 2010d: 552); Congo (Bergmans, 1988; Monadjem et al., 2010d: 552; Bates et al., 2013: 333). HABITAT: Although the natural history of this species is poorly known, it seems to be primarily a savanna species that may range into tropical moist forest (Mickleburgh et al., 2008h; IUCN, 2009). POPULATION: Structure and Density:- It is only known from a few specimens (Mickleburgh et al., 2008h; IUCN, 2009; Fahr and Mildenstein, 2016). Trend:- 2016: Unknown (Fahr and Mildenstein, 2016). 2008: Unknown (Mickleburgh et al., 2008h; IUCN, 2009). Figure 20. Distribution of Epomophorus grandis Epomophorus intermedius (Hayman, 1963) 1889. Epomophorus pusillus: Noack, Zool. Jb. Syst., 4: 206. - Comments: Not of Peters, 1868. Partim. See Kock (1987b: 220). 1953. Micropteropus grandis: Ellerman, Morrison-Scott and Hayman, Southern African mammals 1758 to 1951, 49. - Comments: Not of Sanborn, 1950. In part. See Kock (1987b: 220). 1956. Micropterus grandis: Leleup, Ann. Kon. Mus. Mid. Afr., Zool. Wetensch., 46: 76. Comments: Not of Sanborn, 1950. (Lapsus) *1963. Micropteropus intermedius Hayman, Publ. Cult. Comp. Diamantes Angola, Lisboa, 66: 100, fig. 8. Type locality: Angola: Lunda: Dundo [07 22 S 20 50 E] [Goto Description]. Holotype: BMNH 1962.2073: ad ♀, skull and alcoholic. Collected by: Dr. A. de Barros Machado; collection date: June - July 1953. 1969. Micropteropus pusillus: Kock, Abh. Senckenberg. naturforsch. Ges., 521: 24. - Comments: Not of Peters, 1868. In Part. See Kock (1987b: 220). 2019. Epomophorus intermedius: Van Cakenberghe, in: Wilson and Mittermeier, Handbook Mammals of the World. Vol. 9., 101. (Current Combination) ? Micropterus (sic) intermedius: (Name Combination, Lapsus) ? Micropterus intermedius: (Emendation) TAXONOMY: See Bergmans (1989) and Simmons (2005: 327). COMMON NAMES: Czech: kaloň Haymannův. English: Hayman's Lesser Fruit-bat, Hayman's Lesser Fruit Bat, Hayman's Dwarf Epauletted Fruit Bat, Hayman's Epauletted Fruit Bat. French: Microptère de Hayman, Roussette naine de Hayman. German: Haymans Kleiner Epaulettenflughund. CONSERVATION STATUS: Global Justification Assessment History Global As Micropteropus intermedius:- 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008ba; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004t; IUCN, 2004). 2000: DD (Hilton-Taylor, 2000). 1996: VU (Baillie and Groombridge, 1996). 1994: Rare (Groombridge, 1994). MAJOR THREATS: The threats to this species are not well known, but may include general habitat loss through ongoing 100 ISSN 1990-6471 deforestation (Mickleburgh et al., 2008ba; IUCN, 2009). recorded since the 1950s (Mickleburgh et al., 2008ba; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008ba) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further surveys are needed to locate populations of this species, and to record additional details on natural history and possible threats. Trend:- 2008: Unknown (Mickleburgh et al., 2008ba; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Congo (Democratic Republic of the). GENERAL DISTRIBUTION: This species has only been recorded from four localities in northern Angola (the type locality of Dundo) and southern Democratic Republic of the Congo (Luluabourg, Banana/Netonna, and Thysville). Native: Angola (Bergmans, 1989; Monadjem et al., 2010d: 554); Congo (The Democratic Republic of the) (Bergmans, 1989; Monadjem et al., 2010d: 554). POPULATION: Structure and Density:- It is currently only known from four specimens. Some of these records are old, and the species does not appear to have been Figure 21. Distribution of Epomophorus intermedius Epomophorus labiatus (Temminck, 1837) *1837. Pteropus labiatus Temminck, Monogr. Mamm, 2: 83, pl. 39, figs 1 - 3. Type locality: Sudan: Blue Nile province: Sennaar [15 35 N 33 38 E, 425 m] [Goto Description]. Lectotype: RMNH MAM.27342: juv ♀. Collected by: Dr. Paolo Emilio Botta. See Kock (1969a: 18). - Comments: Type locality originally given as "Abyssinia"; see Horácek et al. (2000: 96). Andersen (1912b: 530) mentions that Temminck (1835) had two specimens, one obtained in London without any indication about the country of origin, and a second from Paris, originating from the collections of M. Botta, the latter is the only one presently in the RMNH. - Etymology: From the Latin "labiatus" for lipped. 1842. Pteropus Schoënsis Rüppell, Mus. Senckenberg., 3 (2): 131. Type locality: Ethiopia: Shoa province: Shoa province [ca. 09 00 N 39 00 E] [Goto Description]. Holotype: SMF 4370: juv ♂, skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell; collection date: 1841. Old catalog II.A.13a; see Kock (1969a: 18); Mertens (1925: 20) mentions it as a juv ♀. - Comments: Andersen (1912b: 531) already mentioned that the mounted specimen is of doubtfull sex. 1860. Epomophorus labiatus: Tomes, Proc. zool. Soc. Lond., 28: 55 - 56. (Name Combination, Current Combination) 1864. Epomophorus anurus Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 31 (7): 12. Type locality: Sudan: Bahr-el-Ghazal province: Bongo [07 15 N 28 42 E]. Syntype: SMNS 1090: imm ♂, mounted skin and skull. Collected by: Martin Theodor von Heuglin; collection date: 1855. Dieterlen et al. (2013: 293) indicate that this specimen was collected in “Bellegas-Tal” [= Belegaz/Belghe], Abyssinia. Syntype: SMNS 1091: ad ♀, mounted skin and skull. Collected by: Martin Theodor von Heuglin; collection date: 1863/1864. Dieterlen et al. (2013: 293) indicate that this animal was collected in Bongo [= country of Bongo, southern part of the province of Bahr-el-Ghazal in the south of Sudan]. Syntype: SMNS 609/661: ad ♂, mounted skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 293) mention the skull bears number 609, and that the mounted specimen has 661 as additional number. - Comments: Thorn et al. (2009: 20) mention 06 40 N 29 40 E. Type locality African Chiroptera Report 2020 1877. 1877. 1877. 1899. 1936. 1950. 1960. 1966. 1971. 1974. 1978. 1988. 1996. 2019. ? 101 originally given as "Gebied zw. Wau am Djur River und dem Getti River im Westen", see Kock (1969a: 18). Two syntypes: SMNS 1090 (♂, imm) and SMNS 1091 (♀, ad). Pteropus (Epomophorus) anurus: Heuglin, Reise in Nordost Afrika, 2: 16 - 17. (Name Combination) Pteropus (Epomophorus) labiatus: Heuglin, Reise in Nordost Afrika, 2: 15 - 16. (Name Combination) Pteropus (Epomophorus) schovanus: Heuglin, Reise in Nordost Afrika, 2: 18. - Comments: Emendation of schoënsis (see Andersen, 1912b: 530; Allen, 1939a: 56; Kock, 1969a: 18). (Emendation) Ep[omophorus (Epomophorus)] doriae Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 54. Type locality: Eritrea: Bogos country [=Keren region] [ca. 15 45 N 38 20 E] [Goto Description]. Holotype: ZMB 10079: Collected by: Gerrard. Presented/Donated by: ?: Collector Unknown. Holotype: ZMB 5384/10079: ad ♀, skin and skull. Vendor Gerrard; see Kock (1969a: 18), and Turni and Kock (2008: 8) [skin: ZMB 5384 + skull: ZMB 10079]. Epomophorus labiatus minor: G.M. Allen and Lawrence, Bull. Mus. comp. Zool., 79 (3): 45. - Comments: (Locality suggestive of labiatus rather than Epomophorus minor Dobson, 1880). (Name Combination) Epomophorus reii Aellen, Rev. suisse Zool., 57 (3): 559. Type locality: Cameroon: Rei Bouba [ca. 08 37 N 14 09 E] [Goto Description]. Holotype: MHNC 32-8-970: ad ♀. Collected by: MSS Cameroun; collection date: 13 September 1947. Presented/Donated by: ?: Collector Unknown. Paratype: MHNC 32-8-969: imm ♀. Collected by: MSS Cameroun; collection date: 13 September 1947. Presented/Donated by: ?: Collector Unknown. - Comments: in part?. Epomophorus labiatus anurus: Harrison, J. East Afr. Nat. Hist. Soc., 23 (7) (104): 287. Publication date: December 1960. (Name Combination) Epomophorus crypturus: Hayman, Misonne and Verheyen, Ann. Kon. Mus. Mid. Afr., Zool. Wetensch., (8) 154: 23. - Comments: (in part: specimen IRSN 10677 from "Kilwezi"- label reads Kilubwezi). Not of Peters, 1852. Epomophorus sp. Hill and Morris, 29. Epomophorus labiatus labiatus: Largen, Kock and Yalden, Mon. Zool. ital., (N.S.) 16 (Suppl. 5): 221, 226. (Name Combination) Epomophevus anupus: Arata and Johnson, in: Pattijn, Ebola Virus Haemorrhagic Fever, 135. (Lapsus) Epomophorus gambianus gambianus: Bergmans, Beaufortia, 38 (5): 84. Epomophorus anarus: Kityo and Kerbis, J. East Afr. Nat. Hist., 85: 58. (Lapsus) Epomophorus labiatuc: Banerjee, Kulcsar, Misra, Frieman and Mossman, Viruses, 11 (1) 41: 6. Publication date: 9 January 2019. (Lapsus) Epomophorus cf. labiatus: TAXONOMY: Includes anurus and minor; see Claessen and De Vree (1991), but see also Bergmans (1988) and Simmons (2005: 322 - 323). Middle Eastern forms reviewed by Horácek et al. (2000). COMMON NAMES: Azande (DRC): Ndima. Chinese: 小 颈 囊 果 蝠 . Czech: kaloň sudánský. English: Little Epauletted Fruit-bat, Ethiopian Epauletted Fruitbat, Ethiopian Epauletted Fruit Bat. French: Petit épomophore, Epomophore d'Ethiopie, Roussette labiée, Roussette labiaire. German: Temmincks Epaulettenflughund. Italian: Epomòforo labiàto. Kinande (DRC): Apopo. Ukrainian: Малий еполетовий крилан [=Malyy epoletovyy krylan]. CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008I; IUCN, 2009; Taylor, 2016a). Assessment History Global 2016: LC ver 3.1 (2001) (Taylor, 2016a). 2008: LC ver 3.1 (2001) [includes E. minor (Mickleburgh et al., 2008I; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004d; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. 102 ISSN 1990-6471 MAJOR THREATS: There appear to be no major threats to this species as a whole. Some populations may be threatened by overharvesting for subsistence food (Mickleburgh et al., 2008I; IUCN, 2009; Taylor, 2016a). CONSERVATION ACTIONS: Taylor (2016a) supports Mickleburgh et al. (2008I) [in IUCN (2009)] who report that this species has been recorded from many protected areas. No direct conservation measures are currently needed for the species as a whole. Additional research is needed into the impact of possible overharvesting of animals on some populations. GENERAL DISTRIBUTION: Epomophorus labiatus is largely distributed in Central Africa and East Africa. It appears to range from northern Nigeria and Cameroon, into Chad, southern and central Sudan, Ethiopia and Eritrea, and from here south into eastern Democratic Republic of the Congo, Uganda, Rwanda and Burundi, Kenya and Tanzania (including the islands of Unguja [=Zanzibar] and Mafia [see O'Brien, 2011: 285]), being recorded as far south as Malawi, northern Mozambique and northern Zambia. There is one isolated record from coastal Congo, although the presence of the species in this country, and also in Chad and Nigeria, needs to be confirmed. It has been recorded from lowland sites up to elevations of over 2,000 m asl (2,200 asl at Gondar, Ethiopia) (Mickleburgh et al., 2008I; IUCN, 2009). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 436,697 km2. Senegal records are probably erroneous (see Bergmans, 1988), although Capo-Chichi et al. (2004: 161) do refer to these records. Native: Burundi; Cameroon; Congo (Bates et al., 2013: 333); Congo (The Democratic Republic of the) (Bergmans, 1988; Monadjem et al., 2010d: 552); Djibouti (Pearch et al., 2001: 392); Eritrea (Yalden et al., 1996); Ethiopia (Yalden et al., 1996); Kenya (Aggundey and Schlitter, 1984; Malawi (Ansell, 1986; Bergmans, 1988; Happold et al., 1988; Monadjem et al., 2010d: 552); Mozambique (Monadjem et al., 2010d: 552; Monadjem et al., 2010c: 376); Rwanda (Baeten et al., 1984); Saudi Arabia (Gaucher, 1992); Sudan (Koopman, 1975; Tanzania; Uganda (Kityo and Kerbis, 1996: 59); Yemen (Benda et al., 2011b: 28); Zambia (Ansell, 1973; Bergmans, 1988; Monadjem et al., 2010d: 552). Presence uncertain: Chad; Congo; Nigeria. SEXUAL DIMORPHISM: Dobson (1873: 247) already indicated that Temminck noticed the absence of shoulder-tufts in the female. MOLECULAR BIOLOGY: Karyotype - Ðulic and Mutere (1973b) recorded the diploid number as 2n = 36. HABITAT: This species is regularly recorded from woodland (including miombo woodland), savanna, bushland, arid or semi-desert grassland habitats and mosaics of these general habitat types. In East African coastal areas it has been recorded from mangroves (Mickleburgh et al., 2008I). HABITS: It has been found resting together in small groups of about a dozen animals (Mickleburgh et al., 2008I). DIET: Seltzer et al. (2013: table S2) provide an overview of the plant species found beneath bat feeding roosts. For E. labiatus this included Ficus natalensis Hochst (Natal Fig - Moraceae). POPULATION: Structure and Density:- This is a common species (Mickleburgh et al., 2008I; IUCN, 2009). Trend:- 2008: Stable (Mickleburgh et al., 2008I; IUCN, 2009). REPRODUCTION AND ONTOGENY: Bergmans (1979a), Kingdon (1974), and Kulzer (1958) [in Krutzsch (2000: 110)] indicate that the male reproductive cycle of E. labiatus [as E. anurus] is apparently in synchrony with the one from the females as there occurs a biseasonal testicular hypertrophy in West Africa. PARASITES: Nycteribiidae: Tripselia blainvillii (Leach 1817) single record (Haeselbarth et al., 1966: 113, host referred to as E. anurus). Teinocoptidae: Teinocoptes epomophori Rodhain, 1923 was reported by Fain (1967: 367) from an E. anurus from Astrida (=Butare), Rwanda and from an E. labiatus minor from Bukavu, DRC. VIRUSES: Willoughby et al. (2017: Suppl.) report the following viruses: Entebbe bat virus and Shamonda orthobunyavirus. In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against West Nile virus (WNV), Dengue 2 virus African Chiroptera Report 2020 (DENV-2), non-specific Flaviviruses, Babanki virus (BBKV), and Rift Valley fever virus (RVFV). Coronaviridae 28.6 % (10 out of 35) of the Kenyan E. labiatus tested positive for CoV in the study by Tao et al. (2017). Nziza et al. (2019: 156) reported the presence of Kenya bat coronavirus/BtKY56/BtKY55 in a rectal swap from a bat from Rwanda. 103 DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Burundi, Cameroon, Chad, Congo, Congo (Democratic Republic of the), Djibouti, Eritrea, Ethiopia, Ghana, Kenya, Malawi, Mozambique, Rwanda, Somalia, South Africa, South Sudan, Sudan, Tanzania, Uganda, Zambia, Zimbabwe. Flaviviridae Pegivirus (BPgv) - 1 out of 3 bats from Kenya examined by Quan et al. (2013: Table S5) were infected by clade k type Pegivirus. Drexler et al. (2012a: Suppl. Table S1) indicated that the single specimen they examined from the Democratic Republic of the Congo did not test positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. UTILISATION: Bobo and Ntumwel (2010: 232) report that, in the Korup area of Cameroon, "E. anurus" is being sold as a medicine on markets, since consuming it would render women fertile. Is hunted for the bushmeat trade (Taylor, 2016a). Figure 22. Distribution of Epomophorus labiatus Epomophorus minimus Claessen & De Vree, 1991 1837. Pteropus labiatus: Temminck, Mon. Mammal., 2: 83 - 84. - Comments: Partim. One of the type specimens, an adult ♂, which later disappeared (Claessen and De Vree, 1991: 216). - Etymology: From the scientific Latin male adjective labiàtus meaning "lipped", referring to the expansible lower lip (see Lanza et al., 2015: 36). 1861. Epomophorus labiatus: Tomes, Proc. zool. Soc. Lond., 29: 11. - Comments: Partim. 1899. Ep[omophorus (Epomophorus)] schoensis: Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 44, 53. - Comments: Partim. Not of Rüppell, 1842. 1930. Epomophorus minor: Zammarano, Le colonie Italiane, 146. - Comments: Not of Dobson, 1880. 1939. Epomophorus labiatus minor: G.M. Allen, Bull. Mus. comp. Zool., 83: 56. - Comments: Partim. Not of Dobson, 1880. 1971. Epomophorus labiatus labiatus: Hayman and Hill, Mammals of Africa: Chiroptera, 6 - 7. Comments: Partim. 1984. Epomophorus labiatus anurus: Aggundey and Schlitter, Ann. Carnegie Mus., 53 (5): 124. - Comments: Animals from Katilo. Not of Heuglin, 1864. *1991. Epomophorus minimus Claessen and De Vree, Senckenb. biol., 71: 216. Type locality: Ethiopia: Harar province: Shewa: Bahadu [10 05 N 40 37 E, 600 m]. Holotype: SMF 44238: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971; original number: 16194. Full grown female with well developed nipples (see Claessen and De Vree (1991: 216). Paratype: SMF 44232: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44233: sad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44234: juv ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44235: juv ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44236: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. 104 ISSN 1990-6471 Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44237: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44239: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44240: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44242: sad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 2015. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44243: juv ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 5 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44245: ad ♀, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44246: sad ♂, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44247: ad ♂, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44248: sad ♂, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44250: juv ♂, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44252: ad ♂, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 - 7 May 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44253: juv ♂, skull and alcoholic. Collected by: Dr. M.J. Largen; collection date: 3 May 1971. Presented/Donated by: ?: Collector Unknown. - Etymology: From the Latin superlative masculine adjective mìnimus meaning "least, smallest" referring to the relatively small size of the species (see Lanza et al., 2015: 41). (Current Combination) TAXONOMY: Included in E. minor by Bergmans (1988). Recognized as a full species by Claessen and De Vree (1991); Simmons (2005: 323). Based on multivariate analyses, Claessen and De Vree (1991) concluded that the type specimen of Epomophorus minor fitted within the variation displayed by E. labiatus and that this name should be considered a synonym of E. labiatus. Consequently, no valid name was available for the smallest form included in the genus Epomophorus at that time. For these specimens, they proposed the new name Epomophorus minimus. However, Brosset and Caubère (1960) provided a further analysis of the problem and rejected this view, and reinstated the name minor. The situation is currently still not completely resolved with both names being used by various authors. The systematics of the small-sized Epomophorus forms, therefore, still remains unclear and needs further investigation. Member of the gambianus species group. COMMON NAMES: Chinese: 侏颈囊果蝠. Czech: kaloň habešský. English: Least Epauletted Fruit-bat, East African Epauletted Fruit-bat, East African Epauletted Fruit Bat. French: Petit Epomophore, Epomophore nain. German: Winziger Epaulettenflughund. Italian: Epomòforo mìnimo. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008j; IUCN, 2009; Webala, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Webala, 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008j; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004j; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There appear to be no major threats to this species. It is locally threatened in parts of its range by deforestation of its habitat for timber and firewood (Mickleburgh et al., 2008j; IUCN, 2009; Webala, 2016). African Chiroptera Report 2020 CONSERVATION ACTIONS: Mickleburgh et al. (2008j) [in IUCN (2009)] report that E. minimus it is not known if the species is present in any protected areas. Some individuals have been captured at Lake Bogoria National Reserve, Kenya (Wechuli et al. unpublished data cited in Webala (2016)). Further studies are needed into the distribution, natural history and possible threats to this species (Webala, 2016). GENERAL DISTRIBUTION: Epomophorus minimus is an East African species which is present in Uganda, northern Tanzania, Kenya, Ethiopia and Somalia. 105 known from collection records, with no information on population trends (Webala, 2016). Trend:- 2016: Stable (Webala, 2016). 2008: Stable (Mickleburgh et al., 2008j; IUCN, 2009). VIRUSES: Drexler et al. (2012a: Suppl. Table S1) indicated that three out of six specimens they tested from the Democratic Republic of the Congo were positive for Rubulavirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Ethiopia, Kenya, Somalia, Tanzania, Uganda. Native: Ethiopia (Yalden et al., 1996, Benda et al., 2020: 2583); Kenya (Aggundey and Schlitter, 1984; Somalia (Funaioli and Simonetta, 1966); Tanzania; Uganda (Kityo and Kerbis, 1996: 58). HABITAT: Recorded from dry and arid savanna habitats. DIET: It is possible that this species has a specialised diet to persist in its arid habitat (Mickleburgh et al., 2008j). POPULATION: Structure and Density:- E. minimus is probably a common species (Mickleburgh et al., 2008j; IUCN, 2009), but patchy distribution may reflect insufficient sampling (Happold, 2013l). It was reported to be the second most abundant species, after Micropteropus pusillus, in northern Uganda (Kityo and Kerbis, 1996). Elsewhere it is only Figure 23. Distribution of Epomophorus minimus Epomophorus minor Dobson, 1880 *1880. Epomophorus minor Dobson, Proc. zool. Soc. Lond., 1879, IV: 715. Publication date: April 1880. Type locality: Tanzania: Zanzibar [ca. 06 10 S 39 12 E]. Holotype: BMNH 1879.9.12.4: juv ♂, skull and alcoholic. Exchanged from Surgon-General G.E. Dobson see Turni and Kock (2008). Paratype: ZMB 5550: ad ♀, skull and alcoholic. Presented/Donated by: George Edward Dobson. See Turni and Kock (2008: 8). Comments: Andersen (1912b: 532) furthermore mentions that paratypes were distributed by Dobson to various museums. Considered a valid species by Kock (1969a: 18 - 24), Koopman (1975: 363) and Ansell and Dowsett (1988: 27), but see Claessen and De Vree (1991). - Etymology: From the Latin "minor" (= smaller). (Current Combination) 1896. Epomophorus pusillus: Pousargues, Ann. Sci. Nat., (8) 3: 255. ? Epomophorus anurus: - Comments: Not of Heuglin, 1864. ? Epomophorus labiatus anurus - Comments: Not of Heuglin, 1864. ? Epomophorus labiatus minor: (Name Combination) ? Epomophorus labiatus: (Name Combination) ? Epomophorus minor minor: (Name Combination) TAXONOMY: Recognized as a full species by Simmons (2005: 323). COMMON NAMES: Czech: kaloň zanzibarský, kaloň menší. English: Minor Epauletted Fruit Bat. German: Zwerg- 106 ISSN 1990-6471 Epaulettenflughund. Yao (applied to all large bats). (Malawi): Lichinji CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Taylor, 2016b). Assessment History Global 2016: LC ver 3.1 (2001) (Taylor, 2016b). 2008: LC ver 3.1 (2001) [as part of E. labiatus] (Mickleburgh et al., 2008I; IUCN, 2009). 2004: LC ver 3.1 (2001) [as part of E. labiatus] (Mickleburgh et al., 2004d; IUCN, 2004). 1996: LR/lc [as part of E. labiatus] (Baillie and Groombridge, 1996). MAJOR THREATS: There appear to be no major threats to this species as a whole. Some populations may be threatened by overharvesting for subsistence food (Taylor, 2016b). Trend:- 2016: Stable (Taylor, 2016b). REPRODUCTION AND ONTOGENY: For Malawi, Happold and Happold (1990b: 564) reported that females are monotocous (having only one pup). The young were born during the late dry season and the wet season. PARASITES: Hirst (1923: 980) described a new mite species (Ancystropus æthiopicus) from an E. minor specimen from Zanzibar. UTILISATION: This species is harvested for the bushmeat trade (Taylor, 2016b). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Ethiopia, Kenya, Malawi, Mozambique, Nigeria, Somalia, Sudan, Tanzania, Zambia. CONSERVATION ACTIONS: Taylor (2016b) reports that this species has been recorded from many protected areas. No direct conservation measures are currently needed for the species as a whole. Additional research is needed into the impact of possible overharvesting of animals on some populations. GENERAL DISTRIBUTION: Ethiopia, Somalia, Sudan, Kenya, Rwanda, SE Dem. Rep. Congo, Zambia, Tanzania, Zanzibar, Uganda, Malawi (Simmons, 2005: 323). POPULATION: Density and Structure:- This is a common species (Taylor, 2016b). Figure 24. Distribution of Epomophorus minor Epomophorus pusillus Peters, 1868 1860. 1860. Epomophorus schoensis: Tomes, Proc. zool. Soc. Lond., 28: 55 - 56. Epomophorus schoënsis: Tomes, Proc. zool. Soc. Lond., 1860, I: 56. Publication date: February - May 1860. - Comments: Not of Rüppell, 1842. *1868. Epomophorus pusillus Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 870 (for 1867). Type locality: Nigeria: S Nigeria: Yoruba. Holotype: ZMB 3438: imm ♀, skull and alcoholic. Collected by: A. Mann; collection date: 1862. Presented/Donated by: F. von Krauss. Holotype designated by Bergmans (1989: 96, 99); see Turni and Kock (2008: 9). - Comments: For a discussion on the type locality, see Bergmans (1989). Grubb et al. (1998: 70) state that "Peters in effect renamed Epomophorus schoënsis Tomes and therefore Tomes's type locality [="Gambia" - fixed by Andersen, 1912b: 559] must stand, even though his syntypes are evidently no longer in existence". However, see Kock et al. (2002: 82) for a discussion on the type locality. - Etymology: From the Latin "pusillus" (=very small) (see Owen-Ashley and Wilson, 1998: 4). (Current Combination) 1974. Micropteropus pussilus: Ayensu, Ann. Missouri Bot. Gard., 61: 706. (Lapsus) African Chiroptera Report 2020 1978. 1990. 2007. ? ? ? 107 Micropterus pusillus: Arata and Johnson, in: Pattijn, Ebola Virus Haemorrhagic Fever, 135. (Lapsus) Micropteropus pusilis: King, Davies and Lawrie, Vet. Microbiol., 23 (1-4): 168. Publication date: June 1990. (Lapsus) Micropteropus prusillus: Lameed, Biodiversity, 8 (4): 8. (Lapsus) Epomophorus (Micropteropus) pusillus: (Name Combination) Micropteropus pusillus: (Name Combination) Nanonycteris veldkampi: - Comments: Not of Jentink, 1888. TAXONOMY: See Bergmans (1989), Owen-Ashley and Wilson (1998) and Simmons (2005: 327). protected areas. In general, no direct conservation measures are currently needed for this species as a whole. COMMON NAMES: Castilian (Spain): Murciélago Enano. Chinese: 非洲小狐蝠. Czech: kaloň nigerijský. English: Peters' Lesser Epauletted Fruit Bat, Peters' Lesser Fruit-bat, Peters's Dwarf Epauletted Fruit Bat, Dwarf Epauletted Fruit Bat. French: Petit microptère, Roussette naine de Peters. German: Peters' Kleiner Epaulettenflughund, Peter's Zwergepaulettenflughund. Kiluba (DRC): Mulima. Bakwo Fils and Kaleme (2016a) further recommends that studies are needed in the species home range countries to have indications on population trends and details on its ecology. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) [as Micropteropus pusillus] in view of its wide distribution, presumed large population, it occurs in a number of protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008q; IUCN, 2009; Bakwo Fils and Kaleme, 2016a). Assessment History Global 2016: LC ver 3.1 (2001) [as Micropteropus pusillus] (Bakwo Fils and Kaleme, 2016a). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008q; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004s; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). MAJOR THREATS: There appear to be no major threats to this species as a whole (Mickleburgh et al., 2008q; IUCN, 2009). It is locally threatened by habitat loss and degradation. In Equatorial Guinea, local people eat E. pusillus (Fa, 2000). This could lead populations to decline (Bakwo Fils and Kaleme, 2016a). CONSERVATION ACTIONS: Bakwo Fils and Kaleme (2016a) supports Mickleburgh et al. (2008q) [in IUCN (2009)] who report that in view of the species wide range it seems probable that it is present in many GENERAL DISTRIBUTION: Distributed in West Africa, Central Africa and East Africa. It ranges from Senegal and The Gambia, throughout much of West Africa to Cameroon, from here it is distributed into Equatorial Guinea (Rio Muni), Gabon, Congo, Central African Republic, the Democratic Republic of the Congo, Sudan, Uganda and western Ethiopia and Kenya. It is found as far south as central Angola and Zambia. Native: Angola (Hayman, 1963; Crawford-Cabral, 1989; Bergmans, 1989; Lopes and CrawfordCabral, 1992; Monadjem et al., 2010d: 554); Benin (Capo-Chichi et al., 2004: 161); Burkina Faso (Kangoyé et al., 2015a: 605); Cameroon; Central African Republic (Morvan et al., 1999: 1195); Chad; Congo (King and Dallimer, 2010: 66); Congo (The Democratic Republic of the) (Schouteden, 1944; Hayman et al., 1966; Dowsett et al., 1991: 259; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 554); Côte d'Ivoire (Heim de Balsac, 1934b: 24); Equatorial Guinea; Ethiopia; Gabon; Gambia; Ghana; Guinea (Fahr and Ebigbo, 2003: 128; Denys et al., 2013: 281; Decher et al., 2016: 261); Guinea-Bissau (Monard, 1939; Bergmans, 1989; Rainho and Ranco, 2001: 28); Kenya; Liberia (Fahr, 2007a: 103); Mali (Meinig, 2000: 104); Niger; Nigeria; Rwanda; Senegal; Sierra Leone; Sudan; Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 59); Zambia (Ansell, 1974; Monadjem et al., 2010d: 554). Presence uncertain: Burundi. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Haiduk et al. (1980; 1981: 226) reported 2n = 35, FN = 64, a subtelocentric X and two Y chromosomes for males from Cameroon (XY1Y2) (see also Nesi et al., 2011: 552). Protein / allozyme - Unknown. 108 ISSN 1990-6471 ROOST: E. pusillus roosts singly or in pairs in the dense leave cover of shrubs and trees (Kunz, 1996: 44). In Ghana, Ayivor et al. (2017: 3) found these bats to roost together with E. gambianus, where pusillus occupied the lower parts of the trees. DIET: Ayensu (1974) reports the following plant species being visited by "Micropteropus pusillus" during foraging: Kigelia africana (Sausage tree), Anacardium occidentale (Cashew tree), Psidium guajava (Guava tree), and Spathodea campanulata (African Tulip tree). Seltzer et al. (2013: table S2) provided an overview of the plant species found beneath bat feeding roosts. For "Micropteropus pusillus" these include Milicia excelsa (Welw.) C. C. Berg (African Teak - Moraceae) and Psidium guajava L. (apple guava - Myrtaceae). PREDATORS: Mikula et al. (2016: Supplemental data) mention the African harrier-hawk (Polyboroides typus Smith, 1829) as diurnal avian predator. POPULATION: Structure and Density:- This is generally considered to be one of the most common African bats (Mickleburgh et al., 2008q; Bakwo Fils and Kaleme, 2016a). Large numbers are often recorded from fruit trees (Mickleburgh et al., 2008q; IUCN, 2009). Epomophorus pusillus is seldom gregarious and mostly roosts alone or in two, but may be found in groups of up to ten individuals (Bakwo Fils and Kaleme, 2016a). These groups are usually wellspaced throughout the roosting site. Trend:- 2016: Stable (Bakwo Fils and Kaleme, 2016a). 2008: Stable (Mickleburgh et al., 2008q; IUCN, 2009). REPRODUCTION AND ONTOGENY: Thomas and Marshall (1984) reported that females mated at the age of six months and gave birth at 12 months. Males reached puberty at the age of seven months. They also found that there is a clear relation between the bimodal births and the bimodality of the rainfall. 11 out of 18 females reported by Bergmans (1979a: 177) from Pointe Noire and Loandjili (Congo) were pregnant between 25 November and 13 December. In Sierra Leone, Weber et al. (2019: 19) found two pregnant females on 30 and 31 March and five lactating on 6 and 7 April and 28 May. MATING: Haft (2002) [in Nesi et al. (2011: 552)] indicates that the male's courting call has a frequency of 2800 Hz. PARASITES: Held et al. (2016: 119) reported one out of 26 examined Gabonese E. pusillus specimens to be infected by the bacterium Staphylococcus aureus. Schaer et al. (2013a: 17416) report the presence of hemosporidian parasites of the genus Hepatocystis in 68 out of 72 investigated bats. In the Amurum forest reserve (Nigeria), Atama et al. (2019: 1550) found Hepatocystis parasites in 42 % of the examined Micropteropus pusillus bats. Hirst (1923: 980) indicates that a specimen from Gambia was the host for a mite that was probably referrable to Ancystropus æthiopicus Hirst, 1923. Fain (1967: 367) reported Teinocoptes epomophori Rodhain, 1923 (Acari: Teinocoptidae) from a "Micropteropus pusillus" specimen from Boma, DRC. He also reported T. auricularis Fain, 1959 on this host species from Kabinda, Boma and Beno, DRC. VIRUSES: Willoughby et al. (2017: Suppl.) report the following viruses: Lagos bat lyssavirus, Rift Valley fever phlebovirus, Zaire ebolavirus. Coronaviridae - Coronaviruses Maganga et al. (2014a: 5) tested 533 animals from Congo and found only two testing positive. Anthony et al. (2017b: Suppl.) mention the betacoronavirus Kenya_CoV_BtKY56. Joffrin et al. (2020: 7) identified the Congolese viruses as Beta-D coronavirus. Filoviridae - Filo viruses Ebolavirus Muyembe-Tamfum et al. (2012: 9) refer to Pourrut et al. (2005) who found ZEBOV-specific antibodies. Marburg Towner et al. (2007) tested 149 individuals from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and 19 individuals for antiMarburg virus IgG antibodies by ELISA; no positives were found. Nairoviridae Orthonairovirus African Chiroptera Report 2020 Only two of the 100 specimens from Gabon and Congo tested by Müller et al. (2016: 3) were positive for Crimean Congo hemorrhagic fever virus (CCHFV). Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that none of the 152 specimens they examined from the Democratic Republic of the Congo, the Central African Republic, Gabon, Congo, and Ghana tested positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. Phenuiviridae Phlebovirus de Jong et al. (2011: 11) and Luis et al. (2013: suppl.) report the occurrence of Rift Valley fever virus. Rhabdoviridae Lyssavirus - Rabies related viruses 2006 in Nigeria, Dzikwi et al. (2010: 269) tested 55 brains by direct fluorescent antibody (DFA) and mouse inoculation test (MTIT) and all tested negative for lyssavirus antigens. Lagos bat virus (LBV) - Sureau et al. (1977) isolated Lagos bat virus in African Central Republic, from where Greenbaum and Carr (2005: 168) also reported monoclonal antibodies. In 2006, 21 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), none tested positive for neutralizing anitibodies (Dzikwi et al., 2010: 269). Mokola virus (MOKV) - King et al. (1990: 168) found monoclonal antibodies for MOKV in bats from the Central African Republic. They also found antibodies for Duvenhage and Denmark bat virus. In 2006, 21 individuals from Nigeria were serum tested by a modified rapid fluorescent focus 109 inhibition test (RFFIT), where none tested positive for neutralizing anitibodies (Dzikwi et al., 2010: 269). In their overview table, Maganga et al. (2014a: 8) report the following viruses were found on E. pusillus: Lagos Bat Virus (LBV), Coronavirus, Zaire Ebola Virus (ZEBOV), Marburg virus (MBGV), Rift Valley Fever virus (RVF). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Burkina Faso, Burundi, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Mali, Nigeria, Rwanda, Senegal, Sierra Leone, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia. Figure 25. Distribution of Epomophorus pusillus Epomophorus wahlbergi (Sundevall, 1846) 1846. Pteropus Haldemani Hallowell, Proc. Acad. nat. Sci. Philad., 3 (3): 52. Publication date: prior to July 1846. Type locality: Liberia [Goto Description]. - Comments: Holotype; ad ♀, SA, in the collection of the Academy of Natural Sciences of Philadelphia, see Andersen (1912b: 524), who further mentions "'W. Africa,' obtained from 'Dr. Goheen, Physician to the American Colonization Society' (from this it might be supposed that the specimen came from Liberia; there can be little doubt, however, that it was obtained elsewhere in W. Africa; no form of Epomophorus with the palate-ridges of the walhbergi type is known to inhabit the Guinea coast west of the Cameroons)." Allen (1939a: 56) suggests that the specimen might possible have come from Liberia. Considered a valid species by Schutt and Simmons (1998: 27). *1846. Pteropus Wahlbergi Sundevall, Öfvers. kongl. Sv. Vet.-Akad. Förhandl., 3 (4): 118. Type locality: South Africa: KwaZulu-Natal: Port Natal [=Durban], near [29 52 S 31 00 E] [Goto Description]. - Comments: Type locality originally: "Prope Port-Natal et in Caffraria interiore occisus" see Acharya (1992: 1). As to the publication date, Andersen (1912b: 526, footnote) mentions "Sundevall's paper was read before the Stockholm Academy on May 13th, 1846, and presumably published in the latter half of May or in June (the 'Öfversigt' 110 ISSN 1990-6471 1870. 1870. 1899. 1899. appears to have been issued regularly for every month in which the Academy held a meeting). Hallowell's description of Pt. haldemani was passed by the Publication Committee on May 26th, 1846, and printed in the May-June issue of the Proceedings of the Philadelphia Academy, which cannot have come out before, in the earliest, in July, since it contains a report of a meeting held on June 30th. The balance of evidence would seem, therefore, to be in favour of the date priority of Pt. wahlbergi". Andersen (1912b: 527) also mentions "Type (by selection) in the Riksmuseum, Stockholm, a well-preserved mounted adult male, skull separate (nearly perfect), Tugela River, Natal, 27 Nov. 1843, J. Wahlberg coll., Reg. No. 1040.". - Etymology: Named after Johan August Wahlberg (1810 - 1856), explorer, hunter and outstanding collector of zoological material in Southern Africa. He was killed in 1856 by a wounded elephant near Savuti in northern Botswana (see Gyldenstolpe, 1933; Taylor, 2005, Lanza et al., 2015: 48). [Epomophorus macrocephalus] var. unicolor Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 125 [Goto Description]. Epomophorus macrocephalus var. unicolor Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 125. Type locality: Mozambique: Zambesi River: Shupanga [=Chupanga] [18 02 S 35 37 E]. Holotype: BMNH 1864.8.16.1: sad ♀, skin and skull. Collected by: Dr. Sir John Kirk; collection date: June. Ep[omophorus (Epomophorus)] neumanni Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 43, 50. Type locality: Kenya: Mombassa [04 03 S 39 40 E] [Goto Description]. Lectotype: ZMB 9967: ad ♂, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: about 1880. Lectotype designated by Andersen (1912b: 525); see Turni and Kock (2008). Paralectotype: ZMB 53873: ♀, skull and alcoholic. Collected by: Michael Rodgers Oldfield Thomas; collection date: 1898. Takaungu, Kenya. Paralectotype: ZMB 53897: ♀, skull and alcoholic. Collected by: Michael Rodgers Oldfield Thomas; collection date: 1898. Takaungu, Kenya. Paralectotype: ZMB 5428: ♀, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: ca. 1876. Mombassa. Paralectotype: ZMB 54367: ♂, skull and alcoholic. Collected by: Michael Rodgers Oldfield Thomas; collection date: 1898. Takaungu, Kenya. See Turni and Kock (2008: 9). Paralectotype: ZMB 5597: juv ♂, skull and alcoholic. Collected by: Fischer. Malini, Kenya. Paralectotype: ZMB 9968: juv ♀, skull and alcoholic. Collected by: Prof. Oscar Rudolph Neumann; collection date: January 1898. - Comments: Mombasa is the locality of the lectotype: see Allen (1939a: 57). Acharya (1992: 1) mentions "No type originally designated. Type locality restricted to "Mombassa, British East Africa (=Kenya) by Andersen (1912b: 524)". Matschie mentioned 14 May 1899 in his introduction. Ep[omophorus (Epomophorus)] stuhlmanni Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 50. Type locality: Tanzania: Usaramo: Vikindo [06 59 S 39 17 E, 120 m] [Goto Description]. Lectotype: ZMB 9982/9983: ad ♂, skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann; collection date: 11 January 1894. Lectotype designated by Andersen (1912b: 527); skull damaged [skin: ZMB 9983 + skull: ZMB 9982]. Paralectotype: ZMB 10000: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Zanzibar. Paralectotype: ZMB 10001: ♀, skull and alcoholic. Collected by: Neumann. Zanzibar. Paralectotype: ZMB 10002: ♂, skull and alcoholic. Collected by: Neumann. Zanzibar. Paralectotype: ZMB 2957: ♂, skull and alcoholic. Collected by: Wessel. Zanzibar. Paralectotype: ZMB 67054: ♀, skull only. Collected by: Dr. Franz Ludwig Stuhlmann; collection date: 1894. Dar es Salaam. Paralectotype: ZMB 67055: ♀, skull only. Collected by: Dr. Franz Ludwig Stuhlmann; collection date: 1894. Dar es Salaam. Paralectotype: ZMB 67056: skull only. Collected by: Dr. Franz Ludwig Stuhlmann; collection date: 1894. "Ushindo" [= ?], Tanzania. Paralectotype: ZMB 9252: ♀, skin and skull. Collected by: Fülleborn. Lindi, Tanzania. See Turni and Kock (2008: 9). Paralectotype: ZMB 9978: ♀, skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9979: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9980: ♂, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9981: ♂, skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9984: skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. African Chiroptera Report 2020 1899. 1990. 2012. 2013. 2016. 2017. ? ? ? ? ? 111 Paralectotype: ZMB 9986: skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9988: skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9989: ♂, skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Vikindo. Paralectotype: ZMB 9990: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Dar es Salaam. Paralectotype: ZMB 9991: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Dar es Salaam. Paralectotype: ZMB 9992: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Dar es Salaam. Paralectotype: ZMB 9993: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Dar es Salaam. Paralectotype: ZMB 9996: ♀, skull and alcoholic. Collected by: Dr. Franz Ludwig Stuhlmann. Zanzibar. Paralectotype: ZMB 9998: ♂, skin and skull. Collected by: Dr. Franz Ludwig Stuhlmann. Dar es Salaam. - Comments: The "type locality" is the locality of the lectotype: see Allen (1939a: 56). Acharya (1992: 1) mentions "No type originally designated. Type locality restricted to Vikindo, Usaramo, German East Africa [=Tanzania] by Andersen (1912b: 527)." In part (see Allen, 1939a: 56). Matschie mentioned 14 May 1899 in his introduction. Ep[omophorus (Epomophorus)] zenkeri Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 44, 46. Type locality: Angola: Cabinda: Tschintschoscho [= Chinchoxo] Station [05 05 S 12 02 E] [Goto Description]. Lectotype: ZMB 9974/9975: ad ♀, skin only. Collected by: Falkenstein; collection date: 1874. Lectotype designated by Andersen (1912b: 524): 8 paralectotypes designated; see Turni and Kock (2008) [skin: ZMB 9974 + skull: ZMB 9975]. Paralectotype: ZMB 3632: ♀, skull and alcoholic. Angola. Ex Mus Lisbon. Paralectotype: ZMB 4801: ♂, skull and alcoholic. Collected by: Falkenstein. Chinchoxo [=Loango Coast]. Paralectotype: ZMB 6129: ♂, skull and alcoholic. Collected by: Büttner. Gabon. Paralectotype: ZMB 9969: ♀, skull and alcoholic. Collected by: von Melchow. Malandje [=Malange], Angola. Paralectotype: ZMB 9970: ♂, skull and alcoholic. Collected by: von Melchow. Malandje [=Malange], Angola. Paralectotype: ZMB 9971: ♀, skull and alcoholic. Collected by: von Melchow. Malandje [=Malange], Angola. Paralectotype: ZMB 9976: ♂, skull and alcoholic. Collected by: Falkenstein. Chinchoxo. Paralectotype: ZMB 9977: ♂, skull and alcoholic. Collected by: Falkenstein. Chinchoxo. See Turni and Kock (2008: 10). - Comments: Allen (1939a: 56) mentioned the lectotype from Chinchoxo, Cabinda, Angola. Acharya (1992: 1) mentions "No type originally designated. Type locality restricted to Chinchoxo, Cabinda, Angola by Andersen (1912b: 524)." Matschie mentioned 14 May 1899 in his introduction. Epomophorus walbergi: King, Davies and Lawrie, Vet. Microbiol., 23 (1-4): 168. Publication date: June 1990. (Lapsus) Epomophorus walhbergi: Monadjem and Reside, Afr. Zool., 47 (2) 321. (Lapsus) Epomophorus angolensis: Van Cakenberghe and Seamark, African Chiroptera Report, 760. - Comments: Not of Gray, 1870 (in part). The specimens not from Angola or Namibia. Epomophorus wahlbergyi: Bahlman, Price-Waldman, Lippe, Breuer and Swartz, J. Anat., 229 (1): 116. Publication date: 1 March 2016. (Lapsus) Epomorphorus wahlbergi: Banyard and Fooks, Microbiol. Aust., 38 (1): 18. Publication date: March 2017. (Lapsus) Epomophorus crypturus: - Comments: Not of Peters, 1852. Epomophorus gambianus: - Comments: Not of Ogilby, 1836. Epomophorus wahlbergi haldemani: (Name Combination) Epomophorus wahlbergi wahlbergi: (Name Combination) Epomophorus wahlbergi: (Name Combination, Current Combination) 112 ISSN 1990-6471 TAXONOMY: IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004f; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Schoeman et al., 2016d). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: There appear to be no major threats to this species as a whole (Mickleburgh et al., 2008k; IUCN, 2009). Figure 26. Epomophorus wahlbergi from Durban, KwaZulu Natal, South Africa. Reviewed by Acharya (1992). (2005: 323). See Simmons COMMON NAMES: Afrikaans: Wahlberg se vrugtevlermuis, Wahlbergwitkolvrugtevlermuis. Castilian (Spain): Zorra Voladora. Chinese: 韦 氏 颈 囊 果 蝠 . Czech: kaloň Wahlbergův. English: Wahlberg's Epauletted Fruit-bat, Wahlberg's Epauletted Fruitbat. French: Epomophore frugivore de Wahlberg, Roussette à épaulettes de Wahlberg. German: Wahlbergs Epaulettenflughund. Italian: Epomòforo di Wàhlberg. Portuguese: Morcego frugivoro de Wahlberg. SIMILAR SPECIES: Taylor and Monadjem (2008: 24) provided data which enabled them to separate E. wahlbergi from E. crypturus. In the first species, the maxillary width is always larger than in the latter (♂♂: > 14 mm; ♀♀: > 13 mm). Adams and Snode (2015) analysed the male mating calls of two groups of epauletted bats (E. wahlbergi and E. crypturus) in the Kruger National Park, and found differences in mean fundamental frequency, mean high frequency, mean low frequency, mean bandwidth, and mean call slope. They hypothesize that these differences may be used to avoid cross-mating between species. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008k; IUCN, 2009; Shoeman, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Shoeman, 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008k; However, MacEwan (2016: 3) reports on E. walhbergi fatalities as a result of encounters with wind turbine blades. While Shoeman (2016) reports that this species is threatened by loss of habitat because of dune mining in KwaZulu-Natal, South Africa. Over the past 5 years a decline in some populations in KwaZulu-Natal have been observed, with the cause suspected to be loss of coastal forest due to dune mining as well as extensive drought. It is also possible that the Lagos bat virus may be affecting the population as a number of bats caught from Durban and the south coast in South Africa have tested seropositive for the virus (Shoeman, 2016). CONSERVATION ACTIONS: Shoeman (2016) supports Mickleburgh et al. (2008k) [in IUCN (2009)] who report that it is present in many protected areas and no direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Epomophorus wahlbergi is found in Central Africa, East Africa and southern Africa (it is broadly distributed across southern Africa). It has been recorded from Cameroon (Aellen, 1952), Equatorial Guinea (Rio Muni), Gabon, Congo and Angola in the west, through southern Democratic Republic of the Congo and Rwanda, being distributed in East Africa from Uganda, Kenya and southern Somalia in the north, through Tanzania (including the islands of Unguja [=Zanzibar], Pemba, and Mafia [see O'Brien, 2011: 285]), Zambia, Malawi and Mozambique into Zimbabwe, eastern and southern South Africa and Swaziland. There is a need to confirm the presence of this species in Cameroon and Equatorial Guinea. It ranges from sea level to around 2,000 m asl (Mickleburgh et al., 2008k; IUCN, 2009). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,327,859 km 2. African Chiroptera Report 2020 In South Africa, its distribution is strongly associated with annual precipitation (Babiker Salata, 2012: 49). Liberian records are probably erroneous. Native: Angola (Bergmans, 1988; CrawfordCabral, 1989; Monadjem et al., 2010d: 552); Botswana; Burundi; Congo (Bates et al., 2013: 333); Congo (The Democratic Republic of the) (Schouteden, 1944; Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 552); Gabon; Kenya; Malawi (Happold et al., 1988; Bergmans, 1988; Happold and Happold, 1997b: 815; Monadjem et al., 2010d: 554); Mozambique (Smithers and Lobão Tello, 1976; Bergmans, 1988; Monadjem et al., 2010d: 554; Monadjem et al., 2010c: 376); Rwanda; Somalia; South Africa (Monadjem et al., 2010d: 553); Swaziland (Monadjem et al., 2010d: 553); Tanzania (Stanley and Goodman, 2011: 37); Uganda; Zambia (Ansell, 1978; Bergmans, 1988; Cotterill, 2002b: 6; Monadjem et al., 2010d); Zimbabwe (Monadjem et al., 2010d: 553). Presence uncertain: Cameroon; Equatorial Guinea, Namibia (Cotterill, 2004a: 260). DENTAL FORMULA: At birth, the following dentition is present: I 2/2 C 1 / 1 P 1/1 = 16 The adult dental formula is: I 2/2 C 1/1 P 2/3 M 1/2 = 28 (Sowler, 1980: 113). Sowler (1980: 113) also reports that the second deciduous premolars appear between the second and third week, making a total of 20 teeth. All of the permanent teeth are present between week 12 and 14. DETAILED MORPHOLOGY: Tongue - Mqokeli and Downs (2012b examined eight E. wahlbergi from Pietermaritzburg, South Africa where all studied had an elongated, muscular tongue of 4.4 ± 0.33 cm (mean ± SD) long and 1.1 ± 0.4 cm (mean ± SD) wide. The anterior tip of the tongue of each was narrower compared with the broader posterior part of the tongue. The anterior tip of the tongues was 0.2 ± 0.05 cm thick with a width of 0.7 ± 0.10 cm (mean ± SD). The mid-region of the tongues had a thickness of 0.6 ± 0.05 cm and a width of 1 ± 0.20 cm (mean ± SD). The posterior regions of the tongues were 0.7 ± 0.00 cm thick and 1.5 ± 0.10 cm wide. The dorsal surface of the tongues possessed two types of mechanical papillae, filiform and conical papillae, and two types of gustatory papillae, fungiform and circumvallate 113 papillae, which varied in distribution. Filiform papillae were highly abundant and occurred on the entire surface of the tongue. These papillae were posteriorly directed, with the small filiform papillae situated at the anterior tip of the tongue. Larger filiform papillae were present as a band posterior to these in the anterior to mid-regions of the tongue. Small filiform papillae with fungiform papillae occurred on either side of the giant filiform papillae band. The posterior part of the E. wahlbergi tongue (behind the giant filiform papillae) had elongated basket-like filiform papillae. These had large projections and occupied most of the posterior region of the tongue. Fungiform papillae occurred among these filifor papillae. The elongated basket-like filiform papillae were symmetrically arranged with respect to the axis of the tongue. The filiform papillae transformed into conical papillae on the left and right side of the posterior base of the tongue and pointed inwards. Short conical papillae occurred on the midportion of the posterior region of the tongue. Fungiform papillae were scattered among the filiform papillae, except in areas of giant filiform and conical papillae. At the posterior end of the tongue occurred a triangular arrangement of three circumvallate papillae directed toward the pharynx surrounded by a deep groove. Brain - Chawana et al. (2013: 160) indicate the average brain mass for 2 specimens was reported as 1.81 g. Physiology - Minnaar et al. (2014b: 475) studied the evaporative water loss and indicate that they can not substantiate previous findings of 5 to 10 % of the gas exchanges being done over the wing membrane. FUNCTIONAL MORPHOLOGY: Makanya and Mortola (2007: 687) and Kovalyova (2014: 326) indicate that the body skin epidermis for this species thick (61 ± 3 µm, SEM), the stratum corneum alone taking a third of it (21 ± 3 µm). The wing membrane was 27.8 ± 3.1 µm thick, of which the epidermis took 9.8 ± 0.7 (9.1 - 10.5) µm, with a stratum corneum (horny layer) of 4.1 ± 0.3 (3.7 4.4) µm. Makanya and Mortola (2007: 687) conclude that the wing web has structural modifications that permit a substantial contribution to the total gas exchange. Coimbra et al. (2016: 191) mention that the eye in this species contains about 200,000 retinal ganglion cells. The minimum angle of resolution was approximately 0.167° (ca. 5-6 mm at 1 m distance). 114 ISSN 1990-6471 SEXUAL DIMORPHISM: Adams and Snode (2013: 54) indicate that males are larger than females and have bulldog-like facial pouches that inflate during mate-calling. MOLECULAR BIOLOGY: DNA - Unknown Karyotype - 2n = 36, FN = 68 (Ðulic and Mutere, 1973b; Peterson and Nagorsen, 1975; Ðulic and Mutere, 1977). Ðulic and Mutere (1973b; 1977) reported BA = 34, whereas Peterson and Nagorsen (1975) reported BA = 36 as they included the X (no male studied). Protein / allozyme - Unknown. HABITAT: Although E. wahlbergi was found in bushed Acacia steppe, Lönnberg (1912: 27) doesn't regard this species as a steppe mammal, since a forest of tall Acacias and other trees was nearby. HABITS: Wickler and Seibt (1976) suggest that male E. wahlbergi's call out to space out other males, but also to attract females. Funakoshi et al. (1995: 504) indicate that their communication calls consist of several harmonics covering the 2 - 8 kHz range, with a duration of 200 - 300 msecs, which also contain a slight click at the beginning. In a study at the University of KwaZulu-Natal, Downs et al. (2015: 7) found that in winter bats generally slept for longer periods of time, and more often than in summer, and that no bats were sleeping when the ambient temperature was above 35°C. During the winter season, the percentage of sleeping bats increased between 9 and 12 o'clock as Ta increased. In both seasons, more bats were sleeping with both eyes closed than with only one open. During the first hours after their nightly activity, sleep with both eyes closed was more often observed, but this diminished later on during the day, with one eye open sleep reaching its peak around 10h00. ROOST: Day roosts are often located in the dense foliage of riverine or gallery forests, but sometimes caves are used too. Three to six individuals roost in small groups, and roosts are switched every five or six days, sometimes only for a few metres (Kunz, 1996: 44). Wickler and Seibt (1976) [in McCracken and Wilkinson (2000:350)] found that E. wahlbergi roosts in palm leaves or thatch roofs in mixed sex colonies, containing from three to over 100 animals, where the individuals are spaced at least 2.5 cm apart (except for female-young pairs). In the Durban area, Taylor et al. (1999: 66) reported E. wahlbergi to roost along the undersurface midribs of royal palm (Roystonia elata (Kunth) O.F. Cook) leaves, and sometimes under eaves of double-storey houses or in recesses under a concrete road bridge. The bats also regularly change their day roost (see Rollinson et al., 2014: 173), e.g. to reduce the distance from the roost site to their feeding sites. During the winter they changed their roosting sites more often than during the summer (p. 179). These authors also reported that the roost temperatures are generally higher than the ambient temperature, and that roosts in man-made structures had usually a higher temperature than those in natural vegetation (e.g. in Bougainvillea, Borassus, Trema orientalis). DIET: Voigt et al. (2011: 355) found E. wahlbergi to be the second most important seed disperser for syringa fruits (Melia azedarach), which is an alian invasive tree species in KwaZulu Natal. Jordaan et al. (2011: 961) fed captive specimens with fruits of Psidium guajava L. (Myrtaceae) and Melia azedarach L. (Meliaceae) during the month of May, and with those of Eriobotrya japonica (Thunb.) Lindl. (Rosaceae) and Morus alba L. (Moraceae) during September. Solanum mauritianum Scopoli (Solanaceae) and Cinnamomum camphora (L.) J. Presl. (Lauraceae) fruits were also made available to the bats, but were not eaten by any of the individuals. In Congo, Kipalu (2009: 16) reported E. wahlbergi visiting/consuming fruits of the Borassus palm (Borassus aethiopum), African peach (Nauclea diderichii), and Mango (Mangifera indica). Seltzer et al. (2013: table S2) provided an overview of the plant species found beneath bat feeding roosts. For E. wahlbergi these include Rauvolfia caffra Sond. (Quinine Tree Apocynaceae), Ficus natalensis Hochst (Natal Fig - Moraceae) and Syzygium cordatum Hochst. (water-berry tree - Myrtaceae). Govender et al. (2016: 315) found that E. wahlbergi prefered ripe figs over green ones. Wickler and Seibt (1976) [in Bonaccorso et al. (2014: 50)] reported E. wahlbergi feeding on leaves of Balanites wilsonianea (Balanitaceae) in East Africa. Bonaccorso et al. (2014: 50) indicated this might also be the case in the Kruger National Park. Based on data from captive feeding experiments Bonaccorso et al. (2014: 50) extrapolated that the Skukuza bat colony (133 African Chiroptera Report 2020 animals in 2007) would have dispersed 3.2 million fig seeds per night. Downs et al. (2011b: 148) indicate that E. wahlbergi's body mass was significantly higher in winter than in summer, which is possibly not attributable to an increase in body fat, but rather to excessive eating as result in increased metabolic costs. Rollinson et al. (2013: 340) found that in the urban environment of Pietermaritzburg, South Africa, the bats had a greater home range in winter than in spring, which was linked with the reduction of fruiting plant species. They also found (p. 343) that male home ranges were smaller than those of females (0.46 km2 versus 0.78 km 2). Their mean foraging distance was 1.45 ± 0.2 km in winter and 0.88 ± 0.08 km in spring. Namah et al. (2019: 2) expected E. wahlbergi to be the pollinator of sausage trees (Kigelia africana (Lam.) Benth.) in the Kruger Park (RSA), but could not find any proof of this. Nor could they confirm pollination by any other species of bat. Coleman and Downs (2012: 431) investigated the effect of sugar type and concentration on E. wahlbergi and suggest that these bats do not appear to have a selective pressure on the sugar composition of the fruit they eat. Mqokeli and Downs (2014: 161) performed some tests with equicaloric solutions (15 % glucose or 15 % sucrose with 0, 2.58, 5.68, 7.23 g soy protein/kg H2O). Volumetric intake of the respective glucose and sucrose solutions varied among individual bats, with total volumetric intake being highest for solutions with no protein. The authors concluded that this suggests E. wahlbergi has low protein dietary requirements, which may be a response to the low-protein fruit available in the wild. They do point out that the bats may supplement their diet by actively preying on insects, but this needs to be investigated. Blood plasma glucose concentrations – Epomophorus wahlbergi‘s blood plasma glucose concentration was lower (5.24 ± 0.38 mmol/l) at 18:00 before feeding and increased during/after feeding (8.19 ± 1.24 mmol/l) but bats appeared to regulate it within limits (Mqokeli and Downs, 2012a: 348). Glucose intake of E. wahlbergi is generally high irrespective of sugar concentration (Coleman and Downs, 2012; Downs et al., 2011a). Mqokeli and Downs (2012a: 349) found that blood plasma glucose concentration differed significantly with time. The greatest individual variation in blood plasma glucose concentrations were recorded at 24:00 possibly because of varying feeding and digestion rates (Mqokeli and Downs, 115 2012a: 350). Mqokeli and Downs (2012a: 349) found that blood plasma glucose concentrations were generally higher than the upper limit of the normal mammalian blood plasma glucose range. Mqokeli and Downs (2012a) showed diet variations in blood plasma glucose concentrations with an increase in concentrations during or directly after feeding. Mqokeli and Downs (2012a: 349) found that the body masses differed significantly at the respective blood sampling times. The lowest mean body mass (± S.E) was at 18:00 (99.69 ± 5.55 g) while the highest mean body mass was at 24:00 (105.05 ± 5.75 g). There was a significant difference in body mass between the 12:00 and 18:00 sampling times, and between the 24:00 and 18:00 sampling times. At 24:00 all bats had fed on the fruit provided and at 06:00 little fruit remained. The increase in body mass at 24:00 was about 6 %. Germination experiments performed with four E. wahlbergi specimens captured on Mount Kilimanjaro (Tanzania), and fed with fig fruit parts of Ficus sur trees resulted in an overal germination percentage of 49.3 % after 13 days (Helbig-Bonitz et al. (2013: 123). PREDATORS: Broadley (in Schätti, 1984: 340) reports that these bats were eaten by Gaboon vipers (Bitis gabonica) kept in captivity. Mikula et al. (2016: Supplemental data) mention the following diurnal avian predators: Dickinson's kestrel (Falco dickinsoni Sclater, 1864), Wahlberg's eagle (Hieraaetus wahlbergi (Sundevall, 1851)), Crowned eagle (Stephanoaetus coronatus (Linnaeus, 1766)); Trumpeter hornbill (Bycanistes bucinator (Temminck, 1824)). POPULATION: Structure and Density:- Common (Mickleburgh et al., 2008k; IUCN, 2009; Shoeman, 2016). Shoeman (2016) reporting that population numbers increasing. Trend:- 2016: Stable (Shoeman, 2016). 2008: Stable (Mickleburgh et al., 2008k; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 10.1 years. REPRODUCTION AND ONTOGENY: Close to the equator, this species has two birth periods in February-March and in OctoberDecember (Bergmans, 1979a; O'Shea and Vaughan, 1980), or an extended period from October to January (Wickler and Seibt, 1976; 116 ISSN 1990-6471 Bernard and Cumming, 1997). Further south in Malawi and Zambia this species has been described as 'aseasonally polyoestrus' with births from September to March (Happold and Happold, 1990b). In southern Africa, this species has been described as 'seasonally polyoestrus' (Taylor, 2005). Although Taylor (1998) reported that in the KwaZulu-Natal province of South Africa 'breeding occurs throughout the year with peaks in July and in the summer'. However, this statement was based on an unpublished dataset by C. Sapsford, and further details were not presented (Taylor, 2005) in [Skinner and Chimimba, 2005]). In Zimbabwe, gravid females have been recorded in June and December (Smithers and Wilson, 1979), suggesting an extended breeding season. By contrast, in Kruger National Park, South Africa, parturition has been recorded in NovemberDecember (Pienaar et al., 1980). In Swaziland, subadults were present throughout the year with two peaks (January-May and August-October), while lactating females were only present between December and May (Monadjem and Reside, 2012: 322). Lönnberg (1912: 47) mentioned that two of the three females captured on 20 March 1911 at the upper Luazomela River, Kenya were carrying young of approximately 6 cm. Stanley and Goodman (2011: 37) reported a pregnant female collected in the South Pare Mountains, Tanzania, on 20 July 1993. One female, taken by Bergmans (1979a: 174) in Congo on 21-22 November was lactating, while three others, taken between 28 November and 13 December, were pregnant. Szekely et al. (2015: Suppl.) report a gestation period of 165 days, and two litters per year. MATING: Wickler and Seibt (1976) [in Krutzsch (2000: 111)] indicate that male mating patterns also include courtship vocalizations (calling territories). Males also establish nocturnal display sites where females visit the males for mating (see McCracken and Wilkinson, 2000: 349). At Skukuza (Kruger National Park), Adams and Snode (2015: 4) found the following values for the male mating calls: mean fundamental frequency: 7.88 ± 0.11 kHz (first harmonic) followed by a second harmonic: 15.71 ± 0.23 kHz. Adams and Snode (2013: 54) investigated the male-male competition for females during the mating season and found (p. 58) that the most dominant individuals were calling nearest to a ripe fig tree, and therefore more likely to be noticed by a foraging female. None of the males they observed were calling from nonfruiting sycamore fig trees. Chaverri et al. (2018: 1942) point out the role of the retractile white shoulder tufts, which are erected during courtship by wing-flapping males. They also suggest that these movements may help spread pheromones. PARASITES: BACTERIA One out of five bats tested by Dietrich et al. (2016b: 3) in Pafuri (RSA) tested positive for Bartonella bacteria, and one out of two bats from Rocktail Bay (RSA) tested positive for Rickettsia, as did one out of 80 tested from Umbilo (RSA). gram-negative Helmick et al. (2004: 88) reported a novel Pasteurella-like organism found in lung tissue of two captive Wahlberg’s epauleted fruit bats, which were recently shipped, and died soon after release from a 30-day quarantine period. HAEMOSPORIDA Garnham (1948, 1950) [in Miltgen et al. (1977: 595)] described some liver schizonts and gametocytes, which he assigned to the genus Hepatocystis. ACARI Teinocoptidae: Fain (1967: 367) reported Teinocoptes epomophori Rodhain, 1923 from an E. wahlbergi haldemani specimen from Boma, DRC, and Teinocoptes auricularis from an E. wahlbergi from Zanzibar. Trombiculidae: Schoutedenichia (Schoutedenichia) cordiformis VercammenGrandjean, 1958 was reported by Stekolnikov (2018a: 101), who also reported (p. 153) Microtrombicula homopholis (Lawrence, 1949). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV: Müller et al. (2007b) tested between 1986 to 1999, two individuals from Mpumalanga Province, South Africa for antibody to SARS-CoV in sera, none were found to be positive. Four out of 63 (6.3 %) Kenyan bats tested by Tao et al. (2017: Suppl.) were positive for CoV. Rhabdoviridae Lyssavirus - Rabies related viruses Lagos bat virus (LBV): Isolated from South Africa (Crick et al., 1982: 211; King et al., 1990: 169; Meredith and Standing, 1981; Swanepoel, 2004; Markotter et al., 2006b; Hayman et al., 2011a: 88; Nadin-Davis et al., 2011: 238; Coertse et al., 2011: 251; Willoughby et al., 2017: Suppl.). Mokola virus (MOKV): King et al. (1990: 168) found monoclonal antibodies for MOKV in bats African Chiroptera Report 2020 117 from South Africa. They also reported antibodies for Duvenhage and Denmark bat virus. Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that none of the 4 specimens they examined from the Democratic Republic of the Congo tested positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Burundi, Cameroon, Congo, Congo (Democratic Republic of the), Equatorial Guinea, Eswatini, Ethiopia, Gabon, Ghana, Kenya, Malawi, Mozambique, Namibia, Rwanda, Somalia, South Africa, South Sudan, Tanzania, Uganda, Zambia, Zimbabwe. Figure 27. Distribution of Epomophorus wahlbergi Genus Epomops Gray, 1866 *1870. Epomops Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 126 [Goto Description]. - Comments: Type species: Epomophorus franqueti Tomes, 1860. - Etymology: Probably an abbreviation of Epomophorus and the Greek "οψ", meaning aspect, referring to its resemblance to that genus (see Palmer, 1904: 268). (Current Combination) 2007. Epomop: Lameed, Biodiversity, 8 (4): 8. (Lapsus) 2014. Hepomops: Nina, Acta Med. Port., 27 (5): 627. Publication date: Sep-Oct 2014. 2016. Epomos: Pancer, Gut and Litwinska, Post. Mikrobiol., 55 (2): 207. 2019. Epomopos: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) TAXONOMY: Reviewed by Bergmans (1989). (2005: 323). See Simmons Andersen (1912b: 487), Grubb et al. (1998: 69) mention "Gray, 1866, Proc. Zool. Soc., Lond.: 65" as first publication in which the name Epomops was used. The results of the phylogenetic analyses by Almeida et al. (2016: 82) showed that the genus Epomops is not monophyletic as a DNA fragment of E. dobsonii shows a close relationship with Epomophorus wahlbergi, which seems to support the similarities or transitional traits of dobsonii with Epomophorus reported by Bergmans (1989: 118). Included in the Epomophorinii tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Hassanin et al. (2020: 5) include it in the Epomophorina subtribe of the Epomophorini tribe. Based on the analyses of mitochondrial and nuclear data, Hassanin et al. (2016: 525) suggest that the genus Epomops contains only one species, which can be divided into two subspecies: E. franqueti franqueti in Central Africa and E. f. buettikoferi in West Africa. Currently recognized species of the genus Epomops: buettikoferi (Matschie, 1899); franqueti (Tomes, 1860). COMMON NAMES: Czech: Grayovi kaloni. English: Clamorous Fruitbats, Noisy Fruit-bats, Singing Fruit-bats, Epauletted bats, Epaulet bats, African Epauletted bats. French: Epomophores de l'ouest. German: Epaulettenflughunde. MOLECULAR BIOLOGY: The Cytb analyses performed by Hassanin et al. (2016: 525) revealed that bats from West Africa, identified as either E. franqueti or E. buettikoferi, share very similar or identical haplotypes, which are divergent from those sequenced for all individuals of E. franqueti collected in Central Africa. 118 ISSN 1990-6471 PARASITES: HAEMOSPORIDA Perkins and Schaer (2016: Suppl.) mention the presence of Hepatocystis epomophori Rodhain, 1926 in Epomops sp. bats. Nine out of 10 (90 %) South Sudanese bats examined by Schaer et al. (2017: 2) were infected by Hepatosystis sp. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Côte d'Ivoire, Gabon, Ghana, Niger. Epomops buettikoferi (Matschie, 1899) *1899. Ep[omophorus (Epomophorus)] büttikoferi Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 43, 45. Type locality: Liberia: Junk river: Schieffelinsville [06 11 N 10 30 W] [Goto Description]. Holotype: RMNH ad ♂, mounted skin (skull missing). Collected by: F.X. Stampfli. - Comments: Matschie dated his introduction to 14 May 1899. - Etymology: In honour of J. Büttikofer, who collected animals in Liberia. 1972. Epomophorus (Epomops) franqueti buettikoferi: Püscher, Z. Säugetierk., 37 (3): 155. (Name Combination) 1996. Epomophorus buettiforteri: Kuhl, Obligate and opportunistic interactions, 52. (Lapsus) 2000. Epomops buetticoferi: Konstantinov, Pema, Labzin and Farafonova, Plecotus et al., 3: 145. (Lapsus) 2013. Epomops buettifoferi: Seltzer, Ndangalasi and Cordeiro, Biotropica, 45 (4): table S2. Publication date: 22 February 2013. (Lapsus) 2016. E[pomops] f[ranqueti] buettikoferi: Hassanin, Nesi, Marin, Kadjo, Pourrut, Leroy, Gembu, Musaba Akawa, Ngoagouni, Nakouné, Ruedi, Tshikung, Pongembo Shongo and Bonillo, C. R. Biologies, 339 (11-12): 525. Publication date: 14 October 2016. (Name Combination) 2019. Epomopos buettikoferi: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) ? Epomops buettikoferi: (Name Combination, Current Combination) TAXONOMY: See Bergmans (1989) and Simmons (2005: 323). VU (Baillie and Groombridge, 1996). 1994: VU (Groombridge, 1994). COMMON NAMES: Czech: kaloň Büttikoferův. English: Buettikofer's Clamorous Fruit-bat, Buettikofer's Epauletted Bat, Buettikofer's Epauletted Fruit Bat, Büttikofer's Fruit Bat. French: Epomophore de Büttikofer, Epomophore de Buettikofer, Roussette de Büttikofer, Roussette de Buettikofer. German: Büttikofers Epaulettenflughund. Regional None known. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in a number of protected areas, has a tolerance of a degree of habitat modification, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008as; IUCN, 2009; Monadjem, 2016a). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem, 2016a). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008as; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bg; IUCN, 2004). 1996: MAJOR THREATS: It is possible that there are no major threats to this species. Populations might be threatened by deforestation in parts of the species range, however, the extent to which it depends on forest cover is unclear (Mickleburgh et al., 2008as; IUCN, 2009; Monadjem, 2016a). CONSERVATION ACTIONS: Mickleburgh et al. (2008as) [in IUCN (2009)] suggest that further studies are needed into the dependence of this species on forested areas. Current conservation efforts This species has been recorded from a number of protected areas in West Africa (Mickleburgh et al., 1992), and in fact appears to thrive outside of these protected areas (Monadjem, 2016a). Conservation needs/priorities Monadjem (2016a) recommends studies are needed on the species’ population sizes, distribution, and extent of occurrence throughout African Chiroptera Report 2020 its range. Monitoring of population sizes and locations over time are also important to establish whether these are stable or experiencing trends of decline. The threats to these bats are poorly understood. Studies are needed on the species’ natural history and habitat requirements and the relationship between habitat and population sizes. Mickleburgh et al. (2008as) [in IUCN (2009)] suggested studies are needed into the dependence of this species on forested areas, and its tolerance of deforestation and habitat degradation (Monadjem, 2016a). GENERAL DISTRIBUTION: Epomops buettikoferi is a lowland West African species which ranges from Senegal and GuineaBissau in the west, to central Nigeria in the east. Bergmans (1975: 147) indicates it is restricted to the West African "moist forest" block. Native: Cameroon (Bakwo, 2009: 12); Côte d'Ivoire (Henry et al., 2004: 24); Ghana (Decher and Fahr, 2007: 10 - 11); Guinea (Denys et al., 2013: 281; Decher et al., 2016: 259); GuineaBissau (Rainho and Ranco, 2001: 26); Liberia (Monadjem and Fahr, 2007: 50); Nigeria; Senegal; Sierra Leone. MOLECULAR BIOLOGY: Denys et al. (2013: 281) determined the karyotype of four specimens from the Guinean Mount Nimba area and found 2n = 36 in females (34 autosomes and two X gonosomes), and 2n = 35 in males (X0 gonosomes); NFa = 68. X = submetacentric or medium-sized. ROOST: E. buettikoferi roosts alone or in small groups, near its feeding area (Kunz, 1996: 52). DIET: Seltzer et al. (2013: table S2) provide an overview of the plant species found beneath bat feeding roosts. For E. buettikoferi this included Milicia excelsa (Welw.) C. C. Berg (African Teak Moraceae). 119 Trend:- 2016: Decreasing (Monadjem, 2016a). 2008: Decreasing (Mickleburgh et al., 2008as; IUCN, 2009). REPRODUCTION AND ONTOGENY: Krutzsch (2000: 93) mentions that males attain puberty at the age of eleven months. Weber et al. (2019: 19) reported one pregnant female on 30 March and six lactating (24, 30, 31 March, 1, 7 April) in Sierre Leone. MATING: The male mating pattern in E. buettikoferi includes courtship vocalizations (calling territories) (see Krutzsch (2000: 111). PARASITES: HAEMOSPORIDA Schaer et al. (2013a: 17416) report the presence of hemosporidian parasites of the genus Hepatocystis in seven out of 25 investigated bats. VIRUSES: Rhabdoviridae Lyssavirus - Rabies related viruses Lagos bat virus - Hayman et al. (2008a) detected antibodies in Ghana. Drexler et al. (2012a: Suppl. Table S1) indicated that the single specimen they examined from Ghana did not test positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. UTILISATION: This species is hunted for bushmeat (Monadjem, 2016a). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Côte d'Ivoire, Ghana, Guinea, Liberia, Nigeria, Senegal, Sierra Leone. POPULATION: Structure and Density:- It appears to remain common in many areas, with the exception of Nigeria (Mickleburgh et al., 2008as; IUCN, 2009). West of Ghana, this species is typically one of the most abundant (in terms of captures) pteropodids in primary and secondary forest, and forest edge habitats (Monadjem, 2016a). Mickleburgh et al. (2008as) in IUCN (2009) suggest that populations may be declining in areas of ongoing deforestation. Figure 28. Distribution of Epomops buettikoferi 120 ISSN 1990-6471 Epomops franqueti (Tomes, 1860) *1860. Epomophorus franqueti Tomes, Proc. zool. Soc. Lond., 1860, I: 54, pl. 75. Publication date: February - May 1860. Type locality: Gabon: "Gabon" [Goto Description]. Holotype: MNHN A.107-776-C.52-1852-257: ad ♂, mounted skin and skull. Collected by: Franquet. Skull has number A6767; see Rode (1941: 233), who uses number 164. 1862. Epomophorus comptus H. Allen, Proc. Acad. nat. Sci. Philad., 13 (11): 158 (for 1861). Type locality: "Africa": West Africa [Goto Description]. - Comments: Andersen (1912b: 497) and Allen (1939a: 57) mentioned "=Gaboon" as type locality. Publication date: Volume 13 issue 11 covers the meetings of June-July 1861, but was probably only printed in 1862. Andersen (1912b: 498) mentions "as having been collected by Du Chaillu the specimen is no doubt from Gaboon." Type in Phildalphia Museum, unmounted skin, skull extracted, ♀. 1910. Epomops franqueti strepitans K. Andersen, Ann. Mag. nat. Hist., ser. 8, 5 (25): 106. Publication date: 1 January 1910. Type locality: Nigeria: 150 mi up the Niger: Asaba [06 11 N 06 43 E]. Holotype: BMNH 1895.5.3.7: ad ♂, skin and skull. Presented/Donated by: Dr. W.H. Cross. See Andersen (1912b: 497). 2017. Epomops frangueti: Dammann, Semin. Cell Dev. Biol., 70: 160. Publication date: 8 July 2017. (Lapsus) 2019. Epomopos franqueti: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) 2019. Epomops frangeti: Kaswera Kyamakya, Nebesse Mololo and Gambalemoke Mbalitini, IJEAB, 4 (2): 330.. Publication date: March-April 2019. (Lapsus) ? Epomophorus (Epomops) franqueti: (Name Combination) ? Epomops franqueti franqueti / strepitans: (Name Combination) ? Epomops franqueti franqueti: (Name Combination) ? Epomops franqueti: (Name Combination, Current Combination) TAXONOMY: See Bergmans (1989) and Simmons (2005: 324). COMMON NAMES: Czech: kaloň Franquetův, kaloň plavý. English: Franquet's Clamorous Fruit-bat, Franquet's Fruit Bat, Franquet's Epauletted Fruit Bat, Singing Fruit Bat. French: Epomophore de Franquet, Chien volant à épaulettes du Congo, Roussette de Franquet. German: Franquets Epaulettenflughund, Franquet's Epaulettenflugfuchs. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008by; IUCN, 2009; Kityo and Nalikka, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Kityo and Nalikka, 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008by; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ci; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Malhi et al. (2013) estimated deforestation rates for Africa to be at 0.29 million ha yr-1 between 2000 – 2010 which figure is much lower than any where else in the tropics of the world. Notwithstanding the foregoing, since the core habitat of the species is forest, reduction in its extent poses an increasing risk of habitat loss (Kityo and Nalikka, 2016). There is currently an increasing perception that bats are playing a major role in the spread of zoonoses, rightly or wrongly, this puts bats into direct line for persecution and extermination (Kityo and Nalikka, 2016). CONSERVATION ACTIONS: Kityo and Nalikka (2016) supports Mickleburgh et al. (2008by) [in IUCN (2009)] who report that E. franqueti is present in many protected areas. No direct conservation measures are currently needed for this adaptable species as a whole. African Chiroptera Report 2020 GENERAL DISTRIBUTION: Epomops franqueti is a widespread species, which is present in much of West Africa and Central Africa, with some records from East Africa. It ranges from Côte d'Ivoire and Ghana in the west, to Cameroon, and then through Central Africa to southern Sudan, Rwanda and Uganda in the east. It has been recorded as far south as northern Angola, southern Democratic Republic of the Congo and northeastern Zambia. The record (MNHN ZM-MO-1984-1165) from Madagascar is a possible misidentification/or incorrect locality information and needs verification. Native: Angola (Bergmans, 1989; Monadjem et al., 2010d: 554); Benin (Capo-Chichi et al., 2004: 161); Cameroon; Central African Republic (Morvan et al., 1999: 1195); Congo (King and Dallimer, 2010: 65; Bates et al., 2013: 333); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 554); Côte d'Ivoire; Equatorial Guinea; Gabon; Ghana; Nigeria; Rwanda; Sudan; Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 59); Zambia (Ansell, 1978; Monadjem et al., 2010d: 554). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data for 7 specimens: Fa: 93.51 ± 5.246 mm, mass: 114.7 ± 35.86 g, wing loading: 19.7 ± 3.81 N/m2, aspect ratio: 5.4 ± 0.43. DETAILED MORPHOLOGY: Dobson (1881: 686) provides a detailed description of the pharynx, larynx, and hyoid bones. Chawana et al. (2013: 160) report an average brain mass of 2.42 g (n = 2). FUNCTIONAL MORPHOLOGY: Coimbra et al. (2016: 191) found that the eye of E. franqueti has a bright yellow tapetum lucidum, which covers almost the entire extension of its dorsal hemisphere. There are about 200,000 retinal ganglion cells. The minimum angle of resolution was approximately 0.167° (ca. 5-6 mm at 1 m distance). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - In Kenya, Peterson and Nagorsen (1975) recorded 2n = 36. Haiduk et al. (1980) reported 2n = 36, FN = 68, BA = 34, a subtelocentric X chromosome and a subtelocentric Y chromosome for specimens from Cameroon. 121 Haiduk et al. (1981: 224) mention 2n = 36, FN = 66. Primus et al. (2006) reported a male without Y chromosome (2n = 35) from Gabon. Protein / allozyme - Unknown. Dammann (2017: 160) refers to Ifuta et al. (1988: 379), who found that levels of T 4 (plasma concentrations of thyroxine) were about 10 times lower in this bat than in other mammals. These low values might be stress related as several hours elapsed between the capture of the bats and taking blood samples. However, these low values were not found for T3 (Plasma concentrations of triiodothyronine). Dammann (2017) suggests that the low levels of T4 might be related to the longevity in bats. Ekeolu and Adebiyi (2018: 391) investigated blood parameters in 17 E. franqueti specimens from Nigeria and found no significant sex-related differences, except for erythrocyte osmotic fragility, which was significantly higher in female than in male bats at 0.1 % NaCl. HABITAT: This species is associated with lowland tropical moist forest, where it has been recorded in both primary and secondary habitat. Populations are also present in the mosaic habitats of tropical moist forest with woodland and grassland (Bergmans, 1989). It is an adaptable species that can be found in disturbed areas, seemingly being only absent from heavily degraded or urban areas. HABITS: Bradbury (1977) [in McCracken and Wilkinson (2000: 351)] found that females and juvenile animals were feeding near the territory of a calling male, which would suggest that additional resources could be available in that area. ROOST: The species roosts alone or in small groups, often close to water, in the dense leave cover of shrubs or trees (Brosset, 1966c; Jones, 1972; Kunz, 1996: 44). DIET: In Congo, Kipalu (2009: 16) reported E. franqueti visiting/consuming fruits of Chlorophora excelsa (African teak) and Ficus mucuso. Gembu Tungaluna (2012: 95 - 97) reported the following fruits being eaten in the Kisangani area (DRC) in the various seasons: First heavy rain season (September - November): Persea americana Mill., 1786 (avocado) (24%), Carica papaya L. (papaya) (16%), Spondias cytherea Sonnerat, 1782 (Ambarella, otaheite apple, great 122 ISSN 1990-6471 hog plum) (14%), Annonidium mannii (Oliv.) Engl. & Diels (Junglesop) (11%), Musanga cecropioides R.Br. & Tedlie (African corkwood tree, umbrella tree) (8%), Ficus leprieuri Miq., 1867 (8%), Myrianthus arboreus P.Beauvois, 1805 (giant yellow mulberry, bush pineapple, corkwoord) (5%), Elaeis guineensis Jacq. (African oil palm, macawfat) (3%), Ficus wildemanniana Warb. (3%), Musa sp. (3%). First mild rain season (December - February): Musanga cecropioides (40%), Elaeis guineensis (20%), Myrianthus arboreus (13%), Carica papaya (13%), Ficus wildemanniana (7%), Ficus vallischoudae Delile (7%). Second heavy rain season (March - May): Carica papaya (23%), Dacryoides edulis H.J. Lam (safou, African pear, bush pear, Nsafu, bush butter tree, butterfruit) (15%), Musanga cecropioides (15%), Myrianthus arboreus (8%), Elaeis guineensis (8%), Oncoba welwitschii Oliv. (8%), Ficus sp3 (8%), Musa sp. (8%), Ficus leprieuri (7%). "Dry" season (June August): Ficus wildemanniana (100%). PREDATORS: Charles-Dominique (1974: 144) reported on a potto Perodicticus potto (Statius Müller, 1766) eating a juvenile E. franqueti. POPULATION: Structure and Density:- This is a common species (Mickleburgh et al., 2008by; IUCN, 2009). Kityo and Nalikka (2016) reports that it is a fairly abundant species in the right habitat. Trend:- 2016: Stable (Kityo and Nalikka, 2016). 2008: Stable (Mickleburgh et al., 2008by; IUCN, 2009). ACTIVITY AND BEHAVIOUR: For a selection of sounds, check http://macaulaylibrary.org/search?taxon=Epomop s franqueti&taxon_id=11069435&taxon_rank_id=67 &tab=audio REPRODUCTION AND ONTOGENY: From southern Sudan, Bates (1905: 72) reports on two females (probably of this species) carrying a half-grown young on 31 August and 1 September. A male with probably active testes was caught on 2-3 November in Pointe Noire (Congo) by Bergmans (1979a: 171), and a lactating female with an embryo was captured in Brazzaville on 23 December. Furthermore, females with developed nipples were collected in March, May, November and December. MATING: Brosset (1966c) and Bradbury (1977) [in McCracken and Wilkinson (2000: 349)] indicate that males establish nocturnal display sites where females visit males for mating. PARASITES: BACTERIA Gram-negative bacterium - Nowak et al. (2017: 6) tested three bats from the Republic of Congo of which two tested positive for Escherichia. coli. Mbehang Nguema et al. (2020: 7) reported the presence of E. coli, Enterobacter cloacae and Klebsiella pneumoniae in feacal samples collected near Makokou in Gabon. HAEMOSPORIDA Lutz et al. (2016: 9) examined 52 E. franqueti specimens from East Africa and found 35 of them infected with Hepatocystis sp. Perkins and Schaer (2016: Suppl.) refer to the presence of Hepatocystis brosetti Miltgen et al., 1977 in Gabon, from where it was described by Miltgen et al. (1977: 589). Landau et al. (2012: 142) also mention it as type host for Hepatocystis epomophori (Rodhain, 1926). Boundenga et al. (2018: 10581) reported 21 out of 160 (13.12 %) bats from the Franceville (Gabon) area to be infected by Hepatocystis sp. Surprisingly, Rosskopf et al. (2018: 31) were unable to find any Hepatocystis parasites in the 21 Gabonese bats they examined and they suggest that this might be linked with the fact their bats were collected during the dry season. DIPTERA Nycteribiidae: Basila tenuispina Theodor 1957 from Stanleyville, Congo (Democratic Republic of the) (Haeselbarth et al., 1966: 111). ACARI Teinocoptidae: Teinocoptes epomophori Rodhain, 1923 was reported from an E. franqueti specimen from Stanleyville (=Kisangani), DRC by Fain (1967: 367). VIRUSES: Willoughby et al. (2017: Suppl.) report the following viruses: Marburg marburgvirus, Rift Valley fever phlebovirus, Shamonda orthobunyavirus, Zaire ebolavirus. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 to 1999 five individuals from Bandundu Province, DRC for antibody to SARS-CoV in sera, none were found to be positive (0/5). Anthony et al. (2017b: Suppl.) mention the betacoronavirus Eidolon_bat_CoV. African Chiroptera Report 2020 123 Joffrin et al. (2020: 7) identified the Congolese viruses as Beta-D coronavirus. Filoviridae - Filo viruses Ebolavirus Pourrut et al. (2007) found homogeneous ZEBOV [Zaire Ebolavirus] infection in the wild populations of E. franqueti in Gabon and the Democratic Republic of the Congo. Eight out of 117 bats had immunoglobulin G (IgG) specific for Ebola in Gabon during a test performed by Leroy et al. (2005: 575). 10 out of 27 bats were found to be positive for EBOV antibodies in the Greater Accra region, Ghana (Hayman et al., 2012d: 1208). Bausch and Schwarz (2014: 1) and Pigott et al. (2014: 9) indicate that E. franqueti is one of the leading candidates for introducing Ebola in Guinea (together with Hypsignathus monstrosus and Myonycteris torquata). However, no proof has been provided that this was indeed the case. Marburgvirus Towner et al. (2007) tested 296 individuals from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and 47 individuals for antiMarburg virus IgG antibodies by ELISA; no positives were found. Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that none of the 100 specimens they examined from Gabon, Congo, Ghana and the Central African Republic tested positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. In their overview table, Maganga et al. (2014a: 8) report that the following viruses were already found on E. franqueti: Zaire Ebola virus (ZEBOV), Reston Ebola virus, Marburg virus (MBGV), Flavivirus. UTILISATION: This species is hunted for the bushmeat trade (Kityo and Nalikka, 2016). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea-Bissau, Kenya, Liberia, Nigeria, Rwanda, Sierra Leone, South Sudan, Tanzania, Togo, Uganda, Zambia. Flaviviridae Flavivirus Maganga et al. (2014a: 5) tested 128 bats from Congo, of which 2 tested positive. Nairoviridae Nairovirus Of the 99 specimens from Gabon and Congo tested by Müller et al. (2016: 3), 7 were positive for Crimean Congo hemorrhagic fever virus (CCHFV). Phenuiviridae Phlebovirus Fagre and Kading (2019: 4) report that Rift Valley fever virus (RVFV) was either isolated from this bat or that molecular evidence was found (see also de Jong et al. (2011: 11) and Luis et al. (2013: suppl.)). Figure 29. Distribution of Epomops franqueti Genus Hypsignathus H. Allen, 1862 *1862. Hypsignathus H. Allen, Proc. Acad. nat. Sci. Philad., 13 (11): 156 (for 1861). Publication date: 1862 [Goto Description]. - Comments: Type species: Hypsignathus monstrosus H. Allen, 1861. Publication date: Volume 13 issue 11 covers the meetings of June-July 1861, but was probably only printed in 1862. - Etymology: From the Greek "υψι", meaning on high or aloft and "γνάθος", meaning jaw, possibly referring to the 'deeply arched mouth' (see Palmer, 1904: 343). (Current Combination) 1862. Sphyrocephalus A. Murray, Proc. zool. Soc. Lond., 1862, I: 8. Publication date: June 1862 [Goto Description]. - Comments: Type species: Sphyrocephalus labrosus Murray,1862 (=Hypsignathus monstrosus H. Allen, 1861). Preoccupied by Sphyrocephala Westwood, 1848, a dipteran and Sphyrocephalus Schmarda, 1859, a 124 ISSN 1990-6471 1862. ? ? turbellarian (see Palmer, 1904: 642; Langevin and Barclay, 1990: 1). Andersen (1912b: 501 - footnote) also mentions "Murray probably originally intended to name this genus Zygænocephalus in allusion to the Hammer-headed Sharks Zygæna, and had that name printed on the plate, but for some reason or other changed it in the text to Sphyrocephalus.". - Etymology: From the Greek "σφύρα", meaning hammer and "κεφαλή", meaning head (see Palmer, 1904: 642; Murray, 1862: 8). Zygænocephalus A. Murray, Proc. zool. Soc. Lond., 1862, I: pl. 1. Publication date: June 1862. - Comments: Type species: Zygænocephalus labrosus Murray,1862 (=Hypsignathus monstrosus H. Allen, 1861). This name was attached to the plate illustrating Murray's description of Sphyrocephalus and serves as a replacement for Sphyrocephalus Murray, 1862: see Langevin and Barclay (1990: 1). - Etymology: From the Greek "ξύγαινα", a hammer-headed shark and "κεφαλή", meaning head (see Palmer, 1904: 717). Epomophorus (Hypsignathus): (Name Combination) Hypsygnathus: (Lapsus) TAXONOMY: Revised by Bergmans (1989). (2005: 324). See Simmons Included in the Epomophorinii tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Included in the Epomophorini tribe and Epomophorina subtribe by Hassanin et al. (2020: 5). Currently (Simmons and Cirranello, 2020) recognized species of Hypsignathus: monstrosus H. Allen, 1861. COMMON NAMES: Czech: kladivohlaví kaloni. English: Hammerheaded bats. French: Hypsignates. German: Hammerkopf-Flughunde. Hypsignathus monstrosus H. Allen, 1862 *1862. H[ypsignathus] monstrosus H. Allen, Proc. Acad. nat. Sci. Philad., 13 (11): 157 (for 1861). Publication date: 1862. Type locality: "Africa": Western Africa: possibly Gabon (see Andersen, 1912b: 506). [Goto Description]. - Comments: Type specimen collected by Mr. Du Chaillu (Murray, 1862: 8); Allen (1861: 158) mentions the name as Duchaillu. Andersen (1912b: 508) mentions the type specimen as being an adult ♂, SS, in the collection of the Philadelphia museum. Publication date: Volume 13 issue 11 covers the meetings of June-July 1861, but was probably only printed in 1862, as Murray (1862: 8) indicates that he received a copy of this publication on 19 February 1862. But we also found a reference to issue 23 of volume 13 (for the meeting of 26 November 1861) having been published on 31 December 1861, which might indicate that issue 11 might have been published in 1861. (Current Combination) 1862. Sphyrocephalus labrosus A. Murray, Proc. zool. Soc. Lond., 1862, I: 8. Publication date: June 1862. Type locality: Nigeria: Eastern region: Old Calabar river [ca. 04 56 N 08 22 E] [Goto Description]. Holotype: BMNH 1862.1.23.1[b]: ad ♂, skull and alcoholic. Collected by: Reverend William C. Thomson. Presented/Donated by: Mr. A. Murray Esq. - Comments: The publication contains 14 January 1862 as date, but see Duncan (1937) for the actual publication date. 1862. Zygaenocephalus labrosus: A. Murray, Proc. zool. Soc. Lond., 1862, I: pl. 1. Publication date: June 1862. (Name Combination) 1876. Epomophorus macrocephalus Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 474. Type locality: Gabon: Dongila. - Comments: Not of Ogilby (see Andersen, 1912b: 507). 1879. Epomophorus monstrosus: Dobson, Proc. zool. Soc. Lond., 1878, IV: 879. Publication date: April 1879. (Name Combination) 1896. Epomophorus (Hypsignathus) monstrosus: Pousargues, Ann. Sci. Nat., (8) 3: 250. (Name Combination) 1899. Hypsignathus haldemani Matschie, Sber. Ges. naturf. Freunde Berlin, 29. - Comments: Not Pteropus haldemani Halowell, 1846: See Langevin and Barclay (1990: 1). 1968. Epomophorus monstruosus: Andral, Brès, Sérié, Casals and Panthier, Bull. Wld. Hlth. Org., 38 (6): 860. (Lapsus) African Chiroptera Report 2020 1975. 1978. 2019. ? ? ? 125 Hypsignathus montrosus: Koopman, Bull. Am. Mus. Nat. Hist., 154 (4): 362. (Lapsus) Hypsignathus monstrousus: Germain, Elsevier / North-Holland Biomedical Press Amsterdam, 133. (Lapsus) Hypsignathus monstruosus: Kaswera Kyamakya, Nebesse Mololo and Gambalemoke Mbalitini, IJEAB, 4 (2): 330. Publication date: March-April 2019. (Lapsus) Epomophorus (Hypsignathus) monstrosus: (Name Combination) Hypsignathus monstrosus: (Current Combination) Hypsygnathus monstrosus: (Lapsus) TAXONOMY: See Langevin and Barclay (1990), Bergmans (1989) and Simmons (2005: 324). COMMON NAMES: Castilian (Spain): Cabeza Martillo. Chinese: 锤 头果蝠. Czech: kaloň kladivohlavý, kaloň štítohlavý. English: Hammer-headed bat, Hammer-headed Fruit Bat. French: Hypsignathe monstrueux, Chien volant à tête en marteau, chauve-souris à tête de marteau. German: Hammerkopf, Hammerkopf-Flughund. Kinande (DRC): Mulina. Tanoboase (Ghana): ahwenekron. Vai (Liberia): Tuña. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008az; IUCN, 2009; Tanshi, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Tanshi, 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008az; IUCN, 2009). 2004: LC ver 3.1 (2001) (IUCN, 2004; Mickleburgh et al., 2004bw). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: This species is locally threatened by habitat destruction, particularly the loss of riverine forests. In parts of its range it is hunted for subsistence food (Mickleburgh et al., 2008az; IUCN, 2009; Tanshi, 2016). It is persecuted for its noise during leks and is hunted for food across its range (Tanshi, 2016). The extent to which this constitutes a threat is unknown and requires investigation. Duke (1934: 74) provided some suggestions on how to shoot this bat as this seemed to be the only way to get rid of its terrible noice. CONSERVATION ACTIONS: Mickleburgh et al. (2008az) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. It has been recorded from Tai National Park (Côte d'Ivoire). Okomu National Park (Nigeria) and Afi Mountain Wildlife Sanctuary (Nigeria), Atewa Forest Reserve (Ghana) and several others across its range (Tanshi, 2016). Mickleburgh et al. (2008az) [in IUCN (2009)] recommend that further studies are needed into the distribution of this species. GENERAL DISTRIBUTION: Hypsignathus monstrosus is widespread in West and Central Africa. It ranges from Guinea-Bissau in the west, to northwestern Angola in the south, and as far east as western Ethiopia and western Kenya (near Lake Victoria). It has been recorded to at least 1,800 m asl. Native: Angola (Crawford-Cabral, 1989; Bergmans, 1989; Monadjem et al., 2010d: 554); Benin; Burkina Faso (Kangoyé et al., 2015a: 605); Cameroon; Central African Republic (Hayman et al., 1966; Morvan et al., 1999: 1195; Monadjem et al., 2010d: 554); Congo (King and Dallimer, 2010: 65; Bates et al., 2013: 333); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 554); Côte d'Ivoire (Heim de Balsac, 1934b: 24); Equatorial Guinea; Ethiopia (Lavrenchenko et al., 2004b: 131); Gabon; Ghana (Decher and Fahr, 2007: 11); Guinea (Denys et al., 2013: 281); Guinea-Bissau (Rainho and Ranco, 2001: 21); Kenya; Liberia (Monadjem and Fahr, 2007: 50); Nigeria (Happold, 1987; Capo-Chichi et al., 2004: 161); Sierra Leone (Grubb et al., 1998); Sudan; Togo; Uganda (Kityo and Kerbis, 1996: 59). Presence uncertain: Gambia. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: The peculiar head of the males of this species, made Johnston (1906: 689) describe it as: "In these bats [Epomophori] the head is unusually large and long; but it is so monstrous and hideous in the biggest of the sub-family - the hammerheaded bat, Hypsignathus - as to provoke the appropriate specific term of monstrosus. This animal is the creation of a nightmare imagination. 126 ISSN 1990-6471 It might be recommended to the designers of an instructive pantomime or Wagnerian opera." Duke (1934: 72) compares its head with the gargoyles of the Nôtre Dame, and described it as half hippo, half horse. From western Uganda, Monadjem et al. (2011: 30) reported the following data: Fa: 115.7 mm, mass: 211 g, wing loading: 21.2 N/m 2, aspect ratio: 5.8. GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: Eisenberg and Wilson (1978: 744) indicate that the cranial volume of H. monstrosus is slightly smaller than was expected from a regression line they calculated for all frugivorous bats. The lower score could be linked with the partially carnivorous diet of these bats as found by Van Deusen (1968). Davies et al. (2013a: 4) provide an illustration of the bony labyrinth of H. monstrosus's inner ear. DETAILED MORPHOLOGY: The average brain mass for two H. monstrosus specimens was found to be 3.81 g (Chawana et al., 2013: 160). FUNCTIONAL MORPHOLOGY: The study of Coimbra et al. (2016: 191) found that the eye of H. monstrosus possesses a bright yellow tapetum lucidum, which covers almost the entire extension of its dorsal hemisphere. Its retinal area is about 200 mm 2 and contains about 3000,000 retinal ganglion cells. The minimum angle of resolution was approximately 0.122° (ca. 40 mm at 10 m distance). Kawashima et al. (2017: 48) examined the postcranial skeleton of six H. monstrosus specimens and found that five had 15 thoracic vertebrae and one had 14; all had seven cervical, three lumbal and four sacral vertebrae. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Peterson and Nagorsen (1975) and Haiduk et al. (1980; 1981: 225) reported 2n = 36, FN = 68, BA = 34, a subtelocentric X chromosome and a subtelocentric Y chromosome for specimens from Kenya and Cameroon, respectively. 2n = 36 was also reported by Primus et al. (2006). Protein / allozyme - Unknown. ROOST: During the day, H. monstrosus roosts in trees, in most cases as singles of either sex, but also in mixed sex groups of up to 17 individuals. Males typically roost at the periphery of larger groups (see Bradbury, 1977). DIET: Duke (1934: 72) indicates that H. monstrosus starts to feed soon after dark, and that it eats fruits of various kinds, especially guavas and certain kinds of wild figs. Gautier-Hion and Michaloud (1989: 1829) reported that 88 % of 133 oral swabs and fecal samples analysed from May to October in Makokou (Gabon) contained figs. They reported (p. 1831) that females forage at about 1 to 4 km from their roosts, whereas males travel up to 10 km, and also that figs only form 38 % of the females diet against 58 % in males. Van Deusen (1968) [in Courts (1998: 190)] observed this bat attacking live chickens and scavenging the carcasses of small birds. Seltzer et al. (2013: table S2) provided an overview of the plant species found beneath bat feeding roosts. For H. monstrosus this included Milicia excelsa (Welw.) C. C. Berg (African Teak Moraceae). From the Kisangani area (DRC), Gembu Tungaluna (2012: 99 - 100) reported the following fruits being eaten during the various seasons: First heavy rain season (September - November): Persea americana Mill., 1786 (avocado) (21%), Annonidium mannii (Oliv.) Engl. & Diels (Junglesop) (18%), Carica papaya L. (papaya) (14%), Dacryoides edulis H.J. Lam (safou, African pear, bush pear, Nsafu, bush butter tree, butterfruit) (14%), Spondias cytherea Sonnerat, 1782 (Ambarella, otaheite apple, great hog plum) (11%), Uapaca guineensis Müll.Arg., 1864 (Sugar plum, red cedar, false mahogany, rikio) (7%), Myrianthus arboreus P.Beauvois, 1805 (giant yellow mulberry, bush pineapple, corkwoord) (4%), Elaeis guineensis Jacq. (African oil palm, macawfat) (4%), Ficus wildemanniana Warb. (4%), Ficus leprieuri Miq., 1867 (3%). First mild dry season (December - February): Myrianthus arboreus (50%), Musanga cecropioides R.Br. & Tedlie (African corkwood tree, umbrella tree) (50%). Second heavy rain season (March - May): Musanga cecropioides (75%), Ficus leprieuri (25%). No data for the "Dry" season (June - August). PREDATORS: Neil (2017: 131) reported on a hammer-headed fruit bat being captured by a Long-tailed hawk, Urotriorchis macrourus (Hartlaub, 1855) in Gabon. The hawk left its wounded prey on the ground, after which the bat was able to escape. POPULATION: Structure and Density:- It is a locally common species. Roosts can contain as many as 25 bats, African Chiroptera Report 2020 but the average roost size is fewer than five animals (Mickleburgh et al., 2008az; IUCN, 2009; Tanshi, 2016). Trend:- 2016: Unknown (Tanshi, 2016). 2008: Unknown (Mickleburgh et al., 2008az; IUCN, 2009). ACTIVITY AND BEHAVIOUR: Bradbury (1977) observed females hover in front of selected males before copulation. Carpenter (1986: 90) and McGuire and Guglielmo (2009: 1291) reported that the respiratory quotient for this species indicates that fat provided 77 - 92 % of the fuel during flight, and a decline in respiratory quotient during long flights may prove that fat is indeed the only fuel source following initial carbohydrate use. Carpenter (1986: 93) also indicates that at airspeeds of 5.5 m/s the heart beats at a frequency of 620 beats per minute, whereas the respiration rate is 293 breadths per minute. The courtship calls, which likely have a territorial function, consist of a simple syllable type in series (Bradbury, 1977 in Smotherman et al., 2016: 538). For a selection of sounds, check http://macaulaylibrary.org/search?taxon=Hypsign athus monstrosus&taxon_id=11075759&taxon_rank_id =67&tab=audio REPRODUCTION AND ONTOGENY: Tuttle and Stevenson (1982: 120, 121) indicate that females attain sexual maturity at 5 to 6 months of age, whereas males need 18 months. Kurta and Kunz (1987: 82) report that, at birth, the young weights about 16.2 % of the mother's body mass (38.0 g versus 234.0 g). MATING: Bradbury (1977) reported that H. monstrosus exhibit lek behaviour at their mating sites: Males form large choruses during the night that are visited by females, and they respond to the presence of females by increasing the repetition rate of their calls and by moving their wings. He also found that females favor larger males with bulbous muzzles and enormous hypertrophied larynxes, which are acting as resonating chambers. Male sites are spaced about 10 m apart, and individual males use the same site every night. Olson et al. (2019: 2) indicate that the lek behaviour might be restricted to the Central African populations, as in West Africa the reproductive strategy seems to be harem-based. 127 In Abidjan (Côte d'Ivoire), Niamien et al. (2015b: 230) found that H. monstrosus males prefered relatively low (5 - 10 m) Terminalia catappa L. trees (Indian almond - Combretaceae) for their leks. The individual trees were about 5 m apart. The males were present from August to October (small dry seaon to start of small wet season), and from February to May (end of large dry season to start of large wet season). In the other months no males were observed. Bradbury (1977) [in Fasel et al. (2016: 2)] reported on uneven copulation rates, whereby harem males mate at a higher frequency than other males. PARASITES: BACTERIA Gram-negative bacterium - Nowak et al. (2017: 6) tested two bats from the Republic of Congo, both of which were positive for E. coli. HAEMOSPORIDA Ayala et al. (1981: 21) examined blood smears from 142 H. monstrosus specimens from Gabon, and found 139 of them to be infected by hemosporidian parasites of the genus Hepatocystis. They also indicate that the bats probably became infected before the age of six months and that the infection rate varied according to sex, age and female reproductive status. Schaer et al. (2013a: 17416) report the same parasites in five out of 10 investigated bats. Perkins and Schaer (2016: Suppl.) mention Hepatocystis carpenteri, described by Miltgen et al. (1980), from Gabon and H. epomophori and H. sp. from various countries. Hepatocystis sp. Infections were also reported by Boundenga et al. (2018: 10581) from bats from Gabon (1 out of 21 examined: 4.8%). Hepatocystis gametocytes were reported from this bat for the first time in the DRC by Rodhain (1913), although he referred the bats erroneously to Epomops franqueti franqueti (see Miltgen et al., 1977: 595). Hepatocystis parasites were also found in the single Hypsignathus sp. from South Sudan examined by Schaer et al. (2017: 2). ACARI Fain (1967: 364) reported H. monstrosus from Epulu, DRC as new host for Teincoptes auricularis Fain, 1959 (Acari: Teinocoptidae). The parasites were found around the nipples of the host. VIRUSES: Willoughby et al. (2017: Suppl.) report the following viruses: Lagos bat lyssavirus, Marburg 128 ISSN 1990-6471 marburgvirus, Mokola lyssavirus, Nipah henipavirus, Reston ebolavirus, Zaire ebolavirus. henipavirus (M74) induce syncytium formation in a kidney cell line derived from H. monstrosus. Nairoviridae Orthonairovirus Only one of the 101 specimens from Gabon and Congo tested by Müller et al. (2016: 3) was positive for Crimean Congo hemorrhagic fever virus (CCHFV). Hayman et al. (2008b) detected antibodies Nipah virus in one bat collected in Ghana in 2007. Drexler et al. (2012a: Suppl. Table S1) reported that 2 out of 4 specimens they examined from Gabon, Congo, Ghana and the Central African republic tested positive for Henipavirus and Rubulavirus. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, 11 individuals from Bandundu Province, DRC for antibody to SARS-CoV in sera, one was found to be positive (9.1 %). Filoviridae - Filo viruses Ebolavirus Pourrut et al. (2007) found homogeneous ZEBOV [Zaire Ebolavirus] infection in the wild populations of H. monstrosus in Gabon and the Democratic Republic of the Congo. Hayman et al. (2010) tested the blood of 3 adult females from Ghana for Ebola virus using indirect flurescent tests for antibodies against EBOV subtype Zaire, none tested positive. However, Hayman et al. (2012d: 1208) found 7 out of 16 bats to be positive for EBOV antibodies in Ghana's Greater Accra region. Four out of 17 bats had immunoglobulin G (IgG) specific for Ebola in Gabon during a test performed by Leroy et al. (2005: 575). Bausch and Schwarz (2014: 1) and Pigott et al. (2014: 8) indicate that H. monstrosus is one of the leading candidates for introducing Ebola in Guinea (together with Epomops franqueti and Myonycteris torquata). However, no proof has been provided that this was indeed the case. Marburgvirus Towner et al. (2007) tested 56 individuals from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and 12 individuals for antiMarburg virus IgG antibodies by ELISA; no positives were found. Hayman et al. (2010) tested the blood of 3 adult females from Ghana for Marburg virus using indirect flurescent tests for antibodies against MARV subtype Leiden, none tested positive. Peterson et al. (2007: 1547) indicate that the geographic distribution of H. monstrosus covers the entire range of the Ebola virus, but not that of the Marburg virus. Paramyxoviridae Henipavirus Krüger et al. (2013: 13889) demonstrated that the surface glycoproteins F and G of an African In their overview table, Maganga et al. (2014a: 8) reported that the following viruses were already found on H. monstrosus: Zaire Ebola virus (ZEBOV), Reston Ebola virus, Marburg virus (MBGV), Coronavirus (SARS-CoV), Nipah virus (NPHV). Bennett et al. (2018: 65) studied viruses in the blood of H. monstrosus from the Republic of Congo and found that four out of five viruses are phylogenetically and genomically allied with viruses of arthropods: a nodavirus ("hypsignathus nodavirus", provisionally genus Hypnovirus, family Nodaviridae), a dicistrovirus ("hypsignathus dicistrovirus", genus Cripavirus, family Dicistroviridae), and two distinct variants of the recently recognized tombus-like viruses ("Hypsignathus monstrosus tombus-like virus 1 and 2", family Tombusviridae). The fifth virus was a novel hepatitis B-like virus ("Hypsignathus monstrosus hepatitis B virus (HMHBV)", family Hepadnaviridae), which is of mammalian origin. They also suggest that the viruses from arthropod origin might have been accidentally ingested, as the insects they are associated with, are themselves parasites of fruits eaten by the bats. (In a Corrigendum to their paper, the authors indicate that the characterization of their putative HBV sequences should be considered inconclusive as contamination of reagents might have occurred.) UTILISATION: Bobo and Ntumwel (2010: 232) report that in the Korup area of Cameroon the whole animal is used for medicinal purposes, as it would render women fertile when consumed. The species is hunted for food, and being Africa’s largest bat, it is targeted, although because it forms small groups or individuals, it is not the most hunted fruit bat species (Mildenstein et al., 2016). ANTHROPOPHILOUS: Jeffreys (1944: 73) [referring to Goddard (1925)] mentions "From Sierra Leone comes an account of the gruesome habits of the large fruit bat. 'One of the most uncanny superstitions is that of the African Chiroptera Report 2020 129 'Boman' in which anthropologists will recognise the vampire of European superstition. This creature is said to suck the blood of sleeping children until they die: it can turn into a stone or snake at will. The 'Boman' is in reality the hammer-headed bat (Hypsignathus monstrosus), the largest fruit bat found in Africa; its dull and monotonous cry at the time when fruit is ripening has struck terror into many a village, whose inhabitants will sally forth from their houses and beat tins to drive it away, cursing its father and mother and all its ancestors the while.'’ DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Burkina Faso, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Nigeria, Sierra Leone, South Sudan, The Gambia, Togo, Uganda. Figure 30. Distribution of Hypsignathus monstrosus Genus Nanonycteris Matschie, 1899 *1899. [Epomophorus (]Nanonycteris[)] Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 35, 58 [Goto Description]. - Comments: Described as a subgenus of Epomophorus Bennett, 1836. Type species: Epomophorus veldkampii Jentink, 1888. Matschie dated his introduction on 14 May 1899. - Etymology: From the Greek "νάνος", meaning dwarf and "νυκτερισ", meaning bat (see Palmer, 1904: 448). 1943. Nannonycteris: Pohle, Sber. Ges. naturf. Freunde Berlin, 78 (for 1942). (Lapsus) 2007. Nnonycteris: Lameed, Biodiversity, 8 (4): 8. (Lapsus) 2019. Nanonycteri: Mbu'u, Mbacham, Gontao, Kamdem, Nlôga, Groschup, Wade, Fischer and Balkema-Buschmann, Vector-Borne Zoon. Dis., xxx: "6".. Publication date: 13 April 2019. (Lapsus) ? Epomophorus: - Comments: Not of Gray, 1836. ? Nanonycteris: (Name Combination, Current Combination) TAXONOMY: Revised by Bergmans (1989). (2005: 328). See Simmons Included in the Epomophorinii tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Hassanin et al. (2020: 5) include it in the Epomophorina subtribe of the Epomophorini tribe. Currently (Simmons and Cirranello, 2020) recognized species of Nanonycteris: veldkampii (Jentink, 1888). COMMON NAMES: Czech: Veldkampovi kaloni. English: Dwarf Fruitbats, Little Flying Cow. French: Roussettes naines. German: Zwergflughunde. Nanonycteris veldkampii (Jentink, 1888) *1888. Epomophorus veldkampii Jentink, Notes Leyden Mus., 10: 51 (for 1887). Publication date: April 1888. Type locality: Liberia: Fisherman Lake: Buluma [06 43 N 11 14 W] [Goto Description]. Holotype: RMNH [unknown]: ad ♀, alcoholic (skull not removed). Collected by: Johan Büttikofer and C.F. Sala; collection date: Jan-April 1882. - Etymology: Jentink (1887b: 52) states: "I call this new species veldkampii, as Büttikofer wishes to connect the name of one of his Liberian friends with this new acquisition. Mr. Veldkamp, at present Consul for the Netherlands in Liberia, has helped our travellers as much as he could, has promoted their investigations in every way and finally assisted to Sala's funeral.". 130 ISSN 1990-6471 2000. 2015. 2018. ? ? Nanonycteris weldkampi: Konstantinov, Pema, Labzin and Farafonova, Plecotus et al., 3: 145. (Lapsus) N[anonycteris] Weldkampi: Sylla, Pourrut, Diatta, Diop, Ndiaye and Gonzalez, Afr. J. Microbio. Res., 9 (22): 1464. Publication date: 3 June 2015. (Lapsus) Nanonycteris veldkompi: Gunnell and Manthi, J. Hum. Evol., Suppl.. Publication date: 6 April 2018. (Lapsus) Epomophorus (Nanonycteris) veldkampii: (Name Combination) Nanonycteris veldkampii: (Name Combination, Current Combination) TAXONOMY: See Bergmans (1989) and Simmons (2005: 328). COMMON NAMES: Czech: kaloň Veldkampův. English: Veldkamp's Dwarf Fruit-bat, Veldkamp's Bat, Little Flying Cow, Flying Calf. French: Roussette naine de Veldkamp, Chauve-souris de Veldkamp, Nanonyctère, Petite vache-volante. German: Veldkamps Zwergflughund. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008be; IUCN, 2009; Monadjem et al., 2017ao). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ao). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008be; IUCN, 2009). 2004: LC ver 3.1 (2001) [as Nononycteris veldkampi] (Mickleburgh et al., 2004cb; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: In general there appear to be no current major threats to this species as a whole. The species is dependent on certain food trees where they visit flowers, and habitat degradation might be a problem in parts of the range (Mickleburgh et al., 2008be; IUCN, 2009; Monadjem et al., 2017ao). CONSERVATION ACTIONS: Mickleburgh et al. (2008be) [in IUCN (2009)] and Monadjem et al. (2017ao) report that, in view of the species wide range, it seems probable that it is present in a number of protected areas. There is a need to determine if there have been any local declines in this species resulting from the loss of food trees. GENERAL DISTRIBUTION: Nanonycteris veldkampii is widely distributed in West Africa and western parts of Central Africa. It ranges from Guinea and Sierra Leone in the west, through each country in West Africa to Cameroon, with a single record from the southern region of Central African Republic. It is generally a lowland species, but has been recorded up to 1,200 m asl. Native: Benin (Capo-Chichi et al., 2004: 161); Burkina Faso (Kangoyé et al., 2012: 6023; 2015a: 605); Cameroon; Central African Republic; Côte d'Ivoire (Heim de Balsac, 1934b: 24); Ghana (Decher and Fahr, 2007: 12); Guinea (Denys et al., 2013: 281); Liberia (Monadjem and Fahr, 2007: 50); Nigeria; Sierra Leone; Togo. HABITAT: Monadjem et al. (2016y: 364) indicate that this species occurs in forested habitats between 500 and 1200 m in the Mount Nimba area of Liberia and Guinea. ROOST: N. veldkampii roosts singly or in small groups of well-spaced individuals (Kunz, 1996: 44). MIGRATION: Thomas (1983) [in Kangoyé et al. (2015a: 605)] reported that this species migrates during the wet season from the forest zone to the northern Sudanian zone. DIET: Ayensu (1974) reports feeding visits to Parkia clappertonia (Dawadawa), Ceiba pentandra (SilkCotton tree or Kapok tree), Psidium guajava (Guava tree), and Carica papaya (Papaya or pawpaw). Fahr (2011a: 114) shows a picture of N. veldkampii feeding on Cola cordifolia. Seltzer et al. (2013: table S2) provide an overview of the plant species found beneath bat feeding roosts. For N. veldkampii this included Psidium guajava L. (apple guava - Myrtaceae). African Chiroptera Report 2020 131 POPULATION: Structure and Density:- This species can be abundant, although this partly depends on migration patterns (Mickleburgh et al., 2008be; IUCN, 2009; Monadjem et al., 2017ao). Drexler et al. (2012a: Suppl. Table S1) reported that none of the 23 specimens they examined from Ghana tested positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. Trend:- 2016: Unknown (Monadjem et al., 2017ao). 2008: Unknown (Mickleburgh et al., 2008be; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Gabon, Ghana, Guinea, Liberia, Nigeria, Sierra Leone, Togo. REPRODUCTION AND ONTOGENY: Weber et al. (2019: 20) reported on five pregnant (25, 30, 31 March, 2 April) and 11 lactating (25, 27, 30, 31 March, 1, 3, 7, 8 April) in Sierra Leone. PARASITES: HAEMOSPORIDA Witsenburg et al. (2012: Suppl. Table 1) report on GenBank data for Hepatocystis, retrieved from N. veldkampii. These parasites were also reported by Schaer et al. (2013a: "2") from six out of 8 investigated bats. VIRUSES: Filoviridae - Filoviruses Ebolavirus One out of four specimens tested by Hayman et al. (2012d: 1208) in the Greater Accra region, Ghana, was positive for EBOV antibodies. Figure 31. Distribution of Nanonycteris veldkampii Paramyxoviridae TRIBE Myonycterini Lawrence and Novick, 1963 *1963. Myonycterini Lawrence and Novick, Breviora, 184: 9. Publication date: 18 April 1963. TAXONOMY: Includes the genera Megaloglossus Pagenstecher, 1885 and Myonycteris Matschie, 1899. Genus Megaloglossus Pagenstecher, 1885 *1885. Megaloglossus Pagenstecher, Zool. Anz., 8 (193): 245. Publication date: 27 April 1885 [Goto Description]. - Comments: Type species: Megaloglossus woermanni Pagenstecher, 1885. - Etymology: From the Greek "μέγας" or "μέγαλο", meaning great or large and "γλοσσα", meaning tongue (see Palmer, 1904: 405). (Current Combination) 1891. Trygenycteris Lydekker, in: Flower and Lydekker, Mammals Living and Extinct, 655. Comments: Type species: Megaloglossus woermanni Pagenstecher, 1885. Proposed to replace Megaloglossus, thought to be preoccupied by Megaglossa Rondani, 1865 (Diptera): see Allen (1939a: 63). - Etymology: From the Greek "τρύγη", meaning ripe fruit and "νυκτερίς", meaning bat (see Palmer, 1904: 696). TAXONOMY: Reviewed by Hill (1983). Bergmans (1997: 69) suggests that Megaloglossus should be included in the tribe Myonycterini Lawrence & Novick, 1963 (together with the genera Myonycteris and Lissonycteris), 132 ISSN 1990-6471 within the subfamily Epomophorinae, and not within the Macroglossinae. Included in the Myonycterini tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Hassanin et al. (2020: 5) include it in the Myonycterina subtribe of the Epomophorini tribe. Currently (Simmons and Cirranello, 2020) recognized species of Megaloglossus: azagnyi Nesi, Kadjo and Hassanin, 2012; woermanni Pagenstecher, 1885. COMMON NAMES: Czech: dlouhojazyční kaloni. English: African Long-tongued Fruit-bats, African Nectar Fruit-bats. French: Mégaloglosses. German: Afrikanische Langzungen-Flughunde. VIRUSES: Filoviridae - Filo viruses Ebolavirus Although the distribution area of the Megaloglossus species largely overlaps with the area where Ebola outbreaks occurred, no seropositive individuals were found yet (Olivero et al., 2019: 1). Megaloglossus azagnyi Nesi, Kadjo and Hassanin, 2012 *2012. Megaloglossus azagnyi Nesi, Kadjo and Hassanin, in: Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, Mol. Phylogenet. Evol., 66(1): 134. Type locality: Côte d'Ivoire: Lagunes region, Azagny National Park [0613007N 0501532W, 125m] [Goto Description]. Holotype: MNHN ZM-2011-993: ad ♀, alcoholic (skull not removed). Collection date: 5 December 2009; original number: G09095. Presented/Donated by: [No Name]. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-2011-1000: ad ♂. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Azagny, near; Gboyo. Paratype: MNHN ZM-2011-1001: sad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Azagny, near; Gboyo. Paratype: MNHN ZM-2011-1002: ad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-2011-994: ad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lamto Reserve. Paratype: MNHN ZM-2011995: ad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lamto Reserve. Paratype: MNHN ZM-2011-996: ad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-2011-997: sad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-2011-998: ad ♂. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-2011-999: ad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-G09052: sad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lamto Reserve. Paratype: MNHN ZM-G09070: ad ♂. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Lagunes region; Azagny National Park; 0613007N 0501532W. Paratype: MNHN ZM-G09133: ad ♀. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Côte d'Ivoire; Azagny, near; Gboyo. Paratype: MNHN ZMGAN005: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Locality: Liberia; Mt. Nimba. Paratype: MNHN ZM-GAN028: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. (Current Combination) TAXONOMY: Nesi et al. (2012) examined cytochrome b sequences and found West African Megaloglossus to be significantly different from Central African specimens. African Chiroptera Report 2020 COMMON NAMES: German: Westlicher Woermanns LangzungenFlughund. CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Monadjem, 2016b). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem, 2016b). MAJOR THREATS: It may be locally threatened by habitat loss and degradation, often resulting from conversion of land to agricultural use or the harvesting of timber and firewood (Monadjem, 2016b). CONSERVATION ACTIONS: Monadjem (2016b) reports that this species has been recorded within a number of protected areas and also survives outside of such areas. 133 HABITAT: In the Mount Nimba area, Happold (2013dh: 364) report this species to occur in old growth or secondary forest at relatively low altitudes (450 690 m) on both the Liberian and Guinean sides. DIET: In Côte d'Ivoire, Pettersson (2005) reported "M. woermanni" to feed on nectar and pollen of and Parkia bicolor (African locust-bean - Fabaceae), Maranthes aubrevillei and M. glabra (Chrysobalanaceae). Furthermore, Anthocleista nobilis (Cabbage tree, Cabbage palm Gentianaceae) was the only tree flowering through out the year, and is probably a keystone species in this area for M. woermanni during periods of low food availability. POPULATION: Density and Structure:- The species is locally abundant (Monadjem, 2016b). Trend:- 2016: Unknown (Monadjem, 2016b). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Côte d'Ivoire, Ghana, Liberia, Nigeria, Sierra Leone, Togo. GENERAL DISTRIBUTION: This species occurs in Côte d'Ivoire, Liberia, and probably Guinea and Sierra Leone to Ghana and Togo. In their study, Nesi et al. (2012) did not include specimens from the area between the Dahomey Gap and the Niger delta, but Herkt et al. (2017: Appendix S9) found their maps predicted moderately high habitat suitability for M. azagnyi in this area whereas suitability for M. woermanni was estimated to be very low. Therefore, they assigned the specimens from this area to M. azagnyi. Native (mostly reported as M. woermanni): Benin; Côte d'Ivoire (Heim de Balsac, 1934b: 24); Ghana (Decher and Fahr, 2007: 12); Liberia (Monadjem et al., 2016y: 364); Nigeria (West of the Niger Delta); Sierra Leone; Togo. Figure 32. Distribution of Megaloglossus azagnyi Megaloglossus woermanni Pagenstecher, 1885 *1885. Megaloglossus woermanni Pagenstecher, Zool. Anz., 8 (193): 245. Publication date: 27 April 1885. Type locality: Gabon: Ssibange farm [00 25 N 09 28 E] [Goto Description]. Holotype: ZMB 54589: ad ♀, mounted skin and skull. Collected by: H. Soyaux; collection date: 21 January 1885; original number: 9489. Bergmans (1997: 60, 63) mentions old number "9489", but this is a writing error in the file card; see Turni and Kock (2008). Etymology: In honour of Mr. Adolf Woermann (see Pagenstecher, 1885c: 128). (Current Combination) 1966. Megaloglossus woermanni prigoginei Hayman, in: Hayman, Misonne and Verheyen, Ann. Kon. Mus. Mid. Afr., Zool. Wetensch., (8) 154: 26. Type locality: Congo (Democratic 134 ISSN 1990-6471 1993. 2003. ? Republic of the): Kiliza [03 42 S 28 10 E, 1 370 m]. Holotype: RMCA 32577: ad ♂, skin and skull. Collected by: Dr. Alexandre Prigogine; collection date: 25 May 1964; original number: 1248. See Bergmans (1997: 60). Paratype: RMCA 20429: ♂. Collected by: Dr. Alexandre Prigogine; original number: 494. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 21586: Collected by: Schepens; original number: 6. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32394: juv ♀. Collected by: Dr. Alexandre Prigogine; original number: 1226. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32578: ♀. Collected by: Dr. Alexandre Prigogine; original number: 1249. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32579: ♀. Collected by: Dr. Alexandre Prigogine; original number: 1250. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32580: ♀. Collected by: Dr. Alexandre Prigogine; original number: 1251. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32581: ♀. Collected by: Dr. Alexandre Prigogine; original number: 1252. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32582: ♀. Collected by: Dr. Alexandre Prigogine; original number: 1253. Presented/Donated by: ?: Collector Unknown. Paratype: RMCA 32583: ♀. Collected by: Dr. Alexandre Prigogine; original number: 1254. Presented/Donated by: ?: Collector Unknown. prigoginii: Koopman, in: Wilson and Reeder, Mammal species of the World (2nd Edition): Chiroptera, 154. (Lapsus) Megaloglossus woermani: Giannini and Simmons, Cladistics, 19 (6): 508. (Lapsus) Megaloglossus woermanni woermanni: (Name Combination) TAXONOMY: See Bergmans and Van Bree (1972), Bergmans (1997) and Simmons (2005: 326 - 327). land to agricultural use or the harvesting of timber and firewood (Mickleburgh et al., 2008p; IUCN, 2009; Bakwo Fils et al., 2016). COMMON NAMES: Castilian (Spain): Murciélago de Lengua Larga. Czech: kaloň dlouhojazyčný. English: Woermann's Long-tongued Fruit-bat, Woermann's Bat, Woermann's Fruit bat, Nectar Bat, African Long-tongued Bat. French: Macroglosse de Woermann, Mégaloglosse de Woermann. German: Woermann's Langzungen-Flughund, Afrikanischer Langzungen-Flughund. CONSERVATION ACTIONS: Bakwo Fils et al. (2016) support Mickleburgh et al. (2008p) [in IUCN (2009)] who report that there appear to be no direct conservation measures in place, however, in view of the species wide range it seems possible that it is present within some protected areas. Further studies are needed into the distribution and natural history of this species. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008p; IUCN, 2009; Bakwo Fils et al., 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Bakwo Fils et al., 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008p; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004w; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). GENERAL DISTRIBUTION: Megaloglossus woermanni was formerly considered to occur throughout much of West and Central Africa. However, Nesi et al. (2012) assigned the West African specimens to a separate species, restricting the distribution area to Cameroon and from here south and east into Equatorial Guinea, Congo, Central African Republic, Democratic Republic of the Congo, Angola and western Uganda. Regional None known. In their study, Nesi et al. (2012) did not include specimens from the area between the Dahomey Gap and the Niger delta, but Herkt et al. (2017: Appendix S9) found their maps predicted moderately high habitat suitability for M. azagnyi in this area whereas suitability for M. woermanni was estimated to be very low. Therefore, they assigned the specimens from this area to M. azagnyi. MAJOR THREATS: It is locally threatened by habitat loss and degradation, often resulting from conversion of Native: Angola (Crawford-Cabral, 1989; Bergmans, 1997; Monadjem et al., 2010d: 554); Cameroon; Central African Republic (Morvan et African Chiroptera Report 2020 135 al., 1999: 1195); Congo (King and Dallimer, 2010: 65); Congo (The Democratic Republic of the) (Schouteden, 1944; Dowsett et al., 1991: 259; Bergmans, 1997; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 554); Equatorial Guinea (Fahr and Ebigbo, 2003: 128); Gabon; Nigeria (East of the Niger delta); Uganda (Kityo and Kerbis, 1996: 58). DIET: Pettersson et al. (2004: 162) indicate that M. woermanni is the only nectar specialist in Africa, whereas there are 14 such specialists in Asia, Australia and Polynesia. Presence uncertain: Bioko. In Cameroon, Grünmeier (1993: 35) reported the following pollen from the gut and faeces: Anthocleista sp. (Gentianaceae), Mucuna flagellipes Hook.f. (Fabaceae), Kigelia africana (Lam.) Benth. (Bignoniaceae), Celtis sp. (Cannabaceae), Citrus sp.(Rutaceae), Bombacaceae indet. DETAILED MORPHOLOGY: Chawana et al. (2013: 160) indicate that the average brain mass is 0.60 g (n = 2). FUNCTIONAL MORPHOLOGY: Coimbra et al. (2016: 191) mention that the retinal area covers about 40 - 50 mm2. The eye contains about 90,000 retinal ganglion cells. The minimum angle of resolution was about 0.227° (ca. 8 mm at 1 m distance). Kulzer and Storf (1980: 409) investigated the thermoregulatory response by measuring body temperature and rate of oxygen consumption. At high ambient temperatures, the bats increase their oxygen consumption and at low ambient temperatures, they enter a lethargic sleep. MOLECULAR BIOLOGY: DNA - Hollar and Springer (1997), Juste B. et al. (1999), Teeling et al. (2000), Colgan and da Costa (2002) and Cantrell et al. (2008). Karyotype - Haiduk et al. (1980, 1981: 228) reported 2n = 34, FN = 62, BA = 30, a metacentric X chromosome and a metacentric Y chromosome for specimens from Cameroon. The same karyotype was reported by Primus et al. (2006) for specimens from Gabon. Protein / allozyme - Unknown. HABITAT: Juste B. and Perez del Val (1995: 144) found M. woermanni to occur to 1,800 m altitude on Bioko Island, where it occurred more frequently in cultivated areas than in lowland rainforests. HABITS: Weber et al. (2009) found that spatial use was characterized by small home ranges and high sitefidelity, mean home range sizes (minimum convex polygon) were larger in females (139.0 and 146.8 ha) than in males (99.8 and 102.9 ha) in Lama Forest Reserve, southern Benin. Kulzer and Storf (1980) [in Lovegrove (2011: 152)] indicate that M. woermanni uses daily torpor. Weber et al. (2009) frequently observed M. woermanni visiting flowers of cultivated bananas. In Uganda, Ssali and Sheil (2019: 1) found M. woermanni to be the second most common flower visitor of African wild bananas or “false bananas" (Ensete ventricosum (Welw.) Cheesman (Musaceae)), after the African dormouse (Graphiurus murinus). As this plant produces flowers (and fruits) year-round, it probably will act as a food resource in periods of the year when other types of food are not available. POPULATION: Structure and Density:- It is a locally abundant species (Mickleburgh et al., 2008p; IUCN, 2009; Bakwo Fils et al., 2016). Trend:- 2016: Stable (Bakwo Fils et al., 2016). 2008: Stable (Mickleburgh et al., 2008p; IUCN, 2009). REPRODUCTION AND ONTOGENY: Czekala and Benirschke (1974: 220) mention an adult female, collected in January 1973 by M. Eisentraut, containing two foetuses of different size, probably due to implantation occurring at two different times (p. 229). Two females collected in Dimonika (Congo) on 10 March were pregnant with a large embryo (Bergmans, 1979a: 181). PARASITES: BACTERIA Gram-negative Enterobacteriae: Mbehang Nguema et al. (2020: 7) reported the presence of E. coli and Klebsiella pneumoniae in feacal samples collected near Makokou in Gabon. VIRUSES: Filoviridae - Filo viruses Marburgvirus Towner et al. (2007) tested 37 individuals from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and 20 individuals for anti- 136 ISSN 1990-6471 Marburg virus IgG antibodies by ELISA; no positives were found. Flaviviridae Pegivirus (BPgV) - Quan et al. (2013: Table S5) reported 1 out of 3 examined specimens from the Democratic Republic of the Congo to be infected by clade H type Pegivirus. Paramyxoviridae Orthorubulavirus Drexler et al. (2012a: Suppl. Table S1) reported that one of the 34 specimens they examined from Gabon, Congo, the Democratic Republic of the Congo, the Central African Republic, and Ghana tested positive for a Rubulavirus that was found to be closely related to human mumps virus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Equatorial Guinea, Gabon, Nigeria, Uganda. Figure 33. Distribution of Megaloglossus woermanni Genus Myonycteris Matschie, 1899 *1899. [Xantharpyia (]Myonycteris[)] Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, 61, 63 [Goto Description]. - Comments: Type species: Cynonycteris torquata Dobson, 1878. Originally described as subgenus of Xantharpyia. Matschie dated his introduction on 14 May 1899. - Etymology: Derived from the Greek "mys" (= mouse) and "nycteris" (= bat) meaning mouse-like bat. 1912. Myonycteris: K. Andersen. - Etymology: From the Greek "μύξ" or "μυόξ", meaning mouse and "νυκτερισ", meaning bat (see Palmer, 1904: 440). (Name Combination, Current Combination) 2012. Lissonycteris: Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, Mol. Phylogenet. Evol., 66 (1): 126. 2015. Cyonycteris: Lanza, Funaioli and Riccucci, The bats of Somalia and neighbouring areas, 60. - Etymology: From the Greek adjective "λισός" (lissós), meaning "smooth" and the feminine Greek substantive "νυκτερίς" (nykterís) [genitive "νυκτερίδος" (nikterídos)] meaning "bat", or combined "smooth bat", referring to the smooth fur of the species (see Lanza et al., 2015: 59). (Lapsus) 2019. Myonicteris: Nziza, Goldstein, Cranfield, Webala, Nsengimana, Nyatanyi, Mudakikwa, Tremeau-Bravard, Byarugaba, Tumushime, Mwikarago, Gafarasi, Mazet and Gilardi, EcoHealth, 17: 155. Publication date: 6 Decembe 2019. (Lapsus) TAXONOMY: Revised by Bergmans (1976; 1997). Lawrence and Novick (1963) separated Lissonycteris from Rousettus because of ethological differences concerning the use of the limbs, and the absence of echolocation in Lissonycteris. Juste B. et al. (1997) presented allozyme evidence corroborating earlier chromosome studies (Haiduk et al. (1980, 1981), DNA-hybridisation results (Kirsch et al., 1995) and cladistic analysis (Springer et al., 1995) that support the recognition of Lissonycteris as a distinct genus. Bergmans (1994: 79; 1997: 136), Lavrenchenko et al. (2004b: 136), and Simmons (2005: 325) also consider Lissonycteris to be a valid genus. Included in the Myonycterini tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Included in the Myonycterina subtribe of the Epomophorini tribe by Hassanin et al. (2020: 5). Currently (Simmons and Cirranello, 2020) recognized subgenera and species of Myonycteris: African Chiroptera Report 2020 Myonycteris Matschie, 1899: angolensis (Bocage, 1898); leptodon K. Andersen, 1908; relicta Bergmans, 1980; torquata (Dobson, 1878). Phygetis K. Andersen, 1912: brachycephala (Bocage, 1889). COMMON NAMES: Czech: límcoví kaloni, měkkosrstí kaloni. English: Collared Fruit-bats, Little Collared Fruitbats, Dwarf Epauletted Bat, Lesser Epauletted Bat, Smooth-haired Fruit-bats, Soft-furred Fruitbats. French: Lissonyctères, Myonyctères, Roussettes à collier. German: HalskrausenFlughunde, Samtfell-Flughunde. 137 parasitized by a Dipseliopoda bat fly (Nycteribiidae), which was infected by Kanyawara virus (KYAV). VIRUSES: Goldberg et al. (2017: 3, 4) described a new virus from Uganda (Kanyawara virus, genus Ledantevirus, Rhabdoviridae, Mononegavirales) from a new nycteribiid bat fly (genus Dipseliopoda Theodor, 1955, which in turn was captured on a (probably) new species of bat belonging to the genus Myonycteris. The bat was originally identified as M. angolensis, but an analysis of its mitochondrial DNA showed it to belong to an outgroup. PARASITES: Szentiványi et al. (2019: Suppl.) indicated that a non specified Ugandan Myonycteris was Myonycteris angolensis (Bocage, 1898) *1898. Cynonycteris Angolensis Bocage, J. Sci. mat. phys. nat., ser. 2, 5 (19): 133, 138, text-fig. Publication date: June 1898. Type locality: Angola: Quibula, Cahata, north of Quanza River: Pungo Andongo [09 40 S 15 35 E, 1 200 m] [Goto Description]. Syntype: BMNH 1897.8.6.1: juv ♀, skull and alcoholic. Collected by: José Alberto de Oliveira Anchieta; collection date: 1891. Presented/Donated by: Lisbon Museum. See Andersen (1912b: 54), Bergmans (1997: 37). Syntype: MLZA T113 [a]: ad ♀, mounted skin (skull missing). Collected by: José Alberto de Oliveira Anchieta; collection date: 1891. Destroyed by fire in 1978; see Bergmans (1997: 37). Syntype: MLZA T113 [b]: juv, mounted skin (skull not removed). Collected by: José Alberto de Oliveira Anchieta; collection date: 1891. Destroyed by fire in 1978; see Bergmans (1997: 37). - Comments: From the feminine scientific Latin adjective angolènsis, meaning "Angolan" referring to the country where the type specimen was collected (see Lanza et al., 2015: 60). - Etymology: Referring to the country where the type specimen was collected. 1920. Rousettus (Lissonycteris) crypticola Cabrera, Bol. r. Soc. espan. Hist. nat. , 20 (3 - 4): 106. Type locality: Equatorial Guinea: Bioko: Malabo, Basilé: San Fernando, cave of [03 44 N 08 49 E, 1 100 m]. Holotype: MNCN 53 / 20-II-25-5: ad ♀, complete skeleton. Collected by: D. Manuel M. de la Escalera; collection date: 26 July 1919; original number: 2.057. See Cabrera (1920: 107); Ibáñez and Fernández (1989: 7); Bergmans (1997: 35). Publication date: March - April 1920. Paratype: MNCN 54: juv ♂, skin only. Collected by: D. Manuel M. de la Escalera; collection date: 26 July 1919. See Ibáñez and Fernández (1989: 7); Bergmans (1997: 35). 2007. Lyssonycteris angolensis: Müller, M. A.; Paweska, J. T.; Leman, P. A.; Drosten, C.; Grywna, K.; Kemp, A.; Braack, L. E. O.; Sonnenberg, K.; Niedrig, M.; Swanepoel, R., Emerg. Inf. Dis., 13 (9): 1368. (Lapsus) 2013. Myonycteris angolensis: Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, Mol. Phylogenet. Evol., 66 (1): 135. Publication date: 10 October 2012. (Name Combination, Current Combination) 2019. Myonicteris angolensis: Nziza, Goldstein, Cranfield, Webala, Nsengimana, Nyatanyi, Mudakikwa, Tremeau-Bravard, Byarugaba, Tumushime, Mwikarago, Gafarasi, Mazet and Gilardi, EcoHealth, 17: 155. Publication date: 6 December 2019. (Lapsus) ? Lissonycteris angolensis: (Name Combination) ? Rousettus (Lissonycteris) angolensis: (Name Combination) ? Rousettus angolensis angolensis: (Name Combination) ? Rousettus angolensis: 138 ISSN 1990-6471 TAXONOMY: Peterson et al. (1995: 48), Schaer et al. (2013a), and Kerbis Peterhans et al. (2013: 189) include angolensis in the genus Myonycteris. Simmons (2005: 325) recognizes goliath, petraea, ruwenzorii and smithii as subspecies of angolensis, indicating that more than one species may be present in this complex. Known subspecies of M. angolensis: angolensis (Bocage, 1898); goliath Bergmans, 1997; petraea Bergmans, 1997; smithii (Thomas, 1908). COMMON NAMES: Afrikaans: Bocage se vrugtevlermuis, Bocagevrugtevlermuis. Castilian (Spain): Ruseta de Angola. Chinese: 安哥拉领果蝠. Czech: kaloň měkkosrstý. English: Angolan Soft-furred Fruitbat, Angolan Long-haired Fruit-bat, Angolan Rousette, Angolan Fruit Bat, Bocage's Fruit-bat, Long-haired Rousette, Canon Smith's Rousette. French: Lissonyctère d'Angola, Roussette d'Angola, Roussette de Bocage, Roussette à longs poils. German: Angolanischer SamtfellFlughund, Angola-Weichfellflughund. Italian: Lissonittèride angolàna, Lissonictèride angolàna. Kiluba (DRC): Mulima. Kinande (DRC): Mulima. Portuguese: Morcego Frugivoro de Bocage. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) [as Lissonycteris angolensis] in view of its wide distribution, presumed large population, its occurrence in a number of protected areas, its tolerance of a degree of habitat modification, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008o; IUCN, 2009). Assessment History Global 2008: LC ver 3.1 (2001) [assessed including L. goliath, L. petraea, L. ruwenzorii, L. smithii as synonyms] (Mickleburgh et al., 2008o; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004z; IUCN, 2004). 2003: LR/lc [assessed as Rousettus angolensis] (IUCN, 2003). 1996: LR/lc [assessed as Rousettus angolensis] (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There appear to be no major threats to this species as a whole (Mickleburgh et al., 2008o; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008o) [in IUCN (2009)] report that this species is present in a number of protected areas. GENERAL DISTRIBUTION: Myonycteris angolensis is widely distributed at elevations ranging from sea level to 4,000 m asl, in West Africa, Central Africa and East Africa, with some distinct populations present within southern Africa. It ranges from Senegal and The Gambia in the west, through most of West Africa to Cameroon; from here it ranges southwards into Congo, eastern Democratic Republic of the Congo and Angola, and eastwards into Central African Republic, southern Sudan, eastern and southern Democratic Republic of the Congo and Rwanda and Burundi. In East Africa it is distributed from Ethiopia in the north through Uganda, Kenya and Tanzania. In southern Africa has been recorded from northwestern Angola, central and southern Cameroon, Central African Republic, central Congo, Equatorial Guinea (Bioko, Mbini), Gabon, eastern Nigeria, and northwestern and southwestern Democratic Republic of the Congo. Native: Angola (Crawford-Cabral, 1989; Bergmans, 1997; Monadjem et al., 2010d: 554); Burkina Faso (Kangoyé et al., 2015a: 605); Burundi; Cameroon; Central African Republic; Congo (Bates et al., 2013: 333); Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 554); Côte d'Ivoire; Equatorial Guinea; Ethiopia; Gambia; Ghana; Guinea (Denys et al., 2013: 281); Guinea-Bissau (Veiga-Ferreira, 1948; Lopes and CrawfordCabral, 1992; Rainho and Ranco, 2001: 35); Kenya; Liberia; Mozambique; Nigeria; Rwanda; Senegal; Sierra Leone; Sudan; Tanzania (Stanley and Goodman, 2011: 40); Togo; Uganda; Zambia (Ansell, 1967; Ansell, 1974; Monadjem et al., 2010d: 554); Zimbabwe. Presence uncertain: Benin; Gabon. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data for 8 specimens: Fa: 77.70 ± 1.634 mm, mass: 78.1 ± 7.31 g, wing loading: 17.7 ± 1.52 N/m 2, aspect ratio: 15.9 ± 0.46. An albino specimen - of "Lissonycteris angolensis?" - was reported from a cave in Uganda by Neal (1971) and Lucati and LópezBaucells (2016: Suppl.). DETAILED MORPHOLOGY: Chernova (2002) studied the structure of the cuticle of guard hairs of several Pteropodid bats and found that the structure of these hairs in African Chiroptera Report 2020 angolensis was nonring-forming as in Epomophorus wahlbergi as oposed to ring-forming as in Myonycteris angolensis [as Rousettus]. FUNCTIONAL MORPHOLOGY: Mainoya and Howell (1979: 164) found no glandular cells in the neck skin patch of male "Rousettus angolensis", and concluded that this patch serves as a visual cue during social encounters. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Haiduk et al. (1980, 1981: 223) reported 2n = 36, FN = 66, BA = 32, and a subtelocentric X chromosome. Denys et al. (2013: 281) also reported 2n = 36 and Fna = 66, but with two medium-sized submetacentric X chromosomes. Protein / allozyme - Unknown. ROOST: Weber and Fahr (2006: 4) indicate that [as Lissonycteris angolensis] largely or exclusively depends on the availability of caves as day roosts. POPULATION: Structure and Density:- This species can be common in some localities (Mickleburgh et al., 2008o; IUCN, 2009). Trend:- 2008: Decreasing (Mickleburgh et al., 2008o; IUCN, 2009). REPRODUCTION AND ONTOGENY: Stanley and Goodman (2011: 38) reported pregnant females collected in the West and East Usambara Mountains (Tanzania) on 1 July 1991, and between 25 July and 5 August 1992. They also found six pregnant females in the South Pare Mountains between 18 and 23 July 1993, and on 25 August 1994. In Sierra Leone, Weber et al. (2019: 20) captured two lactating females on 25 March. PARASITES: Adam (1973: 3) reported Plasmodium voltaicum Van Der Kaay, 1964 (Haemosporida) from the Republic of Congo. He also mentioned that this parasite can be detected in the blood of the bats six to eight days after infection and then persists chronically for an persistent period. Schaer et al. (2013a: 17416) recorded the same parasite in three out of three investigated bats, and Perkins and Schaer (2016: Suppl.) mention it furthermore from Ghana and Guinea. Adam (1973: 4) also reported on a [as Lissonycteris angolensis] specimen that was 139 transported from the Republic of Congo to Paris and that was infected by Hepatocystis perronae. ACARI Teinocoptidae; Fain (1967: 363) described Teinocoptes ituriensis from "Rousettus (Lissonycteris) angolensis" from Saliboko, Lake Albert, Ituri, Congo. Trombiculidae: Trombigastia (Trombigastia) rousetti was described by Vercammen-Grandjean and Fain (1958: 16) from "Rousettus (Lissonycteris) angolensis" from Nyakibanda, Rwanda (Stekolnikov, 2018a: 120). They also described (p. 18) Trombigastia (Trombigastia) hirsuta from bats from Katana, N of Bukavu, DRC (Stekolnikov, 2018a: 118). Fain (1959a: 346) mentions Nycteridocoptes macrophallus Fain, 1958 from "R. angolensis" from Mulungu, DRC. Stekolnikov (2018a: 130, 152) also reported Chiroptella adami Taufflieb, 1972 on "Lissonycteris angolensis smithii" from Senegal, and Microtrombicula hexasternalis VercammenGrandjean, 1965 on bats from Bukavu, DRC. DIPTERA Nycteribiidae: Eucampsipoda africanum Theodor 1955, widely distributed over the Ethiopian region and recorded from localities in Senegal to the Sudan and southwards mirroring the hosts distribution (Haeselbarth et al., 1966: 115). Dipseliopoda biannulate (Oldroyd 1953) distributed over the tropical parts of Africa and recorded from localities in Nigeria, Cameroon, the Congo and the Sudan and Kenya (Haeselbarth et al., 1966: 116). Cyclopodia greeffi Karsch 1884 found in a belt across central Africa, between lat. 10o south and north, while in West Africa it occurs further north up to lat. 20o (Haeselbarth et al., 1966: 117). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) [as Lyssonycteris angolensis] tested between 1986 to 1999, for antibody to SARS-CoV in sera in 16 individuals from Oriental Province, DRC, one was tested positive (1/16, 6.3 %) and two individuals from Bandundu Province, DRC, none tested positive (0/2) (1/18, 5.6 %). None of the nine Kenyan specimens tested by Tao et al. (2017: Suppl.) were positive for CoV. Nziza et al. (2019: 157) found a new Coronavirus in a rectal swap of a "Rousettus angolensis" from the Nyungwe National Park, Rwanda. Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that none of the 3 specimens they examined from the Democratic Republic of the Congo tested 140 ISSN 1990-6471 positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. Polyomaviridae Polyomavirus - Conrardy et al. (2014: 259) tested 11 specimens from the Kakamega cave (Kenya) and found one of them to test positive for this virus. This was also the case for the single specimen from Kisumu. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Burkina Faso, Burundi, Cameroon, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ethiopia, Ghana, Guinea, Kenya, Liberia, Malawi, Mozambique, Nigeria, Rwanda, Senegal, Sierra Leone, Tanzania, Togo, Uganda, Zambia. Figure 34. Distribution of Myonycteris angolensis Myonycteris angolensis angolensis (Bocage, 1898) *1898. Cynonycteris Angolensis Bocage, J. Sci. mat. phys. nat., ser. 2, 5 (19): 133, 138, text-fig. Publication date: June 1898. Type locality: Angola: Quibula, Cahata, north of Quanza River: Pungo Andongo [09 40 S 15 35 E, 1 200 m] [Goto Description]. Syntype: BMNH 1897.8.6.1: juv ♀, skull and alcoholic. Collected by: José Alberto de Oliveira Anchieta; collection date: 1891. Presented/Donated by: Lisbon Museum. See Andersen (1912b: 54), Bergmans (1997: 37). Syntype: MLZA T113 [a]: ad ♀, mounted skin (skull missing). Collected by: José Alberto de Oliveira Anchieta; collection date: 1891. Destroyed by fire in 1978; see Bergmans (1997: 37). Syntype: MLZA T113 [b]: juv, mounted skin (skull not removed). Collected by: José Alberto de Oliveira Anchieta; collection date: 1891. Destroyed by fire in 1978; see Bergmans (1997: 37). - Comments: From the feminine scientific Latin adjective angolènsis, meaning "Angolan" referring to the country where the type specimen was collected (see Lanza et al., 2015: 60). - Etymology: Referring to the country where the type specimen was collected. 2015. Myonycteris angolensis angolensis: ACR. (Name Combination, Current Combination) ? Lissonycteris angolensis angolensis: (Name Combination) GENERAL DISTRIBUTION: Recorded from northwestern Angola, central and southern Cameroon, Central African Republic, central Congo, Equatorial Guinea [Bioko, Mbini], Gabon, eastern Nigeria, and northwestern and southwestern Democratic Republic of the Congo. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Cameroon, Congo, Congo (Democratic Republic of the), Equatorial Guinea, Nigeria. Myonycteris angolensis goliath (Bergmans, 1997) *1997. Lissonycteris angolensis goliath Bergmans, Beaufortia, 47 (2): 46. Publication date: 20 June 1997. Type locality: Zimbabwe: Inyanga: Gleneagles (Estate) [18 17 S 32 54 E] [Goto Description]. Holotype: NHMZ 59831: ad ♀, skin and skull. Collected by: Michael P. Stuart Irving et al.; collection date: 19 March 1970; original number: T1854. 8 paratypes; see Bergmans (1997: 46). Paratype: ZMA MAM.24710: ad ♀, skin and skull. Collected by: S. Irwin; collection date: 19 March 1970; original number: T1853. Presented/Donated by: ?: Collector Unknown. Paratype: ZMA MAM.24719: ♀, skin and skull. Collected by: S. Irwin; collection date: 19 March 1970; original number: T1853. Presented/Donated by: ?: Collector Unknown. Locality: Gleneagles (18 15 - 18 30 S, 32 45 - 33 0 E, 6000ft), see Bergmans (2011: 841). - Etymology: Referring to the biblical giant with the same name. African Chiroptera Report 2020 2001. 2015. 141 Lissonycteris goliath: Cotterill, Durban Mus. Novit., 26: 53. (Name Combination, Current Combination) Myonycteris angolensis goliath: ACR. (Name Combination, Current Combination) TAXONOMY: Bergmans (1997) showed that craniological characters also distinguish Lissonycteris from Rousettus, and assigned the specimens from Zimbabwe to a new subspecies (L. a. goliath). Liberally applying the evolutionary species concept, which holds that any diagnosable or allopatric population is a valid species, Cotterill (2001e) elevated goliath to full species rank. Bergmans (2011: 841) still considers it a subspecies, at most. COMMON NAMES: English: Harrison's fruit bat. French: Lissonyctère goliath. German: Harrisons Samtfell-Flughund. CONSERVATION STATUS: Global Justification Assessment History Global VU A3c+4c ver 3.1 (2001) (IUCN, 2004; Mickleburgh et al., 2004ar); LC ver 3.1 (2001) included as a synonym of L. angolensis (Mickleburgh et al., 2008o; IUCN, 2009). MAJOR THREATS: May be threatened by loss of habitat resulting from logging operations, and disturbance of cave roosting sites. It is possible that some populations of this species are threatened by over harvesting for subsistence food (Mickleburgh et al., 2008o; IUCN, 2009). CONSERVATION ACTIONS: There is a need to protect the habitat (Mickleburgh et al., 2008o; IUCN, 2009). GENERAL DISTRIBUTION: Lissonycteris goliath is known from four localities in south-east Africa (the border area of Mozambique and Zambia) between sea level and 1,800 m asl. Native: Mozambique (Monadjem et al., 2010d: 554; Monadjem et al., 2010c: 376); Zambia; Zimbabwe (Monadjem et al., 2010d: 554). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Malawi, Mozambique, Zimbabwe. Myonycteris angolensis petraea (Bergmans, 1997) *1997. Lissonycteris angolensis petraea Bergmans, Beaufortia, 47 (2): 44. Publication date: 20 June 1997. Type locality: Ethiopia: Ilubabor province: 10 km from Agaro: Jimma, road to [07 50 N 36 38 E] [Goto Description]. Holotype: MNHN ZM-MO-1972-482a: ad ♂, skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4017. Presented/Donated by: ?: Collector Unknown. 10 paratypes; see Bergmans (1997). Paratype: MNHN ZM-MO-1972-482b: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4018. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972-482c: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4019. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972-482d: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4024. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972-482e: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4027. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972482f: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4032. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972-482g: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4044. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972-482h: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4045. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972482i: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4046. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1972-482j: skull and alcoholic. Collected by: Jean Pierre Dorst et al.; collection date: 10 - 13 June 1968; original number: 4051. Presented/Donated by: ?: Collector Unknown. Paratype: ZMA MAM.25180: ad ♂, skull and alcoholic. Collected 142 ISSN 1990-6471 2001. 2015. by: Jean Pierre Dorst et al.; collection date: 10-13 June 1968. Presented/Donated by: ?: Collector Unknown. Locality: 10 km from Agaro (07 50 N 36 38 E), at the road to Jimma (07 39 N 36 47 E), Ethiopia, 1650 m (see: Bergmans, 2011: 842). Lissonycteris petraea: Cotterill, Durban Mus. Novit., 26: 53. (Name Combination) Myonycteris angolensis petraea: ACR. (Name Combination, Current Combination) TAXONOMY: Simmons (2005: 325) states that the status of petraea remains unclear, but recognized it as a subspecies of angolensis. COMMON NAMES: English: Petra fruit bat. French: Lissonyctère d'Ethiopie. German: Äthiopischer SamtfellFlughund. CONSERVATION STATUS: Assessment History Global VU A4c ver 3.1 (2001) (IUCN, 2004; Mickleburgh et al., 2004ab); LC ver 3.1 (2001) included as a synonym of L. angolensis (Mickleburgh et al., 2008o; IUCN, 2009). CONSERVATION ACTIONS: There is a need to protect the habitat (Mickleburgh et al., 2008o; IUCN, 2009). GENERAL DISTRIBUTION: Found only in Ethiopia, where it has been recorded from six localities between 1,190 and 2,600 m asl. Native: Ethiopia. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia. Myonycteris angolensis ruwenzorii (Eisentraut, 1965) *1965. Rousettus angolensis ruwenzorii Eisentraut, Bonn. zool. Beitr., 16 (1/2): 3, fig. 1. Type locality: Uganda: Ruwenzori East [1 575 m] [Goto Description]. Holotype: BMNH 1906.12.4.1: ♂. Presented/Donated by: ?: Collector Unknown. - Etymology: Referring to the country where the type specimen was collected. 2015. Myonycteris angolensis ruwenzorii: ACR. (Name Combination, Current Combination) ? Lissonycteris angolensis ruwenzorii: (Name Combination) COMMON NAMES: English: Ruwenzori Long-haired rousette. French: Lissonyctère du Ruwenzori, Roussette à long poils du Ruwenzori. German: RuwenzoriLanghaarflughund. GENERAL DISTRIBUTION: M. a. ruwenzorii has been recorded from Ethiopia, Kenya, Mozambique, Rwanda, southern Sudan, Tanzania, Uganda, northeastern and southeastern Democratic Republic of the Congo, Zambia and Zimbabwe. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burundi, Cameroon, Congo (Democratic Republic of the), Ethiopia, Kenya, South Sudan, Sudan, Tanzania, Uganda. Myonycteris angolensis smithii (Thomas, 1908) *1908. Rousettus smithii Thomas, Ann. Mag. nat. Hist., ser. 8, 2 (10): 375. Publication date: 1 October 1908. Type locality: Sierra Leone: Freetown [08 30 N 13 15 W] [Goto Description]. Holotype: BMNH 1908.9.11.1: sad ♀. Collected by: Canon F.C. Smith. Presented/Donated by: Canon F.C. Smith. - Comments: Grubb et al. (1998: 67) mention that the type locality is almost certainly Freetown, according to Rosevear (1965: 84). 1974. Lissonycteris angolensis smithi: Bergmans, Bellier and Vissault, Rev. Zool. afr., 88 (1): 38. (Name Combination) 2012. Myonycteris angolensis smithii: Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, Mol. Phylogenet. Evol., suppl. 3. (Name Combination, Current Combination) ? Lissonycteris angolensis smithii: (Name Combination) ? Lissonycteris smithii: (Name Combination) ? Rousettus angolensis smithi: (Name Combination) ? Rousettus smithi: (Name Combination, Alternate Spelling) African Chiroptera Report 2020 TAXONOMY: Mentioned as smithii by Allen (1939a: 63), Meester et al. (1986: 30), Koopman (1993a), Bergmans (1997: 33), Grubb et al. (1998: 67), Kock et al. (2002: 79), but as smithi by Peterson et al. (1995: 48), who also consider it a valid species in the genus Myonycteris. Kock et al. (2002: 79) include it as subspecies in angolensis, due to clinal differentiation (as does Bergmans, 1997: 33). Simmons (2005: 325) follows Kock et al. (2002), who support Bergmans (1997), in treating smithii as a subspecies of angolensis. COMMON NAMES: English: Smith's fruit bat. French: Lissonyctère de Simth. German: Smiths Samtfell-Flughund. CONSERVATION STATUS: Assessment History Global LC ver 3.1 (2001) (IUCN, 2004; Mickleburgh et al., 2004u); LC ver 3.1 (2001) included as a synonym 143 of L. angolensis (Mickleburgh et al., 2008o; IUCN, 2009). GENERAL DISTRIBUTION: Distributed in the lowlands of western Africa, where its distribution follows the savanna and forest zone, it is found up to 1,400 m asl on Mount Nimba. Native: Bioko; Burkina Faso; Ghana; Guinea (Decher et al., 2016: 262); Guinea Bissau; Côte d'Ivoire; Liberia; Nigeria; Sierra Leone; Senegal; Togo. HABITAT: Monadjem et al. (2016y: 364) indicate that the (sub)species is restricted to montane forests in West Africa. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burkina Faso, Côte d'Ivoire, Ghana, Guinea, Liberia, Nigeria, Sierra Leone, The Gambia, Togo. Myonycteris brachycephala (Bocage, 1889) *1889. Cynonycteris brachycephala Bocage, J. Sci. mat. phys. nat., ser. 2, 1 (3): 198. Publication date: December 1889. Type locality: São Tomé and Principé: São Tomé Island. Holotype: MLZA 449a: ad ♀, mounted skin and skull. Collected by: F. Newton and Pires; collection date: 1868. Holotype number "T114"; destroyed by fire in 1978: see Bergmans (1997: 53). 1898. Cynonycteris brachycephalus: Seabra, J. Sci. mat. phys. nat., ser. 2, 5: 170. (Alternate Spelling) 1995. M[yonycteris] brachyptera: Peterson, Eger and Mitchell, : 48. - Comments: Mentioned by Peterson et al. (1995: 48) only with the remark "La situation exacte de ce taxon suppose une étude plus poussée". [The exact situation of this taxon needs a more thorough study]. (Lapsus) ? Myonycteris (Myonycteris) brachyptera: (Lapsus) ? Myonycteris (Phygetis) brachycephala: (Name Combination) ? Myonycteris brachycephala: (Name Combination, Current Combination) TAXONOMY: Reviewed by Bergmans (1997). (2005: 328). See Simmons COMMON NAMES: Czech: kaloň tomášský. English: Sao Tomé Collared Fruit Bat. French: Myonyctère de São Tomé, Chauve-souris frugivore à collier de São Tomé. German Halskrausen-Flughund, São Tomé Halskrausen-Flughund. CONSERVATION STATUS: Global Justification It is assessed Endangered because its extent of occurrence (EOO) is 750 km 2, it is known from less than five locations, and there is continuing decline in habitat quality inferred due to intense logging activities and the conversion of land to agricultural use (Juste, 2016b). This is a is range-restricted species. It is endemic to the islands of São Tomé and Príncipe, where it is known from a single location (Juste et al., 2008a; IUCN, 2009; Juste, 2016b). Assessment History Global 2016: EN B1ab(iii) ver 3.1 (2001) (Juste, 2016b). 2008: EN B1ab(iii) ver 3.1 (2001) (Juste et al., 2008a; IUCN, 2009). 2004: EN B1ab(iii) ver 3.1 (2001) (Juste et al., 2004b; IUCN, 2004). 2000: EN (Hilton-Taylor, 2000). 1996: VU (Baillie and Groombridge, 1996). 1994: VU (Groombridge, 1994). Regional None known. 144 ISSN 1990-6471 MAJOR THREATS: Within the species' relatively limited range it is threatened by ongoing habitat loss, presumably related to logging activities and the conversion of land to agricultural use (Juste et al., 2008a; IUCN, 2009; Juste, 2016b). Rainho et al. (2010b: 257) indicate that hunting might also be a problem for this island endemic. CONSERVATION ACTIONS: Juste et al. (2008a) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas, although is expected in mountain forest within the protected area of Lagoa Amelia (Juste, 2016b). There is a need to protect suitable areas of remaining natural habitat for this species, with additional research required into the persistence of populations in modified habitats (such as cocoa plantations). Further surveys are needed to identify additional localities for this species, and to better understand its natural history (Juste et al., 2008a; Juste, 2016b). indicate that this might be related to a developmental instability associated with the colonization by the island species. POPULATION: Structure and Density:- It appears to be a naturally rare species (Juste et al., 2008a; IUCN, 2009). Nothing is known about the social structure of the species and it seems to show no sexual dimorphism. Individuals were always captured in mountain passes along most possibly commuting flights. Recent bat surveys with mistnets in primary and secondary forests as well as in cocoa plantations have failed to capture the species again (C. Meyer, pers. comm in Juste (2016b)). Trend:- 2016: Decreasing (Juste, 2016b). 2008: Decreasing (Juste et al., 2008a; IUCN, 2009; Rainho et al., 2010b: 257). DISTRIBUTION BASED ON VOUCHER SPECIMENS: São Tomé and Principé. GENERAL DISTRIBUTION: Myonycteris brachycephala is endemic to the uplands of São Tomé Island of São Tomé and Príncipe. It is absent from the northern part of the island. It has only been recorded from three localities, and no colonies have been reported. Individuals have been recorded between 300 and 1,200 m asl. Native: Sao Tomé and Principe (Bergmans, 1997; Simmons, 2005: 328; Rainho et al., 2010a: 18). DENTAL FORMULA: Juste and Ibáñez (1993a: 221) report on an asymmetric dental formula in M. brachycephala, which resulted from the absence of a lower internal incisor, either in the left or right mandible. They Figure 35. Distribution of Myonycteris brachycephala Myonycteris leptodon K. Andersen, 1908 *1908. Myonycteris leptodon K. Andersen, Ann. Mag. nat. Hist., ser. 8, 2 (11): 450. Publication date: 1 November 1908. Type locality: Sierra Leone: "Sierra Leone" [Goto Description]. Holotype: BMNH 1891.2.13.1: ad ♂. Collected by: J. Hickman Esq. Collection date: prior to 1891. (Current Combination) 1971. M[yonycteris (Myonycteris)] t[orquata] leptodon: Hayman and Hill, in: Meester and Setzer, The mammals of Africa. Order Chiroptera. (Name Combination) 1971. Myonycteris torquata leptodon: Jones, J. Mamm., 52 (1): 129. Publication date: February 1971. (Name Combination) ? Myonycteris (Myonycteris) leptodon: (Name Combination) ? Myonycteris (Myonycteris) torquata leptodon: (Name Combination) African Chiroptera Report 2020 TAXONOMY: Based on the study of cytochrome b sequences, Nesi et al. (2012) concluded that West African Myonycteris torquata (Dobson, 1878) differed considerably from those from Central African. COMMON NAMES: German: Sierra-Leone Halskrausen-Flughund ETYMOLOGY OF COMMON NAME: From the Greek λετπτός meaning slender or narrow and όδους meaning tooth. CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Monadjem, 2016c). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem, 2016c). MAJOR THREATS: It may be locally threatened by habitat loss and degradation, often resulting from conversion of land to agricultural use or the harvesting of timber and firewood (Monadjem, 2016c). CONSERVATION ACTIONS: This species has been recorded within a number of protected areas and also survives outside of such areas (Monadjem, 2016c). GENERAL DISTRIBUTION: Nesi et al. (2012) indicated that the M. torquata specimens from West Africa were considerably different from those from the remaining part of the continent, but they did not include specimens from the area between the Dahomey Gap and the Niger delta. Herkt et al. (2017: Appendix S9) were not able to determine a preferential habitat type in that area for either M. torquata or M. leptodon. They were, however, able to determine that the Dahomey Gap was a pronounced break in the habitat suitability predictions of both species. Therefore, they considered the specimens on the western side of the Gap to belong to leptodon and on the eastern side to torquata. Native: Native: Côte d'Ivoire (Heim de Balsac, 1934b: 24); Ghana (Decher and Fahr, 2007: 12); Guinea (Simmons, 2005: 328; Denys et al., 2013: 281; Decher et al., 2016: 263); Liberia (Monadjem and Fahr, 2007: 50); Sierra Leone (Simmons, 2005: 328); Togo. 145 HABITAT: In the Mount Nimba area, Monadjem et al. (2016y: 364) recorded this species from a wide range of forested habitats on both Liberian and Guinean sides, between 450 m and 1,350 m. POPULATION: Structure and Density:- It is a locally abundant species that may constitute the most commonly captured fruit bat at certain localities such as at Tai National Park in Ivory Coast, and sites in Liberia (Monadjem and Fahr, 2007). Trend:- 2016: Unknown (Monadjem, 2016c). REPRODUCTION AND ONTOGENY: Weber et al. (2019: 20) captured one lactating female on 7 April in Sierra Leone. PARASITES: HAEMOSPORIDA Schaer et al. (2013a: "2") report the presence of hemosporidian parasites of the genus Hepatocystis in 11 out of 63 investigated bats. VIRUSES: Filoviridae - Filo viruses Ebolavirus Nesi et al. (2012: 126) suggest that the high nucleotide distance between Ebola virus Côte d’Ivoire and Ebola virus Zaire can be correlated with the Plio/Pleistocene divergence between their putative reservoir host species, i.e., Myonycteris leptodon and Myonycteris torquata, respectively. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Côte d'Ivoire, Ghana, Guinea, Liberia, Sierra Leone, Togo. Figure 36. Distribution of Myonycteris leptodon 146 ISSN 1990-6471 Myonycteris relicta Bergmans, 1980 *1980. Myonycteris relicta Bergmans, Zool. Meded. RMNH, Leiden, 55 (14): 171, 173, figs 1 - 10. Publication date: 4 March 1980. Type locality: Kenya: Coast province: Kwale district: Shimba Hills, Lukore area: Mukanda River [ca. 04 13 S 39 25 E] [Goto Description]. Holotype: RMNH MAM.27909: ad ♂. Collected by: Robert N. Kyongo; collection date: 30 July 1978. See Bergmans (1997: 56). Paratype: ZMB 54936: ad ♂, skull and alcoholic. Collection date: January 1900. See Turni and Kock (2008). Paratype: ZMB 54937: ad ♀, skull and alcoholic. Collection date: January 1900. See Turni and Kock (2008). (Current Combination) ? Myonycteris (Myonycteris) relicta: (Name Combination) TAXONOMY: Bronner et al. (2003) state that Myonycteris relicta was described by Bergmans (1980) based on the re-identification of a specimen from the Nguru Mountains, and two specimens from the Usambara Mountains, in Tanzania. Peterson et al. (1995: 48) regarded M. relicta as a species of Rousettus, in contrast to Koopman (1982), Corbet and Hill (1980), Koopman (1993a: 143), Bergmans (1997) and Simmons (2005: 328) who retained this taxon in Myonycteris. Assessment History Global 2016: LC ver 3.1 (2001) (Taylor, 2016e). 2008: VU A4c ver 3.1 (2001) (Mickleburgh et al., 2008ao; IUCN, 2009). 2004: VU A4c ver 3.1 (2001) (Mickleburgh et al., 2004bd; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). 1994: VU (Groombridge, 1994). COMMON NAMES: Chinese: 孤 领 果 蝠 . Czech: kaloň východoafrický. English: East African Collared Fruit-bat, Relict Collared Fruit-bat, East African Little Collared Fruit Bat, Bergmans's Small Fruit Bat, Bergmans's Collared Fruit Bat. French: Myonyctère d'Afrique orientale, Chauve-souris frugivore à collier d'Afrique orientale, Roussette du Kenya. German: Reliktärer HalskrausenFlughund. MAJOR THREATS: Although the natural history of this species is little known, there is considerable ongoing habitat loss within the species range (through logging, harvesting of firewood and conversion of land to agricultural use) (Mickleburgh et al., 2008ao; IUCN, 2009; Taylor, 2016e). ETYMOLOGY OF COMMON NAME: First described by Wim Bergmans after examining and reclassifying specimens previously ascribed to Rousettus (Lissonycteris) angolensis and following his discovery of the type specimen from the Mukonda River in Kenya (see Taylor, 2005). CONSERVATION STATUS: Global Justification Although lowland forests in Zimbabwe, Mozambique, Kenya and Tanzania are declining due to logging, fuelwood extraction and conversion to agriculture, it is impossible to provide evidence that the decline triggers levels for any Threatened or Near Threatened category (Taylor, 2016e). Further sampling (such as the survey of Monadjem et al. (2010c) in Mozambique) may reveal more populations of this species indicating that the species is not as rare as previously thought (Taylor, 2016e). Regional None known. CONSERVATION ACTIONS: Mickleburgh et al. (2008ao) [in IUCN (2009)] report that there appear to be no direct conservation measures in place for this species. In Zimbabwe it has been recorded from the Haroni and Rusitu protected areas, but the forest has still been declining at these localities over the past three decades. Further surveys are needed to better determine the full extent of this species range. There is a need to conserve remaining lowland forest throughout much of East Africa. GENERAL DISTRIBUTION: Bergmans (1997) reports that the species is known from the Shimba Hills in southeast Kenya, the Usambara and Nguru Mountains in Tanzania. Several other unpublished localities in Tanzania and more recently from eastern Zimbabwe and Mozambique. A female specimen captured in Haroni Forest (Zimbabwe) in 1973 and originally identified incorrectly as Rousettus (=Lissonycteris) angolensis, was re-identified as Myonycteris relicta (Cotterill, 1995; Bergmans, 1997). African Chiroptera Report 2020 Native: Kenya (Simmons, 2005: 328); Mozambique (Simmons, 2005: 328;: 554; Monadjem et al., 2010c: 376); Tanzania (Simmons, 2005: 328; Stanley and Goodman, 2011: 40); Zimbabwe (Simmons, 2005: 328; Monadjem et al., 2010d: 554). HABITAT: All localities known with some accuracy are in East African Zanzibar-Inhambane coastal mosaic: forest patches or in undifferentiated Afromontane vegetation bordering on East African ZanzibarInhambane coastal mosaic (Bergmans, 1997). The holotype specimen was caught over the Mukanda River, bordered by big thorn trees and fig trees, in the Shimba Hills, which are covered with 147 a mosaic of open country and forest patches (Bergmans, 1997). POPULATION: Structure and Density:-There is little known about the population of this species. Nearly all collections have been of single specimens caught in mist nets (Mickleburgh et al., 2008ao; IUCN, 2009; Taylor, 2016e). Trend:- 2016: Decreasing (Taylor, 2016e). 2008: Decreasing (Mickleburgh et al., 2008ao; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, Malawi, Mozambique, Tanzania, Zimbabwe. Myonycteris torquata (Dobson, 1878) 1870. Cynopterus collaris Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 123. Type locality: Angola: "Angola" [Goto Description]. Holotype: BMNH 1843.9.27.2:. - Comments: Not of Kolenati, 1860 [=Rousettus leachi] not of Geoffroy. Cat. Monkeys, Lemurs, Fruit-eating Bats Brit. Mus., p. 123. Type locality "W Africa, Lower Congo; Angola" see Allen (1939a: 58). Andersen (1912b: 581) mentions "The type was obtained 'near Congo' by one Mr. Currer, from whom it came into the hands of Dr. (afterwards Sir) J. Richardson, who presented it to the British Museum. Gray's statement that the specimen is 'young' is incorrect; his quotation of 'Gray, List Mamm. B. M. (1843),' where the specimen is stated to have been mentioned under the name Xantharpyia collaris, does not refer to the printed text of that book, but to a hand-written addition by Gray in his own copy of the book.". *1878. Cynonycteris torquata Dobson, Catalogue of the Chiroptera of the collection of the British Museum, xxxvii, 71, 76, pl. 5, fig. 1. Publication date: June 1878. Type locality: Angola: N Angola: Lower Cuanza region: Golungo Alto [Goto Description]. Lectotype: BMNH 1866.1.20.4: ad ♂, skull and alcoholic. Collected by: F. Welwitsch; collection date: 1853 - 1860. See Bergmans (1997: 49). - Comments: Grubb et al. (1998: 68) indicate "definite restriction of the type locality is not possible at present (Bergmans, 1976: 210 - 211)". Bergmans (1997: 49) indicates that Bergmans (1976) restricted the type locality to "Lower Cuanza Region" and the area was restricted to Golungo Alto by Crawford-Cabral (1989). 2014. Myonycteris tarquata: Ndara R., IJISR, 12 (1): 253. Publication date: November 2014. (Lapsus) ? Myonycteris (Myonycteris) torquata: (Name Combination) ? Myonycteris torquata: (Name Combination, Current Combination) ? Myopterus torquata: (Name Combination, Lapsus) TAXONOMY: See Hayman and Hill (1971), Peterson et al. (1995), Bergmans (1976; 1997) and Simmons (2005: 328). Subspecies follow Koopman (1994), but see Bergmans (1997). In their study on cytochrome b sequences, Nesi et al. (2012) concluded that West African Myonycteris torquata (Dobson, 1878) differed considerably from the specimens from Central African, which resulted in the recognition of M. leptodon as a separate species. COMMON NAMES: Castilian (Spain): Zorra Voladora de Collar. Czech: kaloň límcový. English: Little Collared Fruit-bat, Little Collared Fruit Bat. French: Petit Myonyctère, Petite Chauve-souris frugivore à collier. German: Kleiner Halskrausen-Flughund, Kleiner Halsbandflughund. CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category 148 ISSN 1990-6471 (Mickleburgh et al., 2008bb; IUCN, 2009; Bakwo Fils and Kaleme, 2016b). But due to the taxonomic changes, it is possible that in the near future, this listing might need to be be re-considered in light of results of inventories within the range of the species (Bakwo Fils and Kaleme, 2016b). Assessment History Global 2016: LC ver 3.1 (2001) (Bakwo Fils and Kaleme, 2016b). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bb; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bz; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There are generally no major threats to this species. It is threatened in parts of its range by severe habitat loss, presumably through general logging operations and the conversion of land to agricultural use (Mickleburgh et al., 2008bb; IUCN, 2009; Bakwo Fils and Kaleme, 2016b). CONSERVATION ACTIONS: The species has been recorded in the Dja Biosphere Reserve in Cameroon (Bakwo Fils, 2009) and it seems probable that it is present Central African protected areas (Mickleburgh et al., 2008bb). No direct conservation measures are currently needed for this species as a whole (Mickleburgh et al., 2008bb; Bakwo Fils and Kaleme, 2016b). The reduction of the home range may also influence its conservation status. Even within central Africa, the occurrence is reduced to some suitable habitats that are mostly found in protected areas. Monitoring of populations is important, although it will be challenging in this area. Apart from larger mammals and some taxa such as birds that are mostly used in monitoring programs, bats are less considered in monitoring strategies and even for funding of some activities (Bakwo Fils and Kaleme, 2016b). Monitoring populations is critical to provide baseline information on population trends. In many countries, only tourist appealing mammals or keystone species are more on top for conservation efforts forgetting the ecosystem services of the forgotten ones. Bakwo Fils and Kaleme (2016b) also recommend some studies such as reproduction and ecology of the species in order to provide information on how populations can grow or decline in a time span. GENERAL DISTRIBUTION: Myonycteris torquata used to be considered widespread in West Africa and Central Africa, ranging into western East Africa, but since the recognition of M. leptodon as a separate species, its distribution range starts from Nigeria and Cameroon into Equatorial Guinea (Rio Muni and Bioko), Gabon, Congo, Central African Republic, Democratic Republic of the Congo, Angola, extreme northwestern Zambia, Rwanda, Uganda and possibly southern Sudan. It is a lowland species found up to 800 m asl. Nesi et al. (2012) did not include specimens from the area between the Dahomey Gap and the Niger delta, and Herkt et al. (2017: Appendix S9) were not able to determine a preferential habitat type in that area for either M. torquata or M. leptodon. However, they were able to determine that the Dahomey Gap was a pronounced break in the habitat suitability predictions of both species. Therefore, they considered the specimens on the western side of the Gap to belong to leptodon and on the eastern side to torquata. Native: Angola (Bergmans, 1997; Simmons, 2005: 328; Monadjem et al., 2010d: 554); Cameroon; Central African Republic (Morvan et al., 1999: 1195); Congo (King and Dallimer, 2010: 65); Congo (The Democratic Republic of the) (Bergmans, 1997; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 554); Ethiopia (Lavrenchenko et al., 2004b: 136); Equatorial Guinea (Bioko) (Fahr and Ebigbo, 2003: 128; Simmons, 2005: 328); Gabon; Nigeria (CapoChichi et al., 2004: 161); Rwanda; Uganda (Kityo and Kerbis, 1996: 59; Simmons, 2005: 328); Zambia (Bergmans, 1997; Simmons, 2005: 328; Monadjem et al., 2010d: 554). Presence uncertain: Sudan. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Haiduk et al., 1980; 1981: 223) reported 2n = 36, FN = 66, BA = 32, a subtelocentric X chromosome and an acrocentric Y chromosome for specimens from Cameroon. Identical karyotypes were reported by Primus et al. (2006) for specimens from Gabon. Protein / allozyme - Unknown. DIET: In Cameroon, Grünmeier (1993: 35) found the following pollen in the gut and faeces: Mucuna flagellipes Hook.f. (Fabaceae), Anthocleista sp. (Gentianaceae), Kigelia africana (Lam.) Benth. (Bignoniaceae), Elaeis cf. guineensis Jacq. (Arecaceae), Musa sp. (Musaceae), Bombacaceae indet., and Mimosaceae indet. African Chiroptera Report 2020 In Congo, Kipalu (2009: 16) reported M. torquata visiting/consuming fruits of Chlorophora excelsa (African teak) and Ficus mucuso. Gembu Tungaluna (2012: 102 - 104) reported the following fruits being eaten in the Kisangani area (DRC) during the various seasons: First heavy rain season (September - November): Uapaca guineensis Müll.Arg., 1864 (Sugar plum, red cedar, false mahogany, rikio) (19%), Ficus leprieuri Miq., 1867 (17%), Carica papaya L. (papaya) (11%), Musanga cecropioides R.Br. & Tedlie (African corkwood tree, umbrella tree) (11%), Pseudospondias microcarpa (A. Rich.) Engl. (8%), Persea americana Mill., 1786 (avocado) (8%), Dacryoides edulis H.J. Lam (safou, African pear, bush pear, Nsafu, bush butter tree, butterfruit) (8%), Ficus wildemanniana Warb. (6%), Ficus vallis-choudae Delile (6%), Ficus mucoso Welw. (3%), Spondias cytherea Sonnerat, 1782 (Ambarella, otaheite apple, great hog plum) (3%). First mild rain/dry season (December - February): Elaeis guineensis Jacq. (African oil palm, macawfat) (50%), Myrianthus arboreus P.Beauvois, 1805 (giant yellow mulberry, bush pineapple, corkwoord) (50%). Second heavy rain season (March - May): Pseudospondias microcarpa (17%), Musanga cecropioides (14%), Spondias cytherea (9%), Ficus leprieuri (8%), Ficus sp1 (8%), Uapaca guineensis (8%), Annonidium mannii (Oliv.) Engl. & Diels (Junglesop) (6%), Ficus vallis-choudae (6%), Ficus wildemanniana (3%), Ficus sp2 (3%), Dacryoides edulis (3%), Elaeis guineensis (3%), Parinari excelsa Sabine (Guinea plum) (3%), Musa sp. (3%) Second mild rain/dry season (June - August): Ficus leprieuri (34%), Myrianthus arboreus (33%), Annonidium mannii (Junglesop) (33%). POPULATION: Structure and Density:- Although the species is infrequently caught in ground level nets, it is commonly caught in canopy nets. In Tai National Park (Côte d'Ivoire) its one of the most common fruit bats (Mickleburgh et al., 2008bb; IUCN, 2009). Trend:- 2008: Stable (Mickleburgh et al., 2008bb; IUCN, 2009). REPRODUCTION AND ONTOGENY: In Congo, Bergmans (1979a: 169) caught a female with a large foetus on 25 November, and a male with large, but not active testes on 28 November. 149 PARASITES: BACTERIA Gram-negative bacterium - Of the 40 bats tested by Nowak et al. (2017: 6) in the Republic of Congo, 22 tested positive for E. coli. One isolate was identified as E. albertii. HAEMOSPORIDA Lutz et al. (2016: 9) examined 7 M. torquata specimens from East Africa and found four of them infected with Hepatocystis sp. Perkins and Schaer (2016: Suppl.) mention the presence of Hepatocystis perronae Landau and Adam, 1971 in bats from the Republic of Congo and the Central African Republic (see also Adam, 1973: 4; Miltgen et al., 1977: 595). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999 for antibodies to SARS-CoV in sera of seven individuals from Bandundu Province, DRC, one specimen tested positive (1/7, 14.3 %). Filoviridae - Filo viruses Ebolavirus Pourrut et al. (2007) showed that there is a homogeneous ZEBOV [Zaire Ebolavirus] infection in the wild populations of M. torquata in Gabon and the Democratic Republic of the Congo. Four out of 58 bats had immunoglobulin G (IgG) specific for Ebola in Gabon during a test performed by Leroy et al. (2005: 575). Bausch and Schwarz (2014: 1) and Pigott et al. (2014: 9) indicate that M. torquata is one of the leading candidates for introducing Ebola in Guinea (together with Epomops franqueti and Hypsignathus monstrosus). While Vogel (2014: 140) names M. torquata the leading suspect for the Ebola outbreak in Guinea in the beginnig of 2014. However, no proof has been provided that this was indeed the case. Marburgvirus Towner et al. (2007) tested 264 individuals from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and 55 individuals for antiMarburg virus IgG antibodies by ELISA; no positives were found. Nairoviridae Orthonairovirus Only three of the 100 specimens from Congo tested by Müller et al. (2016: 3) were positive for Crimean Congo hemorrhagic fever virus (CCHFV). Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that three of the 111 specimens (2.7 %) they 150 ISSN 1990-6471 examined from Gabon, Congo and the Democratic Republic of the Congo tested positive for Henipavirus. Rhabdoviridae Kalemba et al. (2017: 409) reported that out of 6 bats examined from the DRC, two tested positive for Lagos Bat Virus neutralizing antibodies. peoples, fruit bats are mostly used as food but microbats are not eaten by many tribes. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Equatorial Guinea, Gabon, Ghana, Nigeria, Uganda, Zambia. In their overview table, Maganga et al. (2014a: 8) reported that the following viruses were already found on M. torquata: Zaire Ebola virus (ZEBOV), Coronavirus (SARS-CoV), Henipavirus. UTILISATION: Contrary to what was reported in the Bats as Bushmeat review (Mickleburgh et al., 2009), Bakwo Fils and Kaleme (2016b) observations in the field suggest this species is affected by hunting. Particularly notable is hunting by pygmy tribes, who are known to eat even what other people do not use as food including fruit bats (Bakwo Fils et al. pers. comm.). There are exceptions, in which some species that are prohibited by traditions or cause some problems in some tribes. However, for most of the local Figure 37. Distribution of Myonycteris torquata Myonycteris torquata torquata (Dobson, 1878) *1878. Cynonycteris torquata Dobson, Catalogue of the Chiroptera of the collection of the British Museum, xxxvii, 71, 76, pl. 5, fig. 1. Publication date: June 1878. Type locality: Angola: N Angola: Lower Cuanza region: Golungo Alto [Goto Description]. - Comments: Grubb et al. (1998: 68) indicate "definite restriction of the type locality is not possible at present (Bergmans, 1976: 210 - 211)". Bergmans (1997: 49) indicates that Bergmans (1976) restricted the type locality to "Lower Cuanza Region" and the area was restricted to Golungo Alto by Crawford-Cabral (1989). 1971. M[yonycteris (Myonycteris)] t[orquata] torquata: Hayman and Hill, in: Meester and Setzer, The mammals of Africa. Order Chiroptera, 12. (Name Combination) ? Myonycteris (Myonycteris) torquata torquata: (Name Combination) ? Myonycteris torquata torquata: (Name Combination) VIRUSES: Nesi et al. (2012: 126) suggest that the high nucleotide distance between Ebola virus Côte d’Ivoire and Ebola virus Zaire can be correlated with the Plio/Pleistocene divergence between their putative reservoir host species, i.e., Myonycteris torquata leptodon and Myonycteris torquata torquata, respectively. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon. Myonycteris torquata wroughtoni K. Andersen, 1908 *1908. Myonycteris wroughtoni K. Andersen, Ann. Mag. nat. Hist., ser. 8, 2 (11): 450. Publication date: 1 November 1908. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Likandi river [=Litaki river] [03 20 N 26 10 E] [Goto Description]. Holotype: BMNH 1907.7.8.25: ad ♂, skin and skull. Collected by: The Alexander-Gosling Expedition; collection date: 18 April 1906. See Andersen (1908c: 450); Bergmans (1997: 50). Paratype: BMNH 1907.7.8.26: ♂. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. - Etymology: In honour of Mr. Robert African Chiroptera Report 2020 1971. 1971. ? 151 Charles Wroughton, who assisted in working out the mammals collected during the Alexander-Gosling Expedition: see Andersen (1908c: 450). M[yonycteris (Myonycteris] t[torquata] wroughtoni: Hayman and Hill, in: Meester and Setzer, The mammals of Africa. Order Chiroptera, 13. (Name Combination) Myonycteris torquata wroughtoni: Jones, J. Mamm., 52 (1): 129. Publication date: February 1971. (Name Combination, Current Combination) Myonycteris (Myonycteris) torquata wroughtoni: (Name Combination, Current Combination) DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the). TRIBE Plerotini Bergmans, 1997 *1997. Plerotini Bergmans, Beaufortia, 47 (2): 69. Publication date: 20 June 1997. TAXONOMY: Includes only the genus Plerotes K. Andersen, 1910 (Almeida et al., 2020: 11). Genus Plerotes K. Andersen, 1910 *1910. Plerotes K. Andersen, Ann. Mag. nat. Hist., ser. 8, 5 (25): 97. Publication date: 1 January 1910. - Comments: Type species: Epomophorus anchietæ Seabra, 1900. - Etymology: From the Greek "πλεροτες", meaning one who renders full or complete. (Current Combination) 2014. Plerote: Ndara R., IJISR, 12 (1): 253. Publication date: November 2014. (Lapsus) TAXONOMY: Revised by Bergmans (1989). (2005: 333). See Simmons Included in the Plerotini tribe by Bergmans (1997: 69) and Almeida et al. (2016: 83), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter. Hassanin et al. (2020: 5) include it in the Epomophorini tribe and Plerotina subtribe. Currently (Simmons and Cirranello, 2020) recognized species of Plerotes: anchietae (Seabra, 1900). COMMON NAMES: Czech: pyložraví kaloni. English: Plerote Fruitbats, Plerotes Fruit-bats. French: Plérotes. German: Breitgesicht-Flughunde. Plerotes anchietae (Seabra, 1900) *1900. Epomophorus anchietæ Seabra, J. Sci. mat. phys. nat., ser. 2, 6: 116. Publication date: August 1900. Type locality: Angola: Benguela district: East of Hanha: Galanga [12 04 S 15 09 E]. Holotype: MLZA 481a: ♀, mounted skin and skull. Collected by: José Alberto de Oliveira Anchieta. Presented/Donated by: ?: Collector Unknown. Neotype: CZL 3159026: imm ♀. Collected by: Jaime V. Santos; collection date: 8 December 1959. Presented/Donated by: ?: Collector Unknown. Designated by Bergmans (1989: 144). Comments: Andersen (1912b: 487) mentions: "Type, in the Lisbon Museum, an adult female, mounted, skull extracted, obtained by Sr. Anchieta.". 2014. Plerote anchetae: Ndara R., IJISR, 12 (1): 253. Publication date: November 2014. (Lapsus) ? Epomophorus anchietae: (Name Combination) ? Epomophorus n. sp.: ? Nanonycteris veldkampii: - Comments: Not of Jentink, 1888. (Lapsus) ? Plerotes ancheilae: (Lapsus) ? Plerotes anchietae: (Name Combination, Current Combination) ? Plerotes anchietai: (Lapsus) 152 ISSN 1990-6471 TAXONOMY: Koopman (1993a: 145) mentions this name as anchietai, whereas Allen (1939a: 59) and Kock (1969a: 114) use anchietae. See Simmons (2005: 333). Almeida et al. (2016: 82) found that the only adult male specimen (SMF 85744 from Malawi) showed some interesting morphological features (i.e. a ruff of enlarged, clustered glandular hairs) that link Plerotes to the myonycterines (Megaloglossus and Myonycteris). COMMON NAMES: Czech: kaloň pyložravý. English: Anchieta's Plerote Fruit-bat, Benguela Fruit-bat, D'Anchieta's Fruit Bat, Anchieta's Fruit Bat. French: Plérote d'Anchieta, Chauve-souris frugivore d'Anchieta. German: Anchietas Breitgesicht-Flughund. Kiluba (DRC): Mulima. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Mickleburgh et al., 2008br; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008br; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004ch; IUCN, 2004). 2000: DD (Hilton-Taylor, 2000). 1996: VU (Baillie and Groombridge, 1996). 1994: Rare (Groombridge, 1994). Regional None known. MAJOR THREATS: Within the Angolan and Malawian parts of the species range it may be threatened by habitat destruction resulting from agricultural development, collection of timber and firewood, and setting of fires by poachers (Mickleburgh et al., 2008br; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008br) [in IUCN (2009)] report that although there appear to be no direct conservation measures in place, it has been recorded from a number of protected areas in Malawi and the Democratic Republic of the Congo. Further field surveys are needed to better determine the current status of this poorly known species. GENERAL DISTRIBUTION: Plerotes anchietae is largely known from scattered records in Central Africa. Most reports of this species are from the 1950's in Angola, the Democratic Republic of the Congo, Malawi and Zambia. A single specimen was collected in Malawi in 1997. It has been recorded between 1,000 and 2,000 m asl (Bergmans, 1989). Native: Angola (Bergmans, 1989; Monadjem et al., 2010d: 555; Taylor et al., 2018b: 60); Congo (The Democratic Republic of the) (Hayman et al., 1966; Bergmans, 1989; Monadjem et al., 2010d: 555); Malawi (Kock et al., 1998a; Monadjem et al., 2010d: 555); Zambia (Bergmans, 1989; Monadjem et al., 2010d: 555). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Harrison (1960c: 68) described the colour of the hairs on the dorsal surface of a subadult male from Zambia as "terminally a uniform light brown (nearest to Suntan Merida+Mayfair Tan-, Plates 13 and 7B, Maerz and Paul, 1950), becoming a little more rufous on the tibiae. The bases of the hairs are paler, buffy coloured". On the ventral surface, the pelage is "pale greyish buff, without any indication of a darker neck band". DENTAL FORMULA: The dental formula is i 1/2 c 1/1 p 3/3 m 1/3. Harrison (1960c: 69) also indicates that almost all known specimens of the genus Plerotes have some difference in the dental formula. FUNCTIONAL MORPHOLOGY: Harrison (1960c: 69) indicates that the tongue is 20 mm long (sub-equal to the mandibular length) and 9.5 mm wide at the base of the epiglottis. The dorsal tip and anterior quarter is covered with filiform papillae. POPULATION: Structure and Density:- It appears to be a rare, or at least a rarely recorded, species (Mickleburgh et al., 2008br; IUCN, 2009). Trend:- 2008: Unknown (Mickleburgh et al., 2008br; IUCN, 2009). VIRUSES: Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that single specimen they examined from Gabon did not test positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. African Chiroptera Report 2020 153 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Congo (Democratic Republic of the), Malawi, Zambia. Figure 38. Distribution of Plerotes anchietae TRIBE Rousettini Andersen, 1912 *1912. Rousettini K. Andersen, Catalogue of the Chiroptera in the British Museum. Publication date: 23 March 1912. TAXONOMY: Includes only the genus Rousettus Gray, 1821 (Simmons and Cirranello, 2020: 11). Genus Rousettus Gray, 1821 *1821. Rousettus Gray, London Med. Repos., 15: 299. Publication date: 1 April 1921. Comments: Type species: Pteropus Egyptiacus E. Geoffroy Saint-Hilaire, 1810. Not Pteropus aegyptiacus Geoffroy, 1818 (see Corbet and Hill, 1992: 67). - Etymology: From the French "rousette" for the rosette of glandular hairs on the neck of males (see Flannery, 1995a: 304) or from the French "rousse" for reddish or reddish brown, referring to the fur colour of the earliest specimens taken in Egypt (see Kwiecinski and Griffiths, 1999: 7). (Current Combination) 1829. Cercopteropus Burnett, Quart. J. Sci., Lit. and Art, pt. 1, art. 5, p. 269. Publication date: April - June 1829. - Comments: Type species: Pteropus aegyptiacus E. Geoffroy SaintHilaire, 1818 (Designated by Andersen, 1912b: 23). Burnett (1829: 269) did mention both Ægyptiacus and Amplexicaud [for amplexicaudata] as being included in his genus. Etymology: From the Greek "κέρκος", meaning tail and "Pteropus" (see Palmer, 1904: 172). 1843. Xantharpyia Gray, Catalogue of the species of mammals of the British Museum, xix, 37. Publication date: 13 May 1843. - Comments: Type species: Pteropus amplexicaudatus E. Geoffroy Saint-Hilaire, 1810. - Etymology: From the Greek "ξανθός", meaning yellow and Harpyia from the characteristic colour (see Palmer, 1904: 710). 1844. Eleutherura Gray, Mammalia, in Voyage "Sulphur", Zool., 1: 29. - Comments: Type species: Pteropus leachii A. Smith, 1829 (=Pteropus aegyptiacus E. Geoffroy Saint-Hilaire, 1810). Andersen (1912b: 24) mentions Pteropus hottentotus Temminck, 1832 as type species, which is a synonym of P. leachi A. Smith 1829. Preoccupied by Eleutherura Gray, 1843; nom. nud. (Andersen, 1912b: 16; Meester et al., 1986: 28; Pavlinov et al., 1995: 61). Allen (1939a: 61) indicates that Gray (1843 - List. Spec. Mamm. Brit. Mus., p. xix) is a nomen nudum, but that it is redescribed in the 1844 publication. - Etymology: From the Greek "έλεύθερος", meaning free and "ούρά", meaning tail, referring to having the tail free from the interfemoral membrane (see Palmer, 1904: 255). 154 ISSN 1990-6471 1852. 1870. 1899. 1906. 1906. 1992. 1999. 2007. 2015. Cynonycteris Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 25 [Goto Description]. - Comments: Type species: Pteropus collaris Liechtenstein [not Illiger] (=Pteropus aegyptiacus E. Geoffroy Saint-Hilaire, 1810). Etymology: From the Greek "κύων" or "κυνός", meaning dog and "νυκτερις", meaning bat, probably referring to the dog-like head (see Palmer, 1904: 212). Senonycteris Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 115. - Comments: Type species: Pteropus seminudus Kelaart, 1850 (=Pteropus lechenaulti Desmarest, 1820). Described as subgenus of Xantharpyia. Mynonycteris Matschie, Die Fledermäuse des Berliner Museums für Naturkunde. 1. Lieferung. Die Megachiroptera des Berliner Museums für Naturkunde, p. 63. Roussettus (Xantharpyia): Johnston, Liberia, Volume II: 690. (Name Combination, Lapsus) Roussettus: Johnston, Liberia, Volume II: 688. (Lapsus) Rosetus: Agrawal, Das, Chakraborty, Ghose, Mandal, Chakrabo, Zoological Survey of India, State Fauna Series 3, 49. - Comments: Lapsus. See Bergmans (1994: 82). (Lapsus) Rosettus: Buffenstein, Maloney and Bronner, S. Afr. J. Zool., 34 (1): 16. (Lapsus) Rousetus: Nel and Markotter, Crit. Rev. Microbiol., 33 (4): 308. (Lapsus) Rousettes: Lee, Kulcsar, Elliott, Khiabanian, Nagle, Jones, Amman, Sanchez-Lockhart, Towner, Palacios and Rabadan, BMC Genomics, 16 (1033): 1. Publication date: 7 December 2015. (Lapsus) TAXONOMY: Meester et al. (1986) mention that the inclusion of Lissonycteris as a subgenus of Rousettus is open to question. Lawrence and Novick (1963) maintain that Lissonycteris should be regarded as a valid genus, more closely related to Myonycteris Matschie, 1899, than to Rousettus. In this, they are followed by Rosevear (1965), Ansell (1978), Corbet and Hill (1980), Almeida et al. (2011a), and Patterson and Webala (2012). Koopman (1975), on the other hand, concludes that although there is a close relationship between Lissonycteris and Myonycteris, the relationship of Lissonycteris with Rousettus is sufficiently close for it to be retained as subgenus. Koopman (1982) again treats Lissonycteris as a subgenus, as is done also by Hayman and Hill (1971). Bronner et al. (2003), following Lawrence and Novick (1963), separated Lissonycteris from Rousettus because of ethological differences concerning the use of the limbs, and the absence of echolocation in Lissonycteris. Juste B. et al. (1997) presented allozyme evidence corroborating earlier chromosome studies (Haiduk et al., 1980, 1981), DNA-hybridisation results (Kirsch et al., 1995) and cladistic analysis (Springer et al., 1995) that support the recognition of Lissonycteris as a distinct genus. Three subgenera are often recognized (Rousettus, Boneia, and Stenonycteris), although Bergmans (1994), rejected the use of subgenera for the African species (Simmons, 2005: 347). A key to the species is provided by Kwiecinski and Griffiths (1999: 1, Mammalian Species, 611). Included in the Rousettini tribe by Almeida et al. (2016: 83) and Hassanin et al. (2020: 5). Currently (Simmons and Cirranello, 2020) recognized species of the genus Rousettus: aegyptiacus (E. Geoffroy Saint-Hilaire, 1810); amplexicaudatus (E. Geoffroy Saint-Hilaire, 1810) – Cambodia, Thailand, Burma, and Laos, Peninsular Malaysia through Indonesia, Java, and Bali, Philippines; New Guinea, Bismarck Archipelago, Solomon Islands (Simmons, 2005: 347); celebensis K. Andersen, 1907- Sulawesi, Mangole, Sanana, Sangihe Islands (Indonesia) (Simmons, 2005: 347); leschenaultii (Desmarest, 1820) – Sri Lanka, Pakistan to Vietnam and southern China, Peninsular Malaysia, Sumatra, Java, Bali, and Mentawai Islands (Indonesia) (Simmons, 2005: 348); linduensis Maryanto and Yani, 2003 – central Sulawesi (Simmons, 2005: 348); madagascariensis G. Grandidier, 1929; obliviosus Kock, 1978; spinalatus Bergmans and Hill, 1980 – Sumatra, Borneo (Simmons, 2005: 348). COMMON NAMES: Czech: egyptští kaloni, noční kaloni, psi noční. English: Rousettes, Rousette Fruit-bats. French: Roussettes. German: Höhlenflughunde. Italian: Rossétti. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Horácek et al. (2013b: 1) report on two tooth fragments from the Middle Pleistocene Israeli site Qesem Cave, which they tentatively attribute to cf. Rousettus sp. If this identification is confirmed, they suggest that the current Mediterranean African Chiroptera Report 2020 population of R. aegyptiacus could be seen as a paleochoric element. However, the teeth they report on are larger than similar teeth in extant R. aegyptiacus specimens, so they might be from a different species or there might have been phenotypic rearrangements such as increased body size (as they suggest on page 13). GENERAL DISTRIBUTION: Stribna et al. (2019: 2337) indicate that the genus probably originated in Asia and that the colonisation of Africa occurred in the late Pliocene or early Pleistocene (ca. 2.6 to 1.7 mya). BIOGEOGRAPHY: Several hypotheses have been proposed about the origin of the genus Rousettus on the phylogenetic inference, and centers of diversity pinpoint a southeastern Asian origin (Juste B. et al., 1999). Three separate routes have been suggested for western expansion of pteropodids into Africa, via a middle Asian-European-Gibraltar route, a middle-Asian, Middle Eastern route, and an Indian subcontinent-western Indian Ocean, east African route (Juste B. et al., 1999). Giannini and Simmons (2003, 2005) support the Middle Eastern route with the Afrotropical clade composed of R. aegyptiacus and R. madagascariensis being the sister group to R. leschenaulti. 155 Almeida et al. (2016: 82) indicate that Rousettus expanded from Asia into Africa in the late Pliocene or early Pleistocene, where the continental R. aegyptiacus already split off in the early Pleistocene, slightly later followed by a split of the two island forms. PARASITES: BACTERIA: Di Cataldo et al. (2020: 2) examined 6 Nigerian Rousettus bats and found 2 to be infected by hemoplasma bacteria. ACARI: Trombiculidae: Stekolnikov (2018a: 50) reported Rousettus sp. to be the type host of Whartonia oweni Vercammen-Grandjean and Brennan, 1957. DIPTERA: Nycteribiidae: Tripselia blainvillii (Leach 1817) single record (Haeselbarth et al., 1966: 113). Morse et al. (2013: Suppl. 1) refer to Dipseliopoda biannulata collected from a non-specified Kenyan Rousettus specimen. This bat fly was itself infected by an Arsenophonus sp. bacteria (Szentiványi et al., 2019: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Gabon, Kenya, South Africa, Tanzania, Uganda. †Rousettus pattersoni Gunnell and Manthi, 2018 *2018. R[ousettus] pattersoni Gunnell and Manthi, J. Hum. Evol., p. "2", "3", fig. 1C-D. Publication date: 6 April 2018. Type locality: Kenya: Kanapoi Bat Site [Goto Description]. Holotype: NMK KP68719: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Holotype is a P4. - Etymology: Named in honour of Brian Patterson, who, in 1966-1967, initiated the early field work in Kanapoi (Gunnell and Manthi, 2018: "3"). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Pliocene (Gunnell and Manthi, 2018: "3"). Rousettus aegyptiacus (E. Geoffroy St.-Hilaire, 1810) *1810. Pteropus Egyptiacus E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 15: 96. Type locality: Egypt: Giza province: Giza: Great Pyramid [30 01 N 31 13 E]. Holotype: MNHN A.70: ad ♀. Presented/Donated by: ?: Collector Unknown. Allotype: MNHN A.69: ♂, mounted skin (skull not removed). Collected by: Etienne Geoffroy-Saint-Hilaire. Presented/Donated by: ?: Collector Unknown. - Comments: Type locality originally "la basse Egypte - le plafond d'une des chambres de la grande Pyramide". See Kwiecinski and Griffiths (1999: 1). Type MNHN A.70 (ad ♀, alcoholic and skull: Qumsiyeh, 1985: 13). - Etymology: The name is derived from the fact that the species was originally described from a specimen from the Great Pyramid of Giza in Egypt (see Smithers, 1983: 63). 1818. Pteropus aegyptiacus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 134, pl. 3, No 2. Holotype: MNHN A.70: ad ♀, skull and alcoholic. Collected by: Etienne Geoffroy-Saint-Hilaire. See Qumsiyeh (1985: 13). Number 167 in Rode (1941). - Comments: Emendation of egyptiacus. Unjustified according to Corbet 156 ISSN 1990-6471 1823. 1825. 1829. 1832. 1862. 1862. 1870. 1906. 1908. 1911. 1951. 1958. 1960. and Hill (1992: 67), justified according to Kock (2001c). - Etymology: From the scientific Latin masculine adjective aegyptiàcus, meaning "Egyptian", as the species was described from an Egyptian specimen (see Lanza et al., 2015: 69). (Emendation) Pteropus collaris Lichtenstein, Verzeichniss der Doubletten des Zool. Mus. K. Univ. Berlin, 3, 5. Type locality: South Africa: "Terra Caffrorum". - Comments: Preoccupied by Pteropus collaris Illiger, 1815; see Kwiecinski and Griffiths (1999: 1). Pteropus geoffroyi Temminck, Monogr. Mamm, 1 (5): 197, pl. 15, f. 14, 15. Type locality: Senegal: Senegal and north coast of Africa. - Comments: Meester et al. (1986: 29) mention that the type locality is restricted to Giza, Egypt by Koopman (1975: 361). Andersen (1912b: 32) mentions it as a replacement name for R. aegyptiacus, since "vu que I'espèce se trouve au Sénégal, et probablement sur toute la côte septentrionale d'Afrique," [= "in view of the fact that the species occurs in Senegal and probably over the the entire northern coast of Africa."]. Pteropus Leachii A. Smith, Zool. Journ., 4 (16): 433. Publication date: May 1829. Type locality: South Africa: Gardens about Cape Town [ca. 33 56 S 18 28 E] [Goto Description]. Paratype: SMF 891: juv ♂, skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Cotype: BMNH 1875.11.3.2: ad ♀, alcoholic (skull not removed). Collected by: Sir Andrew Smith. Exchanged by G.E. Dobson (see Andersen, 1912b: 28). - Etymology: In honour of the celebrated naturalist, Dr. Leach, F.R.S. (see Smith, 1829: 433). Pteropus hottentottusTemminck, in: Smuts, Enumer. Mamm. Capensium, 3. Type locality: South Africa: Cape province: originally ""circa urbem apensem" see Andersen (1912b: 28), restricted to Capt Town.: Cape Town [33 56 S 18 28 E] [Goto Description]. - Comments: Mentioned as "hottentotus" by), Simmons (2005), but as hottentottus by Allen (1939a: 62), Roberts (1951: 55), Meester et al. (1986: 29), Kwiecinski and Griffiths (1999). Andersen (1912b: 29) mentions that the type is in the RMNH, but also indicates that there are "authentic specimens" in the BMNH: 1837.4.28.33-34, 38, 40, 67 (4 ad + 1 imm; sks and skulls), from "near Cape Town" (J. Smuts), and purchased from the Leyden Museum. Xantharpyia ægyptiaca: Gerrard, Catalogue of the bones of Mammalia in the collection of the British Museum, 57. Type locality: Egypt. (Name Combination) Xantharpyia leachii: Gerrard, Catalogue of the bones of Mammalia in the collection of the British Museum, 57. Eleutherura unicolor Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 117. Type locality: Gabon: "Gabon" [Goto Description]. Holotype: BMNH 1862.8.26.1: ad ♂, skin and skull. Purchased Verreaux: see Andersen (1912b: 32). R[oussettus (Xantharpyia)] collaris: Johnston, Liberia, Volume II: 690. (Lapsus) Rousettus sjöstedti Lönnberg, Wiss. Ergebn. Swedischen Zool. Exp. u. Leitung Sjöstedt, Kilimandjaro-Meru, 1 (2): 7. Type locality: Tanzania: near Tanga: Mkulumusi caves. [Goto Description]. Holotype: NRM [unknown]: ad ♀, alcoholic (skull not removed). Collected by: Bror Yngve Sjöstedt; collection date: 4 May 1905. Kilimandjaro-Meru Expedition. See Lönnberg (1908b: 7) and Andersen (1912b: 810). Cynonycteris collaris: Mitchell, Proc. zool. Soc. Lond., 1911, II: 444. Publication date: 6 July 1911. Pteropus amphiplexicaudatus: Roberts, The mammals of South Africa. Type locality: South Africa: Cape. - Comments: Probably a lapsus. Roberts (1951: 55) attributed the name to Temminck (1827) and mentions it as "not of Geoffroy", which would probably refer to amplexicaudatus. However, Temminck (1827) did not use this erroneous spelling. (Lapsus) Rousettus aegypticus: Griffin, Novick and Kornfield, Biol. Bull., 115 (1): 107. Publication date: August 1958. (Lapsus) Rousettus aegyptiacus occidentalis Eisentraut, Bonn. zool. Beitr., 10: 231, figs 8, 13 (for 1959). Publication date: 10 January 1960. Type locality: Cameroon: N side of Mount Cameroon: Mueli [04 23 N 09 07 E, ca. 600 m]. Holotype: ZFMK MAM-1959.0450: ad ♂, skin and skull. Collected by: Prof. Dr. Martin Eisentraut; collection date: 20 February 1958; original number: 648. Paratype: ZFMK MAM-1961.0575: Collected by: Prof. Dr. Martin Eisentraut; collection date: January 1958. Lager IV [= Camp IV], near lake Koto Barombi, 120 m, N side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0576: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, African Chiroptera Report 2020 157 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0577: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0578: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0579: Collected by: Prof. Dr. Martin Eisentraut; collection date: January 1958. Lager IV [= Camp IV], near lake Koto Barombi, 120 m, N side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0580: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM1961.0581: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0582: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0583: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0584: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0585: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0586: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0587: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0588: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0589: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0590: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0591: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0592: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0593: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0594: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0595: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM1961.0596: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0597: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0598: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0599: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0600: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). 158 ISSN 1990-6471 1974. 1993. 1993. 1993. 2006. 2008. 2008. 2011. 2012. 2015. 2015. Paratype: ZFMK MAM-1961.0601: Collected by: Prof. Dr. Martin Eisentraut; collection date: January 1958. Lager IV [= Camp IV], near lake Koto Barombi, 120 m, N side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1961.0602: Collected by: Prof. Dr. Martin Eisentraut; collection date: February 1958. Lager V [= Camp V], above Mueli, 600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0208a: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0208b: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0208c: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0208d: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0208e: Collected by: Prof. Dr. Martin Eisentraut; collection date: March 1958. Lager VI [= Camp VI], Isobi, 30 m, W side Mount Cameroon (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0209a: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0209b: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM1963.0209c: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0209d: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0209e: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Paratype: ZFMK MAM-1963.0209f: Collected by: Prof. Dr. Martin Eisentraut; collection date: November 1957. Lager I [= Camp I], above Buea, 1,600 m (see Hutterer, 1984: 29; Hutterer and Peters, 2010: 10). Rousettus aegyptiacus unicolor: Bergmans, Bellier and Vissault. (Name Combination) Rousettus aegyptiacus thomensis Feiler, Haft and Widmann, Faun. Abh. Mus. Tierk. Dresden, 19 (4): 22. Publication date: 1 July 1993. Type locality: São Tomé and Principé: São Tomé Island: Generosa [ca. 800 m] [Goto Description]. Holotype: SMTD 16753: ad ♂, skin and skull. Collection date: 26 March 1991. See Feiler et al. (1993: 22 - 23). Paratype: SMTD ad ♂, skin and skull. See: 22 - 23). Paratype: SMTD ad ♀, skin and skull. See: 22 - 23). Rousettus aegyptiacus tomensis Juste and Ibáñez, Zool. J. Linn. Soc., 107: 124. Type locality: São Tomé and Principé: São Tomé Island: Bindá (Santa Catarina) [00 16 N 06 29 E]. Roussetus aegyptiacus princeps Juste and Ibáñez, Zool. J. Linn. Soc., 107: 123. Type locality: São Tomé and Principé: Principé Island: 2 km S of Santo Antonio de Principé: Roça Bela Vista [ca. 01 37 N 07 27 E]. Rousettus egyptiacus: Nogales, Rodríguez-Luengo and Marrero, Mammal Rev., 36 (1): 49. Publication date: January 2006. (Lapsus) Rousettus aegyptiacu: Selim, Nahla and Shelfeh, Tishreen Univ. J. Res. Sci. Stud. - Biol. Sci. Ser., 30 (1): 249. (Lapsus) Rousettus occidentalis: Benda, Vespertilio, 12: 109. (Name Combination) Rousettus aegyptiacusi: Hickey, Jüllig, Aitken, Loomes, Hauber and Phillips, Ageing Res. Revs, 11 (2): 249 (for 2012). Publication date: 13 December 2011. (Lapsus) R[ousettus] a[egyptiacus] egyptiacus: Hulva, Marešová, Dundarova, Bilgin, Benda, Bartonicka and Horácek, Mol. Ecol., 21 (24): 6105. Publication date: 24 October 2012. (Lapsus) Rousettus aegypticus leachii: Kaipf, Rudolphi and Meinig, NABU - Biodiversity Assessment in Kafa, Ethiopia, 7. (Lapsus) Rousettus aegypticus occidentalis: Sylla, Pourrut, Diatta, Diop, Ndiaye and Gonzalez, Afr. J. Microbio. Res., 9 (22): 1468. Publication date: 3 June 2015. (Lapsus) African Chiroptera Report 2020 2019. 2019. 2020. ? ? ? ? ? ? ? ? ? ? ? ? ? 159 Rousettus aegypti: Siya, Bazeyo, Tuhebwe, Tumwine, Ezama, Manirakiza, Kugonza and Rwego, BMC Public Health, 19 (136): 1. Publication date: 31 January 2019. (Lapsus) Rousettus aegyptiacusi: Malmlov, Bantle, Aboellail, Wagner, Campbell, Eckley, Chotiwan, Gullberg, Perera, Tjalkens and Schountz, PLoS Negl.Trop.Dis., 13 (2) e0007071: 3. Publication date: 4 February 2019. (Lapsus) Rosettus Aegiptiacus: David, Davidson, Berkowitz, Karniely, Edery, Bumbarov, Laskar and Elazari-Volcani, Vet. Med. Sci., xxx (x): xxx. Publication date: 25 February 2020. (Lapsus) Cynonycteris ægyptiaca: - Comments: see: Günther (1880: 741) as synonym of Cynonycteris collaris. (Name Combination) Pteropus ægyptiacus - Comments: Günther (1880: 741) refers to a list by Kotschy and published in Unger and Kotschy's 'Die Insel Cypern'. (Alternate Spelling) Rousettus (Rousettus) aegyptiacus leachii: (Name Combination) Rousettus (Rousettus) aegyptiacus: (Name Combination) Rousettus aegyptiacus aegyptiacus: (Name Combination) Rousettus aegyptiacus arabicus: (Name Combination) Rousettus aegyptiacus leachi: (Name Combination) Rousettus aegyptiacus leachii: (Name Combination) Rousettus aegyptiacus leacii: (Lapsus) Rousettus aegyptiacus: (Name Combination, Current Combination) Rousettus arabicus: Rousettus collaris: - Comments: . Rousettus leachi: TAXONOMY: Figure 39. Rousettus aegyptiacus, Pufuri, Kruger NP, South Africa. Andersen (1912b) recorded R. aegyptiacus, R. arabicus and R. leachii as species in the genus Rousettus. Miller (1912) and Ellerman and Morrison-Scott (1951) recognized R. arabicus as a valid species. Harrison (1964a) stated that R. aegyptiacus is represented by the subspecies R. a. aegyptiacus and R. a. arabicus. Bergmans (1994) recognised R. a. aegyptiacus, R. a. leachii, R. a. unicolor, and R. a. arabicus as valid subspecies. Corbet and Hill (1992) corrected Geoffroy SaintHilaire's spelling of Rousettus aegyptiacus to R. egyptiacus, a change endorsed by Koopman (1993a: 152) and Bergmans (1994). Kock (2001c), however, has presented a detailed and convincing case for regarding egyptiacus as an incorrect original spelling and treating aegyptiacus as the valid name (Bronner et al. (2003). Kock (1969a: 16), Qumsiyeh et al. (1992: 104) and Horácek et al. (2000) state that Geoffroy SaintHilaire (1818: 134) already corrected the spelling to aegyptiacus. Allen (1939a: 62) also indicates that it was a misprint in the 1810 publication, which was corrected in the 1818 publication. Meester et al. (1986) state that, while leachii is clearly the Southern African subspecies extending well beyond the limits of this subregion, Hayman and Hill (1971) mention that intergradation with aegyptiacus may occur in the equatorial zone. Koopman (1975: 361) indicates that in the Sudan there is a hiatus between the ranges of these two sub-species, and he remarks that if they make contact it will be in Ethiopia, north of the equator. See Simmons (2005: 347). Horácek et al. (2010: 175) indicate that the specimens from the Mediterranean area differ with more than 10 % in their mtDNA from the SubSaharan populations, which might suggest that these specimens represent a different species. Benda et al. (2012b: 1230) showed that, although two size morphotypes were found, the Palaearctic populations of R. aegyptiacus represent only one form, which is uniform in genetic traits (mitochondrial genome [nd1 and cyt b]) but plastic in metric traits. Avery and Avery (2011: 539 - 540) report that a separation between the ancestor of R. aegyptiacus and those of genera such as Myotis and Kerivoula occurred some 64 million years ago. The separation with Rhinolophus and Hipposideros occurred slightly less than 60 million years ago, and that with Pteropus some 22 million years ago. 160 ISSN 1990-6471 COMMON NAMES: Afrikaans: Egiptiese vrugtevlermuis. Albanian: Lakuriq nate egjiptian i frutave. Arabian: Khafash El Fawakeh, Khafash Masri, Khaffash, Wat-Wat. Armenian: Եգիպտական մեծաչղջիկ. Azerbaijani: Misir meyvə yarasası. Basque: Fruitu-saguzar egiptoar. Belarusian: Крылан егіпецкі. Bosnian: Egipatski letipas. Breton: Lgodenn-dall rous Egipt. Bulgarian: Египетски плодояден прилеп. Castilian (Spain): Murciélago de la fruta egípcio, Ruseta Egipicia. Catalan (Spain): Ratpenat egipci. Chewa (Malawi): Karuru. Chinese: 北 非 果 蝠 . "Congolese": Kubukubo, Papo. Croatian: Egipatski plodojedi šišmiš. Czech: Kaloň egyptskýý, Upír egypeckýý, Kaloň nilskýý. Danish: Egyptisk flyvehund. Dutch: Nijlrousette, Nijlrousettus. English: Egyptian Fruit Bat, Egyptian Rousette, Egyptian Rousette Bat, West African Rousette, Egyptian Xantharpy. Estonian: Egiptuse öötiibur. Finnish: Ronkko. French: Roussette egyptienne, Roussette d'Égypte. Frisian: Nylrûsette. Galician (Spain): Morcego exipcio. Georgian: ეგვიპტური მფრინავი ძაღლი. German: Nilflughund (Ägyptischer Höhlenflughund), Egyptischer fliegender Hund. Greek: Αιγυπτιακή φρουτονυχτερίδα. Hebrew: Atalef Perot. Hungarian: Nílusi repülőkutya. Italian: Rossétto egiziàno, Rossétto del Nìlo, Rossetta. Irish Gaelic: Ialtóg meas Éigipteach. Kinande (DRC): Mulima. Latvian: Ēģiptes augļsikspārnis. Lithuanian: Egiptinis vaisėdis. Luxembourgish: Nilfluchhond. Macedonian: Египетски овошен лилјак. Maltese: Farfett ilLejl tal-Frott ta' L-Eġittu. Montenegrin: Egipatski letipas. Norwegian: Afrikagrotteflygehund. Polish: Rudawka nilowa. Portuguese: Morcegofrugívoro-egípcio, Morcego frugivoro do Egipto. Rhaeto-Romance: Chaun sgulant da l'Egipta. Romanian: Liliacul fructivor egiptean. Russian: Крылан египетский. Serbian: Oбични пећински летипас [= Obični pećinski letipas]. Scottish Gaelic: Ialtag meas Èipheiteach. Slovak: Kaloň egyptský. Slovenian: Egipč anski leteči pes. Swedish: Nilflyghund. Turkish: Mısır Meyve Yarasası. Ukrainian: Крилан єгипетський. Welsh: Ystlum-ffrwythau yr Aifft. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Hildebrand et al. (2010: 268) recorded this species from the Kumali site in southern Ethiopia. Van Neer (1990) reported on the presence of archeozoological material from this species in a number of localities in Central Africa. CONSERVATION STATUS: Global Justification This species is broadly distributed and abundant, and is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category, hence is listed as Least Concern (LC ver 3.1 (2001)) (Benda et al., 2008a; IUCN, 2009; Korine, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Korine, 2016). 2008: LC ver 3.1 (2001) (Benda et al., 2008a; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ct; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Markotter et al., 2016). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: This species faces a number of threats, but none of them are considered a serious threat to the species at the global level. Hunted for food in some cave systems in Africa. Cave disturbance and persecution are also a problem in parts of the range. The species is considered as a pest by fruit farmers and consequently cave roosts have been fumigated and destroyed in Israel, Turkey and Cyprus, resulting in incidental killing of many insectivorous bats of the genera Rhinolophus and Myotis (Benda et al., 2008a; IUCN, 2009; Korine, 2016). The Israeli Nature Conservation Society is trying to prevent this lethal control (D. Kock, pers. comm., in Benda et al., 2008a, in IUCN, 2009), as campaigns during the 1950s led to the unintended death of 90 % of the insectivorous bats that roosted in the same caves (Makin and Mendelssohn, 1985 [in Guyton and Brook, 2014: 77]). Sherwin et al. (2012: 174) identified that R. aegyptiacus's frugivory and long-distance dispersal are major risk factors linked with climatic change. MacEwan (2016: 3) reports on R. aegyptiacus as fatalities of wind turbine blades. Monadjem et al. (2016y: 364) report that the species has disappeared from the Liberian side of Mount Nimba, possibly as a result of hunting. CONSERVATION ACTIONS: Benda et al. (2008a) [in IUCN (2009)] and Korine (2016) report on International legal obligations for protection through the Bonn Convention (Eurobats) in areas where this applies. Included African Chiroptera Report 2020 in Annex IV of the EU Habitats Directive in areas where this applies. Occurs in a number of protected areas. There is a need to enforce measures to protect this species, especially to prevent the fumigation of caves. GENERAL DISTRIBUTION: Rousettus aegyptiacus is patchily distributed across sub-Saharan Africa and North Africa; also ranges outside of Africa through south-west Asia to Iran, Pakistan and India; also on Cyprus. In the Western Palaearctic it occurs as two forms: R. a. arabicus (Iran, southern Arabia, Pakistan), R. a. aegyptiacus (rest of range). For more details on the Palaearctic distribution, see Benda et al. (2011a). In sub-Saharan Africa occurs as four forms: subspecies leachi is found in SW Ethiopia, S Sudan, E and S DR Congo, Uganda, Kenya, Tanzania (including the islands of Pemba and Unguja [see O'Brien (2011: 286]), Zambia, Malawi, Zimbabwe, Mozambique, and the extreme E and south of South Africa, including Swaziland and Lesotho; subspecies princeps is endemic to Principe Island in the Gulf of Guinea; subspecies tomensis is endemic to São Tomé; and subspecies unicolor is found from Senegal and the Gambia east to Liberia, Côte d'Ivoire, Ghana, Togo, south to northern central Nigeria and W Cameroon and Bioko Island, then south from Gabon and Congo to western DRC and W Angola. Elevational range: from sea level to 4,000 m asl, although Dongmo et al. (2020: 51) only reported it in Cameroon from elevations below 1,250 m. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 692,896 km 2. In the RSA, its distribution is best predicted by geology (Babiker Salata, 2012: 50). Nogales et al. (2006: 52) report on a population of R. aegyptiacus, which escaped in 2000 from two zoos on Tenerife, where they were kept from around 1992. This population was eradicated in 2009 (see Simons et al., 2014: 2091). Native: Angola (Crawford-Cabral, 1989; Bergmans, 1994; Monadjem et al., 2010d: 555); Burkina Faso (Kangoyé et al., 2012: 6024; 2015a: 607); Burundi; Cameroon; Central African Republic (Morvan et al., 1999: 1195); Congo; Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 555); Côte d'Ivoire; Cyprus; Egypt (Benda et al., 2008f: 1); Equatorial Guinea (Fahr and Ebigbo, 2003: 128); Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 135; Benda et al., 2020: 2582); Gabon; Gambia; Ghana; Greece (Strachinis et al., 2018: 611); Guinea (Denys et al., 2013: 281; Decher et al., 2016: 263); Guinea- 161 Bissau (Rainho and Ranco, 2001: 33); India (Rainho et al., 2010a), Iran, Islamic Republic of; Israel; Jordan; Kenya; Lebanon; Lesotho; Liberia (Monadjem and Fahr, 2007: 50); Libyan Arab Jamahiriya; Macaronesian islands (Tenerife island (Fajardo, 2002; Rodriguez-Luengo and Garacia Casanova, 2002; Trujillo, 2003; Nogales et al., 2006; Masseti, 2010)); Malawi (Happold et al., 1988; Ansell and Dowsett, 1988: 28; Bergmans, 1994; Monadjem et al., 2010d: 555); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 555; Monadjem et al., 2010c: 377); Nigeria (Capo-Chichi et al., 2004: 161); Oman; Pakistan; Palestinian Territory, Occupied; Rwanda; São Tomé and Principé (Juste and Ibáñez, 1993b: 906; Rainho et al., 2010a: 19 as R. a. tomensis); Saudi Arabia; Senegal; Sierra Leone; South Africa (Monadjem et al., 2010d: 555); Sudan; Syrian Arab Republic; Tanzania (Stanley and Goodman, 2011: 40); Togo; Turkey; Uganda (Kityo and Kerbis, 1996: 59); United Arab Emirates (Anonymus, 2008: 38); Yemen (Benda et al., 2011b: 26) ; Zambia (Ansell, 1969; Bergmans, 1994; Cotterill, 2004a: 260; Monadjem et al., 2010d: 555); Zimbabwe (Cotterill, 2004a: 260; Monadjem et al., 2010d: 555). BIOGEOGRAPHY: Horácek et al. (2013a: 71) suggest a few hypotheses about the presence of R. aegyptiacus in the eastern Mediterranean. These populations might be the result of immigration from subSaharan Africa or southern Arabia, although they also indicate that the Mediterranean form might have been the source population for the spread of R. aegyptiacus towards Africa. Hulva et al. (2013: 74) indicate that the colonisation of northeastern Africa and the Middle East was connected with the spread of agricultural plants, and might therefore have an anthropogenic component. DETAILED MORPHOLOGY: Baculum - See Albayrak et al. (2008) for samples from Turkey. Brain - Chawana et al. (2013: 160) reported the average brain mass for two specimens to be 2.01 g. Maseko et al. (2007) used immunohistochemical techniques to describe the nuclear parcellation and neuronal morphology of the cholinergic, catecholaminergic and serotonergic systems within the brain of three adult female R. aegyptiacus, captured from a cave adjacent to Legalametse Nature Reserve, Limpopo Province, South Africa. Naumann et al. (2015) investigated the medial entorhinal cortex architecture in layer 2 of the bat's brain. 162 ISSN 1990-6471 The cytoarchitecture of R. aegyptiacus's brain was investigated by Rodenas-Cuadrado et al. (2018: 1237) to determine the expression patterns of FoxP2, FoxP1, and Cntnap2, which are implicated in human speech and language phenotypes. Schönenberger and Lane (1971: 653) studied the peripherical nervous system in the wings of R. aegyptiacus. The nerve bundle consists of a number of fasciles and each of these contains both myelinated and non-myelinated fibres. They also found that some of these nerves react positively to an histochimical test for catecholamine. The accompanying veins and arteries were studied by Lane et al. (1971). Tongue - The arrangement of mechanical filiform papillae and gustatory papillae is regarded as being useful for the efficient uptake of semiliquid food and also an adaptation to a fruit diet (Jackowiak et al., 2009). Abumandour and ElBakary (2013: 230) report four types of lingual papillae: two mechanical (filiform and conical)l and two gustatory (fungiform and circumvallate) in which the shape, size, number, distribution, nomenclature and orientation of these lingual papillae are region specific according and in relation to feeding habits and food types. Abumandour (2014: 1407) distinguishes four types of lingual papillae, of which the filiform papillae can be subdivided into six subtypes: three on the anterior part (small, conical and giant), two on the middle part (cornflower and leaf-like papillae), and rosette shape filiform papillae on the posterior part. El-Mansi et al. (2019: 24, 29) subdivided the mechanical papillae in conicals (small and large) and filiforms (giant, rousette, leaf-like and spearlike filimorms). The gustatory papillae are either fungiform (with one or two taste buds) or vallate (with a single, barrel-shaped taste bud). See also Massoud and Abumandour (2020). Stomach - The histomorphological characters of the stomach were studied by Abumandour and Pérez (2017), who concluded that this had special characters different to other mammals. The Cshaped stomach is divided into three parts: an elongated fundic cecal region with a blunt end, a long and narrow tubular pyloric part, and a small cardiac vestibular part. These characteristics of frugivorous species will help in their adaptations to accumulate large quantities of bulky food material. Eye - Bojarski and Bernard (1988: 155) found that the retina shows a marked folding due to papillae projecting inwards from the inner choroid layer. These papillae are directed towards the nodal point of the dioptric apparatus, which makes that light coming in from an angle onto the papillae produces no shadow on the retina, which would otherwise result in the formation of an incomplete image. The eye of R. aegyptiacus possesses a tapetum lucidum, which increases the visual sensitivity by reflecting more light onto the retina and causes a faint red eyeshine observed while collecting the specimens. The eyes of this bat contain an average of 160,000 retinal ganglion cells (Coimbra et al., 2016: 191). These authors also calculated the minimum angle of resolution as about 0.167° (ca. 5-6 mm at 1 m distance). Surprisingly, the eye of R. aegyptiacus contained a horizontal streak, which is generally found in species which roost in open environments (e.g. E. helvum, E. franqueti or E. wahlbergi. The horizontal streak of high retinal ganglion cell density in megachiropterans may afford increased resolution across the horizon, potentially assisting with predator surveillance (Coimbra et al., 2016: 195). Skull - Curtis and Simmons (2018: 1390) described the extensive calcification of the external rostral cartilages surrounding the dorsolateral sides of the nostrils. These elements do not seem to be composed of bone as no porosity nor trabeculae were found. Michelmore et al. (1998: 319) and Meng et al. (2016: 1) reported that the endocrine component of the pancreas occupied 9.1 % of the total volume, which is at least nine times higher than in humans. The former authors (p. 324) also indicated that the endocrine structure of the pancreas is more similar to that of snakes than that of other mammals. FUNCTIONAL MORPHOLOGY: Holland and Walters (2005: 83) report on the movements of the pinnae in free flight and demonstrated that the ears have a greater sensitivity to click stimuli in front of the animal when directed forwards than when directed back and to the side. They also indicated (p. 87) that echolocation signals are significantly more likely to be emitted during the forward sweep of the ears. Lancaster et al. (1997: 110) studied the wing temperature during flight and found that, on average, the wing surface temperatures were about 15 °C below the body temperature, and only some 1.8 (± 0.5) °C above the ambient temperature. Barclay et al. (2017: 572) found that the bat's body temperature exhibited a circadian cycle, with an average temperature of 37.7 °C when returning from foraging and decreasing to 35.5 °C by mid-day. Just before leaving for foraging, the temperature was on average 37 °C. Korine and Arad (1993: 61) indicated that the African Chiroptera Report 2020 thermoneutral zone ranged between 31 and 36 °C, and that the bat's oxygen consumption in this range averaged 0.95 ± 0.15 ml g-1 h-1. Experiments executed by Czenze et al. (2020: 4) indicated that R. aegyptiacus started to show signs of severe heat stress once the body temperature reached about 38 °C, which was 2 °C lower than the value reported for the tree roosing Epomophorus wahlbergi. In their study on the kinematics of the chiropteran shoulder girdle in flight, Panyutina et al. (2013: 393) found that the scapula and clavicle are making a very minor contribution to the wing movement as compared to the input of humerus movements relative to scapula in the shoulder joint. The shoulder girdle moves relative to thorax as a crank mechanism, where the clavicle plays the role of crank, and the scapula the role of connecting rod. Amador et al. (2015: 447) refer to Norberg (1972), who provided a detailed description of the left wrist of R. aegyptiacus. From this (and their own study of the handwing of a number of yangochiropteran bats), they concluded that the "dactylopatagium brevis" (the small piece of wing between the thumb and the second digit) functionally belongs to the propatagium (the piece of wing in front of the arm, between the thumb and the body). Greville et al. (2018: 974) studied the flight membrane healing in R. aegyptiacus and found that wounds in the plagiopatagium took about 50 % longer to heal than wounds in the chiropatagium. The interaction between vision and echolocation was investigated by Danilovich and Yovel (2019), who found that the bats learn the 3D shape of objects through vision and that they can classify objects using echolocation. During navigation, both senses are used: vision to decide where to fly and echolocation when approaching an obstacle. Hostnik et al. (2020: 897) used CT scans to investigate the differences in lung volume between bats in their natural (head-down) and inverted (head-up) position. In head-up position, the lung volume was significantly larger. SEXUAL DIMORPHISM: See Albayrak et al. (2008) for samples from Turkey. In South Africa, Jacobsen and Du Plessis (1976: 272) found that females were on average 18 g lighter than males during winter and spring. Females had a slightly larger average forearm length (92.3 versus 89 mm). 163 ECHOLOCATION: Griffin et al. (1958: 112) performed a number of experiments in total darkness where R. eagyptiacus bats were able to avoid vertically placed metallic wires of 3 mm diameter in 85 % of the time. For 1.07 mm wires, the succes percentage was reduced to 68 %, and for 0.46 mm wires it dropped to 45 %, which was still better than chance. Rainho et al. (2010a: 21) report the calls for 8 individuals from Saõ Tomé. Waters and Vollrath (2003: 209) found that the sonar clicks have a duration of about 250 micoseconds, with most of its energy occurring during the first 100 microseconds. With these clicks, the bats can detect and avoid wires of at least 6 mm in diameter. Yovel et al. (2011b: 515, 522) indicate that the sonar clicks produced by Rousettus species do not have a long duration as was generally believed, but are actually extremely short: 50 - 100 microseconds. The peak energy of the click is at 30 kHz, and the bandwidth is about 57 kHz. These signals could be, in some parameters, as good or even better than the signals used by Eptesicus fuscus (see Geva et al., 2010: 147). Davies et al. (2013b: Table S8) report a peak energy frequency of 35.03 kHz and a range between 12 and 70 kHz. Heffner et al. (2019: 584) mention a frequency range from 14 to 60 kHz, with peak energy between 25 and 30 kHz. At 60 dB, the hearing range for these bats extends from 2.25 to 64 kHz. Benda et al. (2012a: 182) report the following data for Iranian specimens: Start frequency: 29.7 ± 3.5 (25.7 - 31.8) kHz, end frequency: 23.4 ± 4.0 (19.1 - 26.9) kHz, FMAXE: 30.7 ± 3.9 (23.3 - 35.0) kHz, pulse duration: 1.4 ± 0.2 (1.0 - 1.5) msec, interpulse interval: 29.8 ± 16.5 (7.0 - 66.3) msec. Yovel et al. (2011a: 2) found that the intensity of the click emissions alters when they approach and lock their sonar on a target, and that R. aegyptiacus uses an undescribed strategy to change the spatial region ("field-of-view") their sonar covers: They increase the angle between the sonar beam axes of the click-pairs they emit. Lee et al. (2017) studied this further and concluded that the observed beam features can be captured by inducing changes to the pattern of phase differences through moving tongue location (see also Kling, 2018). Danilovich et al. (2015: R1124) also found that the click rate increases prior to landing, indicating that 164 ISSN 1990-6471 they adjust biosonar sampling in a task-dependent manner. From tests in a laboratory environment, Danilovich et al. (2014: S32) found that R. aegyptiacus can transfer information acquired using echolocation to the visual modality, suggesting they can create an echo-based representation of the world that is accessible to other sensory modalities. As to the value of echolocation for fruit-eating bats, Fenton (1994: 27) suggests it permits access to the relative safety of dark, underground roosts, a situation analogous to the nesting situations of the echolocating birds. However, Danilovich et al. (2015: R1124) found that R. aegyptiacus not only relies on echolocation in dark caves, but also during orientation and foraging in diverse light environments. The click rate and intensity increased at lower light levels, where visual information was limited. Fenton (2013: 1) indicated that it remains to be determined if the tongue click signals can be used to monitor the activities of this species as these calls are much less conspicuous than the high intensity call used by other bats. Boonman et al. (2014: 2965) mentioned that they recently recorded wing clicks produced by wild R. aegyptiacus bats while they were flying in the field. Burchardt et al. (2019: 2) indicate that wingbeat and echolocation calls in R. aegyptiacus are tightly coupled around 5 - 12 Hz (the so-called theta frequencies: brain wave frequencies which are known to play a role in active movement and stimulus intake), suggesting a neuronal correlation. A large amount of vocalizations was collected by Prat et al. (2017b) (for samples see: https://doi.org/10.6084/m9.figshare.c.3666502). MOLECULAR BIOLOGY: DNA - See Ali (2011). Teeling et al. (2017: 32) mention the estimated genome size for R. aegyptiacus to be 2.11 C, and the assembly size 1.91 Gb. Szczesniak et al. (2013) presented the complete sequence of the mitochondrial genome of "R. leschenaulti" (16,704 nt, GenBank record KC702803). However, as pointed out by BoteroCastro et al. (2014), the sequence actually pertains to R. aegyptiacus. Lee et al. (2015: 2) reported on a "de novo transcriptome assemby of R. aegyptiacus" and assessed its quality and biological validity. In their phylogenetic study, Hassanin et al. (2016: 524) found that the Cytb from individuals collected in the eastern Mediterranean region (Cyprus and Egypt) differed by 13 mutations from populations of sub-Saharan Africa. Additionally, some Cytb haplotypes of southeastern Africa were found to be highly divergent from the others, whereas the population from West Africa did not differ significantly from that of Central Africa. Stribna et al. (2019: 2336) distinguished two mitochondrial haplotypes: Type I is widespread in regions where there is an extensive tree cover (e.g. tropical rain forests) and has close relationships to isolated lineages in the Middle East and islands in the Gulf of Guinea. The second type or haplogroup is sister to the rest of the R. aegyptiacus radiation and occurs in southern and eastern Africa, northwards to the Sudanian savanna, where it is found in habitats with semi-open land cover. Karyotype - 2n = 36, FNa = 66, BA = 32, and a submetacentric X chromosome (Ðulic and Mutere, 1973b; 1977; Haiduk et al., 1981: 223 from South Africa; Albayrak et al., 2008 for samples from Turkey). Albayrak et al. (2008) reported FN = 70 for a female and Fna = 66 from Turkey. Sayed (2011: 658) reported FN = 68 for Egyptian specimens and indicates the X chromosome is a medium sized submetacentric and the Y is a minute acrocentric one. She also reported that the metacentric autosomal pair 10 has a secondary constriction near its centromere and is of variable size within the same genome. Pairs 7 and 8 also show polymorphism. Pairs 9, 14, and 15 show a G-negative band in their short arm, and pair 10 has a broad G-negative band around its centromere. Pair 17 has complete heterochromatin, and displays polymorphism, with sometimes one autosome being subacrocentric and the other submetacentric. Denys et al. (2013: 281) report the karyotype of four specimens from the Guinean Mount Nimba area as 2n = 36, Fna = 66, with 12 metacentric pairs, two subtelocentric pairs and one small acrocentric pair. The Y chromosome is the smallest in the set. Protein / allozyme - Unknown. Hoenerhoff and Williams (2004: 592) reported hepatic copper values for 7 individuals, which ranged from 5.42 to 99.20 parts per million on dry weight basis. Águeda-Pinto et al. (2019: 4) studied the S100A7 (S100 Calcium Binding Protein A7 = psoriasin) African Chiroptera Report 2020 165 gene in a number of bats and found that R. aegyptiacus possessed three copies of this gene, which might contribute to the bat's enhanced tolerance for infections. 469) furthermore found that native forest was prefered over residential areas (59.1 % versus 40.9 %) to forage in, but there were great individual differences. Stasiak et al. (2018: 683) indicate that hemochromatosis (iron storage disease) is is a frequent cause of liver disease and mortality in captive Egyptian fruit bats. They found that Hepcidin mRNA expression increased in response to iron administration in healthy bats. Tsoar et al. (2010: 300) report that tracked fruit bats flew over relatively long distances (14.6 ± 3.7 km) at a speed of 33.0 ± 5.2 km/hr and at relatively high altitudes (108 ± 52.6 m) to commute between their roosting and feeding sites. Geva-Sagiv et al. (2015: 99) indicate that there are five main types of navigational strategies: beaconing, route following (or route guidance), path integration, cognitive map, map and compass. They also suggest that combinaties of these strategies can be used. HABITAT: Sclater (1868: 404) reported on a "Cynonycteris collaris" specimen that was captured at sea off the St. John's River in Natal on 1 March 1868. Arumoogum et al. (2019: 188) simulated the role of biotic and a-biotic factors in view of climate change and found that the current suitable habitat is primarily mediated by temperature, but under both moderate and extreme future climate change scenarios the most influencial factor would be the distribution of freestanding fig trees. HABITS: Jacobsen and Du Plessis (1976: 271) report that, in South Africa, radio-tracked R. aegyptiacus specimens travelled 24 km between their roost and foraging sites, which took them about 1 hour and 29 minutes at an average speed of 16.2 km/h (see Jacobsen et al., 1986: 207). They also found that these bats foraged on Ficus spp. inside the forest canopy and that seeds were dispersed while flying and consuming fruit on perches. The bats left their roosts between 20 and 40 minutes after sunset and started to return at 2 a.m., with a peak around 3.45 a.m. and most were back at 5 a.m. In Cape Town, Barclay and Jacobs (2011: 468) report that individuals left the roost cave between 21 and 183 minutes after sunset, and that bats left their roosts later when it was raining at their usual emergence time. Furthermore, males left later than females in September - October (during early pregnancy), but not in November - January (during late pregnancy and lactation). The difference in emergence time between females and males in September - October is attributed to the males having to defend their roosting territory (Barclay and Jacobs, 2011: 471), and not to females having to reach fruit resources before males, or to travel to more distant fruit patches, because of the greater energy demands as was also suggested by Korine et al. (1994) for R. aegyptiacus in Israel. Barclay and Jacobs (2011: 469) also observed that the individuals left their roost in a consistent order. The bats returned to their roost between 217 and 21 minutes before sunrise, without any differences among the sexes. Barclay and Jacobs (2011: In the Dakhla Oasis in Egypt; Lucan et al. (2011: 36) reported that some individuals made quite long flights of about 60 km at the beginning of their night activity. They also found that spatial activities (e.g. roost changes and night exploration of foraging grounds) was strongly reduced by the end of winter, when few food resources remained available. Bachorec et al. (2020: 1), however, observed a decrease in home range size during spring, when food availability was lowest. Using an automated tracking system, Rerucha et al. (2014: 14) found that the feeding time showed a much larger variation in summer than in spring or winter, whereas the search time per night remained more or less stable. The opposite was true for winter, where the total feeding time remained more or less stable, but the total search time showed a large variation. Bachorec et al. (2020: 11) also found that, when food was abundant, Egyptian fruit bats choose to forage with more related individuals. Jahelková and Vašícková (2010: 182) describe some socal calls and social behaviour (e.g. protection of mother and juvenile by a male during lactation) in a captive population. Prat et al. (2016: 1) examined the vocalizations of these bats and found that these contained information about identity of the emitter, the context of the call, the behavioral response to the call, and even the call's addressee. These vocalizations are composed of sequences containing 1 to 20 broad-band multiharmonic syllables. Almost all of the communication calls were emitted during aggressive pairwise interactions. Mann et al. (2011: 416) indicated that males do not form random associations with nearby conspecifics, but that they establish stable groups, in which some form of dominance hierarchy exists. They also found that R. aegyptiacus can make a distinction between familiar and unfamiliar 166 ISSN 1990-6471 conspecifics, most likely by means of olfactory and/or visual cues (Mann et al., 2011: 411). ROOST: Type - Caves (Albayrak et al., 2008); abandoned factories (Albayrak et al., 2008). Microclimate - Unknown. Orozco (2012: 379), referring to Braack (1989), indicates that no scarab beetles were found in over 40 South African caves inhabited by R. aegyptiacus. Braack (1989: 77), however, does mention the presence of reduviid bugs (Platymeris guttatipennis (Stal)) preying on cockroaches (Gyna caffrorum (Stal, 1856), Gyna spp., Herbardina spp., Symploce incuriosa (Saussure, 1899)), parasitic evaniid and chalcidoid wasps (Hockeria sp.), trogid beetles (Trox zumpti Haaf, 1957) utilizing dead bats, tenebrionid (Tenebrio guineensis Imhoff, 1843) and ptinid beetles, thysanurans, as well as numerous haematophagous argasid ticks (Ornithodoros faini Hoogstraal, 1960) and nycteribiid flies (Eucampsipoda africana Theodor, 1955) using the bats as food source. Other "visitors" of cave, inhabited by R. aegyptiacus, were ants (Pheidole crassinoda Emery, 1895), spiders (Smeringopus sp.), troglophilic crickets, In South Africa, Herzig-Straschil and Robinson (1978: 104) found R. aegyptiacus roosting together with Miniopterus natalensis [as M. schreibersi], Rhinolophus capensis, a few M. fraterculus and possibly R. clivosus. Similar associations were also reported from Zambia (Leopard's Hill cave) by Grafitti et al. (2003: 70), who reported on a colony of approximately 10,000 R. aegyptiacus, together with three colonies of M. schreibersi of about 50 - 300 animals, and single specimens of R. hildebrandtii, R. simulator and Nycteris sp. DIET: In Turkey, Albayrak et al. (2008) found two wild fruits were important (Ficus elastica (Rubber tree) and Melia azadirachta (Persian lilac)). In South Africa, Barclay and Jacobs (2011: 467) report that the diet includes the fruit of multiple species of figs, but also those of Cape ash (Ekebergia capensis Sparrm.), saffronwood (Cassine crocea (Thunb.) Kuntze), yellowwood (genus Podocarpus Persoon), Diospyros L., and Syzygium R.Br. Ex Gaertn. Herzig-Straschil and Robinson (1978: 101) also included Bushman's poison (Acokanthera oppositifolia (Lam.) Codd) and Mistletoe (Viscum obscurumi Thunb.). Lamb et al. (2010: 215) reported that Cypriotic R. aegyptiacus specimens fed on Agave americana (Century plant) during mid-summer, but moved on to Ficus carica (figs) when these became available. During late-winter, the most important food items were Phoenix dactylifera (dates), Melia azedarach (Persian lilac), and flowers of eucalypt, Myrtus communis (common myrtle), mandarines and lemons. Del Vaglio et al. (2011: 283) identified 11 plant species in the droppings of Cypriotic specimens, including Melia azedarach (Perzian lilac or Chinaberry tree), Morus spp. (Plum tree), Ceratonia siliqua (Carob tree or St John's-bread), Eryobotria japonica (loquat, Japanese medlar or Japanese plum), Ficus carica / F. microcarpa (Common fig tree / Laurel fig), Arbutus andrachne (Eastern or Greek Strawberry Tree), Washingtonia filifera (Washingtonia), Crataegus azarolus (Mediterranean hawthorn), Citrus sinensis (Orange tree), and Styrax officinalis (Stirax). All of these plant species had limited economic importance. Baum (1995: 339) mentioned that R. aegyptiacus is a frequent visitor of Adansonia digitata (Baobab), especially in the first two hours after dusk, where they licked nectar from the petal bases. Jacobsen and Du Plessis (1976: 271) found that in South Africa this bat fed mainly on Ficus spp. [F. capensis, F. petersii, F. sansibarica], although other fruits (e.g. Litchis chinensis - litchis, Syzgium gerrardii - water pear, Syzgium jamba jamba, Syzygium cordatum - water berry, Harpephyllum caffrum - kafflI plum, Ekebergia capensis - Cape ash, Prunus africana - bitter almond/red stinkwood, and possibly Olea capensis and O. africana) were eaten in other seasons. Nader and Kock (1983c: 271) also reported that captive bats ate 8 to 9 litchis per night or 97 g of fruit (bananas). The bats were probably also feeding on the nectar of Aloe dolomitica as they found a number of animals covered in pollen and their faeces contained pollen too. Seltzer et al. (2013: table S2) provided an overview of the plant species found beneath bat feeding roosts. For R. aegyptiacus these include Ficus mucuso Welw. ex Ficalho (Ipoh's terrible tree - Moraceae), Ficus sur Forssk. (Broom cluster fig Moraceae) and Eriobotrya japonica (Thunb.) Lindl. (Loquat - Rosaceae). Amitai et al. (2010: 2693) used stable carbon isotope ratio in exhaled CO2 to assess the origin of metabolized substrates in 16 R. aegyptiacus, which were maintained on a diet of C3 plants. Amitai et al. (2010: 2693) found that both resting and flying individuals directly fueled their metabolism with freshly ingested exogenous substrates. In Cape Town, Barclay et al. (2006: 549) observed R. aegyptiacus deliberately feeding on Scarabid garden fruit chaffer beetles (Pachnoda sinuata). African Chiroptera Report 2020 Del Vaglio et al. (2011: 285) also reported the presence of coleopterans, lepidopterans (caterpillars), dipterans (fruit flies) and ticks in the bat's droppings, but these were most probably ingested by accident. In his study on R. aegyptiacus from the Kruger Park, Braack (1989: 79) found that they were feeding on the fruits of Ficus sycomorus (sycamore fig or fig-mulberry), Diospyros mespiliformis (Jackalberry, African Ebony, jakkalsbessie), and Xanthocercis zambesiaca (Mashatu tree or Nyala tree). In the Kisangani area (DRC), Gembu Tungaluna (2012: 106 - 108) followed the fruit intake by R. aegyptiacus during the different seasons and came up with the following list: First heavy rain season (September - November): Carica papaya L. (papaya) (28%), Musanga cecropioides R. Br. & Tedlie (African corkwood tree, umbrella tree) (14%), Dacryoides edulis H.J. Lam (safou, African pear, bush pear, Nsafu, bush butter tree, butterfruit) (11%), Persea americana Mill., 1786 (avocado) (8%), Oncoba welwitschii Oliv. (8%), Myrianthus arboreus P. Beauvois, 1805 (giant yellow mulberry, bush pineapple, corkwoord) (8%), Spondias cytherea Sonnerat, 1782 (Ambarella, otaheite apple, great hog plum) (6%), Annonidium mannii (Oliv.) Engl. & Diels (Junglesop) (6%), Ficus wildemanniana Warb. (5%), Ficus leprieuri Miq., 1867 (3%), Ficus vallis-choudae Delile (3%). First mild rain season (December - February): Musanga cecropioides (36%), Myrianthus arboreus (22%), Ficus vallis-choudae (10%), Annonidium mannii (6%), Spondias cytherea (6%), Carica papaya (6%), Uapaca guineensis Müll.Arg., 1864 (Sugar plum, red cedar, false mahogany, rikio) (6%), Elaeis guineensis Jacq. (African oil palm, macaw-fat) (2%), Ficus wildemanniana (2%), Pseudospondias microcarpa (A. Rich.) Engl. (2%), Musa sp. (2%). Second heavy rain season (March - May): Persea americana (19%), Annonidium mannii (16%), Carica papaya (16%), Dacryoides edulis (11%), Spondias cytherea (10%), Musanga cecropioides (7%), Oncoba welwitschii (7%), Myrianthus cecropioides (7%), Uapaca guineensis (3%), Ficus vallis-choudae (3%), Musa sp. (3%), Parinari excelsa Sabine (Guinea plum) (1%), Ficus sp. (1%). "Dry" season (June - August): Oncoba welwitschii (29%), Ficus vallis-choudae (15%), Parinari excelsa (14%), Carica papaya (14%), Myrianthus arboreus (14%), Musanga cecropioides (14%). Izhaki et al. (1995: 340) report, that in Israel, R. aegyptiacus ingested only 2 % of the small seeds (< 4 mg), all other seeds were spat out between 25 and 400 m from the parent tree. Korine et al. 167 (1998: 151) indicate that R. aegyptiacus feeds on two types of fruit ("syndromes"). The first one is similar to that eaten by most fruit bats: variable size (range 0.53 - 32.3 g), relatively high water content (57 - 87 %), relatively low fat content (1.3 - 4.1 %), low protein content (1.1 - 6.75 %), and relatively high sugar content (69 - 86 %). The second type is only found in eastern Mediterranean habitats and is characterized by the extremely low water content: 13 - 24 %. This type of food is generally eaten during winter, when other food sources are scarce. A case of hemochromatosis or ISD (iron storage disease or excessive burden of iron) was reported by Clauss and Paglia (2012: S8) as the result of an accidental excessive dietary iron supplementation. Stasiak et al. (2014: 111) investigated whether mutations in the hepcidin gene might be the reason for the increased susceptibility to ISD, but they were unable to identify any differences in the sequence of the portion of the hepcidin gene that encodes the mature peptide. In their ISD study, Leone et al. (2016: 45) provided evidence of a positive correlation between hemochromatosis and HCC in any species other than humans. In a study of R. aegyptiacus this was not evident as its blood plasma glucose concentrations were observed to be greater than 40 mmol/l after glucose uptake (Keegan, 1977; Michelmore et al., 1998), this was not after natural feeding but during an oral glucose tolerance test (Mqokeli and Downs, 2012a: 351). Herrera M. et al. (2015: 409) investigated the relationship between food and energy intakes, salt content and sugar types. PREDATORS: Jacobsen and Du Plessis (1976: 273) indicate that R. aegyptiacus has relatively few natural enemies in South Africa, but that genets (Genetta sp.) and Lanner falcon (Falco biarmicus Temminck, 1825) took them as prey. Mikula et al. (2016: Supplemental data) mention the following avian predators: Black sparrowhawk (Accipiter melanoleucus Smith, 1830), Lanner falcon (Falco biarmicus Temminck, 1825), Peregrine falcon (Falco peregrinus Tunstall, 1771), Bat hawk (Macheiramphus alcinus Bonaparte, 1850); Black-and-white-casqued hornbill (Bycanistes subcylindricus (Sclater, 1871)). Mikkola (2018: 4) shows a photo of a dead bat (with sucking pup) brought to the nest by an Eagle owl (Bubo bubo (L., 1758)). 168 ISSN 1990-6471 POPULATION: Structure and Density Common in parts of Africa; generally uncommon in SW Asia although locally abundant in Israel and Jordan (Z. Amr pers. comm., 2005 in Benda et al., 2008a in IUCN, 2009 and Korine (2016)). In Africa it occurs in large colonies of up to 40,000 to 50,000 individuals. In SW Asia colonies generally number 50 to 500 individuals, although up to 3,000 individuals were recorded in a cave in Jordan. The population in Turkey is estimated at 5,000 to 10,000 individuals; the population there may be decreasing due to control measures in caves (A. Karatas pers. comm., 2007 in Benda et al., 2008a in IUCN, 2009 and Korine (2016)). In Syria only a single locality is known of 1,000 to 2,000 animals (A. Karatas pers. comm., 2007 in Benda et al., 2008a in IUCN, 2009 and Korine (2016)). Also see Albayrak et al. (2008). Horácek et al. (2010: 175) indicate that there is a clear decline in the Cypriotic population, whereas the situation in other Mediterranean countries (Lebanon, Jordan, Egypt) is more or less stable. They estimate the total Mediterranean population to include at most 75,000 specimens. Lucan et al. (2014c: 111) report a population decline of 90 % on Cyprus between 2005 and 2013 (from 8,000 to 800 animals). Trend 2016: Stable (Korine, 2016). 2008: Stable (Benda et al., 2008a; IUCN, 2009). R. aegyptiacus populations within the Asia Minor and Levant region are predicted to have a stable population trend, under climate change scenarios (Bilgin et al., 2012: 433). LIFESPAN: Mitchell (1911: 444) mentions for 13 captive specimens (as Cynonycteris collaris) an average lifespan of 18 months and a maximum lifespan of 108 months (= 25 yrs). Cushing et al. (2013: 794) reported on a captive female of over 11 years of age that needed to be euthanised following a deterioriation of its health due to a metastatic pancreatic carcinoma. Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 22.9 years. Olds et al. (2015: 325) found the following tumor types in a captive population of male R. aegyptiacus: fibrosarcoma, cutaneous lymphoma, benign focal bronchioloalveolar neoplasm, anaplastic sarcoma, and sebaceous epithelioma. ACTIVITY AND BEHAVIOUR: Carpenter (1986: 93) mentions that the heart beat for this species is as high as 728 beats per minute during a flight with airspeeds of 5 m/s. During displacement experiments, Tsoar et al. (2011: E722) found that R. aegyptiacus primarily uses self-triangulation based on multiple distal visual landmarks to return to their roost cave. Ulanovsky et al. (2010: 302) provide evidence that the bats used a large-scale navigational map to return to their roosts after having been displaced over 44 km. From flight corridor tests, Riskin et al. (2012: 2946) determined the average flight speed to be 4.41 ± 0.54 m/s. In captive groups, Carter and Leffer (2015: 3) found that a group consisting of nine females and two castrated males spent 0.26 % of their awake time on social grooming. In a group of five males, no social grooming was observed at all. REPRODUCTION AND ONTOGENY: Korine and Arad (1999) [in Andrianaivoarivelo et al. (2011a: 77)] mention that the newborns of R. aegyptiacus are completely dependent on their mothers for the first 6 weeks of life. At birth, the pup weights about 18.0 % of the mother's body mass (22.7 versus 126.0 g) (Kurta and Kunz, 1987: 82). Barclay and Jacobs (2011: 467) indicate that the lactation period lasts from 9 to 12 weeks in South Africa. Davis (2007: 4) indicates that young are dependent until three months after birth in the UAE. At that time, they are able to fly, but do not leave the family colony. In Uganda, Mutere (1968) observed a bimodal breeding pattern (pregnancies from December to March and from July to October and births in March and October), which was correlated to two rainfall peaks (April-May and October-November). Bartonicka et al. (2013: 17) report that in both the Levant (Turkey, Cyprus, Lebanon and Jordan) and in the Dakhla Oasis in Egypt, births take place during most of the year, except for the winter months. The births peak in April-May and in August-September. For Malawi, Happold and Happold (1990b: 564) reported births in mid-October, but they had no data for other months. In Sierra Leone, Weber et al. (2019: 20) captured one lactating female on 7 April. Lucan et al. (2014a: 110; 2014b: 1036) investigated the reproductive cycle in populations from the Mediterranean region and the Egyptian desert and found that both had a bimodality in birth timing regarless of climatic differences. The differences in birth times between the two populations were linked with differences in fruiting phenology of their major dietary plants. They also African Chiroptera Report 2020 hypothesize that food abundance is an important trigger of male sexual activity. Spitzenberger and Bauer (1979: 440, 448) also found a bimodal reproduction cycle for R. aegyptiacus on Cyprus, but remarked that the succession of the two phases is so quick that females only take part in one cycle. Kozhurina and Ilchenko (2016: 13) found a bimodal reproducion cycle in captive R. aegyptiacus with a "tropical" polyoestrous continuous reproductive pattern, but found their cycle to take 11 months rather than 12. They also found that females usually began to produce offspring at the age of 14 - 15 months up to 2 years. The gestation period was very variable and lasted from 95 to 129 days. Mahmoud et al. (2018: 293) tested 3monochloropropane-1, 2-diol (3-MCPD) to sterilize male rousettes to assess its use as pest management tool. 169 Laboratory tests performed by Taub et al. (2014: S122) provided strong evidence for vocal learning in this species. Khannoon et al. (2020: 309, 313) described 13 developmental stages from middle pharyngula through immediately before birth, indicating that the wings are formed during stages 19 to 20. The nasal pits appears at stages 14 to 15, and at stage 17, the face starts to look 'fox-like' to attain its final shape at stage 19. Until stage 22, the crownrump length of the embryo is very similar to that of embryos of other bat species, but then it starts to grow very rapidly. MATING: Jacobsen and Du Plessis (1976: 272) and Barclay and Jacobs (2011: 467) mention that in South Africa, R. aegyptiacus mates in spring (July to September) and, after a gestation of approximately 4 months, a single pup is born in summer (end October to December). Occasionally, Start (1969: 272) recorded the birth of twins. Stanley and Goodman (2011: 40) reported on three adult males with scrotal testes collected in the first half of August 1992 in Tanzania. Two females collected in the first halfs of August 1992 and 1993 were pregnant with a single embryo. Trentin and Rovero (2011: 49) report a lactating female captured in December 2005 in the Uzungwa Scarp Forest Reserve, Tanzania. For West Africa, Krutzsch (2000: 110) indicates that the male reproductive cycle is apparently in synchrony with the female, since they undergo biseasonal testicular hypertrophy. In a colony in the Prague zoo, Jahelková et al. (2013: 76) found courtship behaviour and matings taking place throughout the year, but parturitions were strongly synchronized in a one-week period in September/October and March/April. In the southern Cape province of South Africa, Herzig-Straschil and Robinson (1978: 108) calculated that births took place from October to June. At Matlapitsi (RSA), Dietrich et al. (2018b: 4) found that 64 % of the females they sampled in September 2015 were pregnant, indicating that there were also non-pregnant females and both scrotal and non-scrotal males. In January 2016, they found large amounts of juveniles and lactating females, whereas in April 2016, most of the bats (83 %) were flying juveniles. POSTNATAL DEVELOPMENT: Prat et al. (2013: 123) studied R. aegyptiacus pups raised in small colonies and in acoustic isolation and found that the latter displayed delayed vocal development and that their calls differed from young of the same age raised in colonies. The first sounds produced by young in both groups were isolation calls (Prat et al., 2015: 1). These calls are innate and appear at postnatal day 1. Prat et al. (2017a) investigated vocal learning in pups that were exposed to playback calls, and found that their calls demonstrated distinct dialects, which were related to the playback calls they were acustomed to rather than to the calls produced by their mothers. In the United Arab Emirates, Davis (2007: 4) indicated that mating takes place from June to September, and that a single young (exceptionally twins) are born in October-December after a gestation period of 115 to 120 days. Szekely et al. (2015: Suppl.) report a gestation period of 116 days and they mention that weaning occurs after 122 days. During that period, the weight of the young has increased from 21.06 g to 73.8 g (adult: 125 g). In a captive colony in Israel, Harten et al. (2019: 1895) determined the paternity of pups and found that females gave birth to young of males from which they used to scrounge food [=obtaining food from the mouth of the male], supporting the foodfor-sex hypothesis in this species. Genzel et al. (2019: 1) studied the vocal plasticity in adult bats and found that adult bats are able to modify distinct parameters of their vocalizations when exposed to broad-band, acoustic perturbation. The changes can persist months after the noise has ceased. 170 ISSN 1990-6471 PARASITES: BACTERIA Gram-negative bacterium - Pinus and Müller (1980) and Sara (2002: 41) reported Citrobacter freudii, Enterobacter cloacae, Escherichia coli, Klebsiella pneumonia, Proteus morganii, and Proteus rettgeri. Han et al. (2010) isolated a pure sample of Kluyvera ascorbata from blood of an individual found dead at a zoo in Seoul, Korea, which had originally been imported (possibly illegally) from Egypt. Childs-Sanford et al. (2009: 8) reported Yersinia pseudotuberculosis from a colony of bats kept in a zoo in Syracuse, New York. Two of three bats from the Republic of Congo, tested by Nowak et al. (2017: 6) were positive for E. coli. Hughes et al. (2018: 867) reported the presence of Yersinia pseudotuberculosis in bats from a zoo colony. Gram-positive bacterium - Three out of 55 tested R. aegyptiacus bats from Gabon tested positive for Staphylococcus aureus and two for S. schweitzeri (Held et al., 2016: 119). The positive bats were captured at the fringe of the forest, in the vicinity of humans. None of the bats that were captured in the forest were infected by this bacterium. Bartonellae - Bacteria Bartonella - Kosoy et al. (2010: 1878) found 15 Bartonella spp. gltA sequences in R. aegyptiacus, and also found six unique gltA varients of Bartonella which clustered in a monophyletic genogroup. Kosoy et al. (2010: 1877) and Bai and Kosoy (2012: 58) reported a prelevence of Bartonella spp. in R. aegyptiacus from Kenya of 22/105 (21.0%) cultured from blood samples. See also Kosoy (2010: 719) for further information. Dietrich et al. (2016b: 3) found one positive testing bat (out of 7) in Mahune (RSA). Bai et al. (2018: 2319) identified a new Bartonella species (B. rousetti) from bats collected in the Idanre Hills area in Nigeria, which can infect humans as they also found antibodies against this bacteria in people living in surrounding communities. Lei and Olival (2014: Suppl.) mention R. aegyptiacus as host for Leptospira bacteria. These were further identified as L. interrogans and a second species as having sequences closely related to L. borgpetersenii. Nakamura et al. (2012: 410) report on an outbreak of yersiniosis (caused by Yersinia pseudotuberculosis) in a colony of captive R. aegyptiacus in a zoological park in the Kanto region in Japan. Kolodny et al. (2018: 116) examined the microbial composition of fur and gut samples of a captive and free-living colony of Egyptian fruit bats and found that the fur samples of different animals from the same colony taken at the same time were more similar than samples taken from the same animal at different time. The gut samples, however, were more individualized and sex related. Dietrich and Markotter (2019: 1733, Suppl.) investigated the differences between direct and indirect sampling of urine and feces in South African R. aegyptiacus. During this study, they found the following bacteria in the urine (in descending LDA [Linear discriminant analysis] order): direct: Streptococcus, Bacillus, Rhizobium, Brevudimonas, Rhizobiales, Corynebacterium, Achromobacter, Acinetobacter, Phenylobacterium, Deinococcus, Methylobacterium, Paenibacillus, Lyninibacillus, Cellulomonas, Cellulosimicrobium, Planococcaceae, Nocardiodes, Serratia, Devosia, Solirubracacter, Providencia, Brucella, Sphingomonadacea, Stenotrophomonas, Oxalobacteraceae, Gp4, Terribacillus, Gemmatimonas, Ochrobactrum, Gp6, Blastococcus, Comamonadaceae, Microvirga, Sphingobacteriaceae, Microbacteriaceae, Gp3, Solirubrobacterales, Massilia, Arthrobacter, Geodermatophilaceae, Microbacterium, Leucobacter, Beijerickiaceae, Planococcaceae, Burkholderiales, Shinella, Rubellimicrobium, Gp16, Novosphingobium, Alcaligenaceae, Rubrobacter, Betaproteobacteria, Dyadobacter, Actinomyces, Agromyces, Ruminococcaceae, Polyangiaceae, Lysobacter, Bradyrhizobiaceae, Moraxella, Aurantimonas, Acetobacteraceae, Flavobacterium, Ramlibacter, Caenispirillum, Bordetella, Leifsonia, Paenibacillaceae_1, Bacillaceae_1, Xanth0obacteraceae; indirect: Planctomycetes, Lentibacillus, Actinocatenispora, Xanthomonadales, Marinomonas, Ignavibacterium, Jiangellaceae, Methylohalomonas, Fodinicurvata, Salinicoccus, Nocardiopsis, Deinococcales, Ectothiorhodospiraceae, Nitrosospira, Nocardia, Saccharopolyspora, Nocardiopsaceae, Nakamurellaceae, Ureoplasma, Pntibaca, Phycisphaera, Nitrococcus, Actinophytocola, Nesterenkonia, Luteibacter, Enterobacteriaceae, Jiangella, Brevibacteriu, Bacteroidetes, Gammaproteobacteria, Salinisphaera, Actinomycetales, Gracillimonas. In the faeces, the following bacteria were found: direct: Actinobacillus, Pasteurellaceae, Flavobacteriaceae, Salinisphaera, Gp1, Gammaproteobacteria, Kytococcus, Mycobacterium, Gracilimonas, Chryseobacterium, Mycoplasma, Geodermatophilaceae, Chloroflexi, Rhizobium, unclassified Bacteria, Helicobacter, Rhodococcus, Blastococcus, Deinococcus, Chitinophagaceae, African Chiroptera Report 2020 Acinetobacter, Brevundimonas, Sphingobacteriales, Comamonadaceae, Sphingomonas, Mosesorhizobium, Comamonas, Gemmatimonas, Proteobacteria, Achromobacter, Agrmyces, Nitrosococcus, Devosia, Gp10, Gp3, Flavobacterium, Roseomonas, Marmoricola, Gp6, Ectothiorhodospiraceae, Campylobacter, Solirubrobacterales, Isoptericola, Nocardioidaceae, Deltaproteobacteria, Gordonia, Sphingomonadaceae, Neisseriaceae, Luteibacter, Actinomadura, Erythrobacteraceae, Acidimicrobiales, "3_genus_incertae_sedis", Nocardioides, Spirosoma, Truepera, Sphingomonadales, Moraxella, Solirubrobacter, Tessaracoccus, Alphaproteobacterai, Myxococcales, Paenibacillus, Bacteroidetes, Oxalobacteraceae, Stenotrophomonas, Phenyobacterium, Caulobacteraceae, Gp7, Arthrobacter, Polyangiaceae, Geminicoccus, Rubellimicrobium, Gp4, Azospirillum, Sphingobacteriaceae, Rhodospirillaceae, Dermacoccus, Nitrospira, Pseudomonadaceae, Acetobacteraceae, Cellulosimicrobium, Lysobacter, Dyadobacter, Sphingopyxis, Betaproteobacteria, Ornithobacterium; indirect: Granulicatella, Carnobacteriaceae, Streptococcaceae, Alkalibacterium, Nesterenkonia. Modesto et al. (2019: 8, 9) described two new species of Bifidobacterium from the feaces of a captive colony of R. aegyptiacus: B. vespertilionis and B. rousetti. HAEMOSPORIDA Perkins and Schaer (2016: Suppl.) mentioned the presence of Plasmodium roussetti van Riel, l'Hoest et l'Hoest, 1951 in bats from the DRC. Adam (1973: 8) mention on an infection by Trypanosoma megadermae Wenyon, 1909. Barbosa et al. (2016: 215) and Espinosa-Álvarez et al. (2018: Suppl.) reported a specimen from Gabon being infected by "Trypanosoma sp. bat". Atama et al. (2019: 1550) reported the first case of a Hepatocystis parasite present in R. aegyptiacus (in one of 32 bats from the Amurum forest reserve, Nigeria). ACARI Spinturnicidae: Hirst (1923: 979) indicates that Kolenati stated that either R. microphyllum or Pteropus ægyptiacus [=Rousettus aegyptiacus] is the host for Ancystropus zelebori Kolenati, 1856 (Acari). Another mite, he reported (p. 987) from R. aegyptiacus is Ancystropus (Meristaspis) lateralis Kolenati. Taufflieb (1962: 112) also reported Ancystropus leleupi Benoit, 1959. 171 Trombiculidae: Stekolnikov (2018a: 50) reported this species as host for Whartonia novemsetosa Goff, 1982 in Tanzania. Korine et al. (2012: 1435) experimentally infected R. aegyptiacus specimens with the flea Xenopsylla ramesis, whose natural host is Sundevall's jird (Meriones crassus). They found that female fleas engorged the same amount of blood from these bats as from the jirds. Male fleas, however, sucked much less blood. From the Lanner Gorge cave [22°27'S 31°09'E], in the Kruger National Park, Braack (1989: 81) reported argasid ticks (Ornithodoros faini Hoogstraal, 1960), nycteribiid flies Eucampsipoda africana Theodor, 1955), spinturnicid mites (Ancystropus zeleborii, Kolenati, 1857), and the flea Thaumapsylla breviceps Rothschild, 1907 (Ischnopsyllidae). In Jordan, Benda et al. (2010b: 201) found the following ectoparasites: Argasidae: Argas vespertilionis (Latreille, 1802), Ornithodoros (Reticulinasus) salahi (Hoogstraal, 1953). Nycteridocoptes pteropodi Rodhain and Gedoelst, 1923 (Acari: Sarcoptidae) was reported by Fain (1958: 236) from "Rousettus leachi" specimens from Kivu, DRC. Fain (1958: 244, 245) also described Nycteridocoptes lavoipierrei, Nycteridocoptes macrophallus and Nycteridocoptes rousetti from the same bat species collected at Mahuysa cave, Katana-Kivu, DRC. Another species (Nycteridocoptes rousetti Fain,1958) was mentioned by Fain (1959a: 345) from R. aegyptiacus from Thysville (=MbanzaNgungu), DRC. Fain (1960: 83) described Stomatodex rousetti (Acari: Demodecidae) from "R. leachi" from Mahyusha cave, near Katana, DRC. Fain (1967: 367) furthermore reported on the presence of Teinocoptes rousetti Fain, 1959 on R. aegyptiacus leachi from Mahyusha cave, Kivu, DRC. Reinhardt et al. (2008: 173) report that the cimicid bug Afrocimex constrictus Ferris and Usinger, 1957 infested R. aegyptiacus colonies at Makingeny and Ngwarisha (Mount Elgon region, Kenya) and that these bats were bitten between 0.5 and 1.5 times a day, and that to up to 28 µl of blood was removed each time. Afrocimex leleupi Schouteden, 1951 was reported by Kock and Aellen (1987: 877) the Kakontwe cave (DRC). In Iran, Benda et al. (2012a: 185) report the presence of Meristaspis lateralis (Kolenati, 1856) (Spinturnicidae). This mite was already reported 172 ISSN 1990-6471 by Delfinado (1963: 913) to be widely distributed on R. aegyptiacus in Egypt and Palestine. Bianco et al. (2019: 552) indicate that demodicosis, caused by Demodex mites (Demodicidae, Prostigmata) was reported from zoo-kept Egyptian fruit bats. DIPTERA Streblidae: Brachytarsina africana (Walker, 1849) recorded in Katanga, but Haeselbarth et al. (1966: 100) suggests that this may be accidental and R. aegyptiacus is not its true host. Brachytarsina alluaudi (Falcoz, 1923) recorded in Katanga (Haeselbarth et al., 1966: 101), Gabon (ObameNkoghe et al., 2016: 5). Brachytarsina bequaerti (Jobling, 1936) collected in Tanzania and Kenya (Haeselbarth et al., 1966: 101). Nycteribiidae: Eucampsipoda africana (Theodor, 1955), widely distributed over the Ethiopian region and recorded from localities in Senegal to the Sudan and southwards to the Cape (Haeselbarth et al., 1966: 115; Duron et al., 2014: 2108; ObameNkoghe et al., 2016: 5). This bat fly might be a carrier of Bartonella (Zhu et al., 2014: 2160; Szentiványi et al., 2019: Suppl.) and Arsenophonus bacteria (Szentiványi et al., 2019: Suppl.), and the fungus Anthrorhynchus eucampsipodae Thaxter, 1901 (see Balazuc, 1971: 214; Haelewaters et al., 2017: Suppl. for a record from Gabon; Haelewaters et al., 2018: 794; Szentiványi et al., 2019: Suppl.), as well as the blood parasite Polychromophilus melanipherus and Mahlapitsi and Wolkberg viruses [Szentiványi et al., 2019: Suppl.). Eucampsipoda hyrtlii (Kolenati, 1856) has been identified with certainty only from the Middle East (Arabia, Egypt, Palestine and Syria), but probably also occurs in East Africa (Haeselbarth et al., 1966: 115). Balazuc (1971: 214) reported this bat fly from a "Chiroptera" from Egypt, which was infected by the fungus Anthrorhynchus eucampsipodae Thaxter, 1901. However, as pointed out by Blackwell (1980: 151), E. hyrtlii is only known as parasiting on R. aegyptiacus (see also Haelewaters et al., 2018: 794). Dipseliopoda biannulate (Oldroyd 1953) distributed over the tropical parts of Africa and recorded from localities in Nigeria, Cameroon, the Congo and the Sudan and Kenya (Haeselbarth et al., 1966: 116). Morse et al. (2013: Suppl. 1) refer to Eucampsipoda africana (Diptera: Hippoboscoidea: Nycteribiidae: Cyclopodiinae) collected from a Kenyan R. aegyptiacus. This bat fly was also reported from Sierra Leone by Blackwell (1980: 146), where it was found to be infected by the fungus Arthrorhynchus eucampsipodae Thaxter, 1901. In Jordan, Benda et al. (2010b: 201) found Eucampsipoda aegyptia (Macquart, 1851). They also mention that two other Nycteribiidae were previously found in Palestine: Nycteribia pedicularia Latreille, 1805 and N. schmidlii Schiner, 1853. The latter species (as Nycteribia schmidlii scotti Falcoz, 1923) was also reported from Gabon by ObameNkoghe et al. (2016: 5). Jansen van Vuren et al. (2016: 1) describe a new orthoreovirus, which was collected from a bat fly (Eucampsipoda africana (Theodor, 1955)), which in turn was captured from a R. aegyptiacus bat from South Africa. SIPHONAPTERA Ischnopsyllidae: Thaumapsylla breviceps Rothschild 1907, where the nominate subspecies T. b. breviceps Hopkins and Rothschild, 1956, was found in the Cape, Natal and Transvaal provinces of South Africa, Kenya and DRC (Haeselbarth et al., 1966: 186). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Rousettus aegyptiacus: Adenoviruses, Astroviruses, Caliciviruses, Coronaviruses, and Rotaviruses. Willoughby et al. (2017: Suppl.) mention the following viruses: Chikungunya virus, European bat 1 lyssavirus, Israel turkey meningoencephalomyelitis virus, Kasokero orthonairovirus, Lagos bat lyssavirus, Marburg marburgvirus, Rousettus bat coronavirus HKU9, Shamonda orthobunyavirus, West Nile virus, Yogue virus, Zaire ebolavirus. Nieto-Rabiela et al. (2019: Suppl.) mention the following viruses: Bat coronavirus, Bat flavivirus, Bat paramyxovirus, Bat pegivirus, Betacoronavirus, Betaherpesvirus, Chikungunya virus, Cytomegalovirus, European bat lyssavirus, Gammaherpesvirus, Kasokero virus, Lagos bat virus, Marburg virus, Polyomavirus, Rousettus aegyptiacus papillomavirus, Sosuga virus, Yogue virus. In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against Yellow fever virus (YFV), non-specific Flaviviruses, Babanki virus (BBKV), non-specific Alphaviruses, and Rift Valley fever virus (RVFV). Adenoviridae Mastadenovirus Jansen van Vuren et al. (2018: 1) isolated a new virus (RaegAdV-3085) from an apparently healthy wild-caught Egyptian fruit bat in South Africa. African Chiroptera Report 2020 Coronaviridae - Coronaviruses SARS - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in 29 individuals from Limpopo Province, South Africa, 11 were tested positive (11/29, 37.9 %) and 142 individuals from Oriental Province, DRC, 17 tested positive (17/142, 12 %) (28/171, 16.4 %). In Nigeria during June 2008, Quan et al. (2010) screened the gastrointestinal tissue for the presence of coronaviruses by PCRs, none tested positive. Van Doremalen et al. (2018: 3, 10) inoculated 12 adult bats from a captive colony with WIV1-CoV (SARS-like WIV1-coronavirus) and were unable to detect efficient replication of the virus and only modest seroconversion in two of the bats. Tong et al. (2009) found several alpha- and betacoronaviruses in Rousettus aegyptiacus collected in Kenya in 2006 based on detection of coronavirus RNA in fecal swabs. Some of the coronaviruses were highly divergent, whereas others showed similarity to Rousettus sp. coronaviruses identified in China. Shehata et al. (2016) tested 265 Egyptian bats and found 12 infected with HKU9-like viruses. Anthony et al. (2017b: Suppl.) mention betacoronaviruses Bat_CoV_HKU9 and Kenya_CoV_BtKY56. 19 out of 397 (4.8 %) of the Kenyan bats tested by Tao et al. (2017: Suppl.) were positive for CoV. Nziza et al. (2019: 156) found Bat coronavirus HKU9 and Kenya bat coronavirus/BtKY56/BtKY55 in rectal swaps from Rwandan bats. The coronaviruses found in the Kenyan bats were identified as Beta-D coronavirus by Joffrin et al. (2020: 7). Filoviridae - Filo viruses Ebolavirus Muyembe-Tamfum et al. (2012: 9) refers to Pourrut et al. (2005) who found ZEBOV-specific antibodies in this species. Paweska et al. (2016: 1) experimentally inoculated R. aegyptiacus bats and were unable to find any viral RNA in oral, nasal, ocular, vaginal, penile and rectal swabs from any of the experimental groups. They concluded (p. 9) that these bats are unlikely to maintain and perpetuate EBOV in nature (see also Mandl et al., 2018: 7), which would qualify them as 'dead-end' hosts (Schuh et al., 2018: 1716). Marburgvirus Towner et al. (2007) tested 283 individuals from Gabon for Marburg virus [MARV] RNA by conventional and real-time RT-PCR and 242 individuals for anti-Marburg virus IgG antibodies by ELISA; 4 individuals (1.5 %) were PCR positive 173 and 29 (12 %) were IgG positive. Peterson and Holder (2012: 1831) indicate that the Gabonese clade represents a virus form that has not been reported from human infections. Kuzmin et al. (2010b) detected RNA in a female R. aegyptiacus bat collected at Kitum cave in July 2007. Adjemian et al. (2011: S796 - S798) report on four cases of MHR [Marburg hemorrhagic fever] in miners, which they contracted while working in a mine tunnel at Kitaka mine in Ibanda District, Uganda, where thousands of R. aegyptiacus bats lived. These cases resulted in the destruction of the residing bat population. Van Paassen et al. (2012: 635) report on the death of a Dutch woman, who probably contracted the virus after visiting a cave in Uganda, where R. aegyptiacus bats were living. Amman et al. (2013: 6) indicate that a longterm study at this site showed that 2 to 5 % of the bat population was actively infected by the MARV virus. Amman et al. (2014: 1762) estimated that both the cave and the mine contained about 40,000 to 100,000 R. aegyptiacus bats at the time the miners and the tourist were infected. The bats were exterminated by November 2008, but in November 2012 a small colony was (1 - 5 % of the original numbers) was found in the re-opened Kitaka Mine. Maganga et al. (2011: S800) indicate that MARV is enzootic in Gabon due to the fact that MARVpositive bats were captured in between 2005 and 2009. Kuzmin et al. (2011a: 4) report that MARV RNA was found in bats from Gabon, Uganda, and Kenya. Paweska et al. (2012: 1) inoculated captive R. aegyptiacus specimens - captured in northeastern South Africa - and were able to confirm the susceptibility of this species to infections with MARV irrespective of sex and age. In a subsequent study, Paweska et al. (2015) recovered the virus from 15 different tissues and plasma, but only sporadically in mucal swab, urine and fecal samples. Amman et al. (2015b) also experimentally infected bats and found viruses in liver, spleen, blood, kidneys, salivary glands, bladder, large intestine as well as in oral and rectal swaps. The infected bats did not show any significant changes in daily food consumption, body weight, or daily body temperature, and no signs of overt morbidity or behavioral changes such as waning appetite, overly aggressive behavior, separation from cage mates, lethargy, or reduced or abandoned grooming (p. 121). Schuh et al. (2017c) found prove for horizontal transmission of MARV as virus RNA was found in contact bats that were kept together with MARVinoculated bats. Schuh et al. (2017b), furthermore, reported that following infection of ERBs with MARV, virus-specific IgG antibodies 174 ISSN 1990-6471 are induced, but after 3 months the bats are seronegative again. Amman et al. (2012: 1) and Albariño et al. (2013a: 99) link an outbreak of MARV in October 2012 in Uganda with the second annual virus pulse in R. aegyptiacus populations, more specifically in older juvenile bats of about six months of age. The viral genome sequences collected from infected humans were approximately 99.3 % to two MARV isolates captured in 2008 and 2009 in the Python Cave in the Queen Elisabeth National Park. Amman et al. (2013: 6) indicate that at the time of these pulses, the juvenile bats are about 6 months old and are 5 to 6 times more likely to be infected by the virus than the adults (see also Olival and Hayman, 2014). Storm et al. (2018: 1) found that maternal immunity against MARV was lost in juvenile bats by 5 months of age, and that antibodies to MARV remained present in 84 % of naturally exposed bats at least 11 months after capture. Peterson and Holder (2012: 1832) state that R. aegyptiacus is now seen as central in the picture of Marburg virus maintenance, but they also are somewhat confused about the fact that two lineages of Marburg virus are maintained in the same cave or mine in Uganda. Carroll et al. (2012: 2613) indicate that Rousettus aegyptiacus has been identified as a potential natural reservoir for Marburg and Ravn viruses (Marburg marburgvirus). Jones (2015: 3420) experimentally inoculated R. aegyptiacus specimens with Marburg and five Ebola viruses (Sudan, Ebola, Bundibugyo, Taï Forest, and Reston). In the Ebola virus groups, viral RNA tissue distribution was limited, wheras in the Marburg virus group, the virus was widely dissimated. They concluded that this bat species is very unlikely to be a source for Ebola viruses in nature and that there is support for a single filovirus - single reservoir host relationship (here with Marburg virus). Paweska et al. (2018: 1135) tested 1,431 South African bats and found 53.04 % (759) to be positive for antibodies against MARV. These percentages varied between 17.75 % in April 2013 and 82.1 % in October 2013. In adults, these numbers varied between 39.6 % in August 2013 and 100 % in February 2013 (overal 71.56 %), whereas in young bats, they varied between 1.3 % in June 2013 and 77.3 % in January 2014 (overal 30.6 %). An extensive test by Changula et al. (2018: S313S315) showed 158 out of 290 Zambian R. aegyptiacus bats was positive for anti-filovirus antibodies, but no viral RNA genomes were detected. They also found that the seroprevalence peaked in November-December and decreased in January-February. The seroprevalence was also lowest in animals weighing 51 - 60 g, and increased when the bats grew, suggesting that maternal antibodies might have disappeared. Kajihara et al. (2019: 1578), however, were able to extract viral RNA from two Zambian bats, which showed close relationship to the virus found in Uganda. Jones et al. (2019: 1) inoculated Egyptian fruit bats with MARV to investigated the clinical and pathologic effects. They found a mild elevation in alanine aminotransferase (ALT) at 3, 6 and 7 days after inoculation. Nine days after inoculation, Lymphocyte and monocyte counts were slightly elevated and the liver histology revealed small foci of inflammatory infiltrate, similar to lesions previously described in wild, naturally-infected bats. Prescott et al. (2019: 3) suggest that a coevolved strategy exists between the virus and the antiviral respones in R. aegyptiacus. The role of the argasi tick Ornithodoros faini as transmission vector between R. aegyptiacus specimens in Uganda was rejected by Schuh et al. (2016: 1) as none of the 3,125 ticks examined tested positive for the Marburg virus. Pavlovich et al. (2020: 1) investigated the antiviral activity of type I interferons (IFN-ω) in Rousettus aegyptiacus Flaviviridae Flavivirus Uganda S virus- reported in 1947 from Uganda by Butenko (1996). This virus was mentioned by Luis et al. (2013: suppl.). Maganga et al. (2014a: 5) tested 305 animals from Gabon, of which only one tested positive for a Flavivirus. Malmlov et al. (2019: 3) referred to Simpson et al. (1968) who experimentally inoculated three Egyptian fruit bats with Zika virus, of which one became viremic and two seroconverted. Pegivirus (BPgV) Quan et al. (2013: Table S5) found 3 out of 26 Kenyan and 9 out of 91 Nigerian bats examined to be infected with clade K type Pegivirus (overal 10.3 % infection). Herpesviridae Jánoska et al. (2011: 118) reported two viruses, from captive specimens from the Budapest Zoo, belinging to the Betaherpesvirinae and Gammaherpesvirinae. Nairoviridae Orthonairovirus de Jong et al. (2011: 12) and Luis et al. (2013: suppl.) mention the presence of an unassigned Kasokero virus, as well as an unassigned Yogue African Chiroptera Report 2020 virus. The latter was also reported by Kemp (1975: 617) from Senegal in June 1968. Of the 1979 specimens from Gabon tested by Müller et al. (2016: 3), 48 were positive for Crimean Congo hemorrhagic fever virus (CCHFV). Jansen van Vuren et al. (2017: 935) discovered an orthobynyavirus (Wolkberg virus - WBV) in a bat fly (Eucampsipoda africana) collected on a R. aegyptiacus from South Africa. WBV was detected in one serum pool from bats, and at relatively low concentration. Orthomyxoviridae Kandeil et al. (2019: 1) reported a new influenza A virus (IAV) from bats collected in the Nile delta. Orthoreoviridae Harima et al. (2019: 165) found a new Orthoreovirus in a colon swap from a Zambian R. aegyptiacus. Papillomaviridae McKnight et al. (2006: 193) reported some carcinomas near the eye and on wing membranes of a R. aegyptiacus kept in captivity in the USA. These were caused by a putative novel close-toroot papillomavirus, belonging to a new genus. Tse et al. (2012: 7) refer to a Rousettus aegyptiacus papillomavirus type 1 (RaPV1), belonging to the genus Psipapillomavirus (GenBank: DQ366842). Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) reported that three of the 213 specimens (1.4 %) they examined from Gabon and Congo tested positive for Henipavirus. 15 of these bats (7.0 %) were positive for Rubulavirus. Conrardy et al. (2014: 259) found one out of 10 tested animals from the Kitum cave (Kenya) and one of six from the Watamu cave to test positive for Paramyxovirus. Two out of 84 Kenyan R. aegyptiacus specimens tested by Mortlock et al. (2015: 1481) were found to test positive for Paramyxovirus sequences. Amman et al. (2020: 2) reported on Ugandan bats that tested positive for Sosuga virus (SOSV) RNA. They experimentally infected 12 R. aegyptiacus bats and found all of them to be seropositive after 21 days, without showing any overt clinical illness. Orthorubulavirus In South Africa, Mortlock et al. (2019b: 13) found a number of putative Rubula- and related virus RNA in spleen (9.54 %) and urine samples. In October, the virus observations peaked, which coincides with females aggregating in the cave during late stages of gestation. Additionally, high values were also found during December-January, when the bats were lactating. A second excretion peak was found in June-July, probably linked with 175 the juveniles being no longer protected by maternal antibodies. Pararubulavirus Amman et al. (2015a: 776) tested 1331 Ugandan R. aegyptiacus specimens of which 62 (4.6 %) tested positive for Sosuga [pararubula] virus (SOSV) (see also Markotter et al., 2020: 6). Phenuiviridae: Phlebovirus - Balkema-Buschmann et al. (2018: 3) infected three bats from a captive colony with Rift Valley fever phlebovirus (RVFV) and found that viruses could be isolated from the spleens of all three animals and from the liver of one of them, proving that the virus can replicate in the bats. Polyomaviridae: Polyomavirus - Initially, two polyomaviruses were detected from Kenya in 2006 by Tao et al. (2012). Further surveillance identified four more from Kenyan caves (including Watamu cave) where three of the 11 individuals tested positive (Conrardy et al., 2014: 259). Poxviridae: David et al. (2020: 1) discovered a new pox virus in a captive colony at Tel Aviv Zoo: Israel Rousettus aegyptiacus pox virus (IsrRAPXV). Reoviridae: Orbivirus - Fagre et al. (2019: 1) described Bukakata orbivirus (BUKV) from a bat from Uganda. Rotavirus - Sasaki et al. (2018: 106) isolated Group A Rotavirus (RVA) from R. aegyptiacus in Zambia, which was very similar to the strain they collected from Eidolon helvum specimens from that country, suggesting interspecies transmission and genetic reassortment between these bat species. Rhabdoviridae Lyssavirus - Rabies related viruses In Nigeria during June 2008, Quan et al. (2010) did not detect any lyssavirus-specific antigens from the brains by use of direct fluroescent antibody testing. European bat lyssavirus 1 is reported by Van der Poel et al. (2000: 1919), Wellenberg et al. (2002: 349), Omatsu et al. (2008:169), de Jong et al. (2011: 9), Luis et al. (2013: suppl.). Lagos bat virus: Kuzmin et al. (2008a) found a 29 - 46% seroprevalence of Lagos bat virus neutralizing antibodies in Kenya. Kuzmin et al. (2011b: 1467) reported 37 seropositive Kenyan bats on a total of 79 examined (46.8 %). PicardMeyer et al. (2004) isolated the virus from a R. aegyptiacus from an unknown location in Africa. 176 ISSN 1990-6471 Hayman et al. (2011a: 88) and Kuzmin et al. (2011a: 3) refer to a R. aegyptiacus specimen from France, which came originally from either Egypt or Togo. Shimoni bat virus: Kuzmin et al. (2011b: 1467) found 26 bats to be seropositive for this virus on a total of 79 specimens examined in Kenya between 2009 and 2010. Horton et al. (2014: Table S1) tested 54 Kenyan R. aegyptiacus specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). Togaviridae Alphavirus: de Jong et al. (2011: 10) and Luis et al. (2013: suppl.) reported Chikungunya virus occurring on R. aegyptiacus. In their overview table, Maganga et al. (2014a: 8) report the following viruses were already found on R. aegyptiacus: Lagos Bat Virus (LBV), Bat Gammaherpesvirus (1, 2, 4, 5, 6, 7), Bat Gammaherpesvirus 3, Betaherpesvirus, Marburg virus (MBGV), Coronavirus, Zaire Ebola virus (ZEBOV), Yogue virus, Kasokero virus, Chiropteran Papillomavirus, Henipavirus, Rubulavirus, Flavivirus. In their literature review, Fagre and Kading (2019) reported that seriologic evidence was found for the following viruses: Crimean-Congo Hemorrhagic Fever Virus (CCHF), Rift Valley fever virus (RVFV) and Bunyamwera virus (BUNV). The latter virus was also isolated or molecular evidence was found, which was also the case for Kasokero (KKOV) and Yogue (YOGV) viruses. as food and some are transported to markets in other parts of the country to be sold as bushmeat. During the Yam hunting festival in Buoyem, Ghana, fruit bats (possibly R. aegyptiacus were collected (Anti et al., 2015: 1418). ANTHROPOPHILOUS: In Turkey, Albayrak et al. (2008) observed 11 of 15 roosts (73%) were destroyed by human efforts to exterminate the bats over a five-year period. Centeno-Cuadros et al. (2017: 6234) found that, in the Mediterranean area, R. aegyptiacus benefits from human-mediated habitats for daily movements and foraging behaviour. On the other hand, they also found that dispersal (promotor of gene flow between colonies) is apparently not influenced by human-altered habitats. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Burundi, Cameroon, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Egypt, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Malawi, Mali, Mozambique, Nigeria, Rwanda, São Tomé and Principé, Senegal, Sierra Leone, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. Goldberg et al. (2017: 2) and Feng et al. (2017: 99) mention that two viruses were identified from Nycteribiid bat flies (Eucampsipoda africana), which were retrieved from R. aegyptiacus: the putative orthoreovirus Mahlapitsi virus (from South Africa) and the putative orthobunyavirus Wolkberg virus. UTILISATION: Vora et al. (2014: 334) report on an annual bat festival in Idanre, Nigeria, where males of all ages enter caves inhabited by R. aegyptiacus to capture these bats. Many of these animals are prepared Figure 40. Distribution of Rousettus aegyptiacus Rousettus madagascariensis G. Grandidier, 1929 *1929. Rousettus madagascariensis G. Grandidier, Bull. Acad. Malgache, (N.S.) 11: 91, pl. 4 textfigs (for 1928). Type locality: Madagascar: E Madagascar: Beforona: between Tananarive [=Antananrivo] and Andevoranto: Grand Forêt de l'Est. Holotype: MCZ 45432: ad ♂, skull and alcoholic. Collection date: 1917. Presented/Donated by: Guillaume Grandidier. Received from the "Académie Malgache" in 1917; see Bergmans (1994: 119); Peterson et al. (1995: 49); Helgen and McFadden (2001: 141). (Current Combination) African Chiroptera Report 2020 2016. ? ? 177 R[ousettus] madagascar: Lengkong, Arisoesilaningsih, Hakim and Sudarto, Online J. Biol. Sci., 16 (2): 98. Publication date: 14 April 2016. (Lapsus) Rousettus madagascariensis madagascariensis: (Name Combination) Rousettus sp. cf. R. madagascariensis TAXONOMY: Considered a subspecies of lanosus by Hayman and Hill (1971: 12); but see Bergmans (1977a) and Simmons (2005: 348). Bergmans (1994: 119) indicates that the year of publication is often cited as 1928, but Tome XI of the Bulletin de l'Academie Malgache, Nouvelle Série, for 1928, in which it appeared, was published in 1929. Helgen and McFadden (2001: 141) indicate that it might even be published in 1930. COMMON NAMES: Czech: kaloň Grandidierův. English: Madagascan Rousette, Madagascar Rousette. French: Roussette de Madagascar. German: Madagaskar-Höhlenflughund. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Samonds (2007: 57 - 58) found Rousettus sp. cf. R. madagascariensis within the deposits (NCC-1 locality) dating estimated as 69,600 to 86,800 years old within the Anjohibe cave. Samonds (2007: 60) found samples of R. madagascariensis in deposit (SS2) which was collected from the surface near the main cave entrance, the dating attemps revealed this sample was contaminated, therefore no date is known for Anjohibe cave, Madagascar. Gunnell et al. (2014: 3) also refer to Goodman and Junkers (2013), who report on fossil material from the Andrahomana cave in Madagascar. CONSERVATION STATUS: Global Justification This species is listed as Near Threatened (NT ver 3.1 (2001)) as it is believed to have undergone a decline in the region of 20-25% over the past 15 years or so primarily as a result of chronic hunting. Almost qualifies for listing as threatened under criterion A2d. This species is not thought to be declining fast enough to place it in a category of higher threat, although a review of this assessment may be warranted in future, especially with further information on the degree to which it may be impacted by habitat loss (Andriafidison et al., 2008q; IUCN, 2009). Assessment History Global 2008: NT ver 3.1 (2001) (Andriafidison et al., 2008q; IUCN, 2009). 2000: LR/nt ver 2.3 (1994). 1996: VU ver 2.3 (1994). Regional None known. MAJOR THREATS: Andriafidison et al. (2008q) [in IUCN (2009)] report that the extent to which the destruction and degradation of natural forest threatens this species is poorly understood, while Andrianaivoarivelo et al. (2011b: 81) indicate that this is an indirect threat. Although Goodman et al. (2005a) suggested that R. madagascariensis is not dependent on relatively intact forest there are insufficient data on its annual dietary requirements to decide this matter. It is clear, however, that some of the roosting colonies are located some distance from intact forest and additional study is therefore needed on the mobility of this species. The main threats to R. madagascariensis are to its roosts where it is subject to hunting pressure (Jenkins et al., 2007a; Rakotonandrasana and Goodman, 2007; Jenkins and Racey, 2008; Andrianaivoarivelo et al., 2011b: 78, 81) in virtually all sites that are not inside protected areas (Goodman et al., 2005a) or considered sacred (Rakotoarivelo and Randrianandrianina, 2007). Jenkins et al. (2011: 1), however, indicate that the taboo on hunting in sacred areas is rapidly eroding. It appears to be hunted exclusively for subsistence and bats are harvested using locally made traps as well as being knocked down from the cave ceiling with wooden batons (Rakotonandrasana and Goodman, 2007). R. madagascariensis can legally be hunted between the first of May and the first of September (Anonymus, 2007b: 6). On Ile Sainte Marie the reported offtake was between 360 and 480 bats per year (Rakotonandrasana and Goodman, 2007). Jenkins et al. (2007b: 213) point out that R. madagascariensis is very vulnerable to hunters because it roosts in caves, which can easily be covered by a trap system. They mention that one local farmer was able to capture 170 bats in a single night. Cardiff et al. (2012: 479) investigated the influence of tourist disturbance on roosting R. madagascariensis. They found that the bats were slightly less disturbed when ambient light was present. Directly illuminating the bats resulted in higher disturbance. Also in roosts which were previously visited by hunters, the bats were more easily disturbed. They suggest keeping a 178 ISSN 1990-6471 minimum visit distance of 12 m, not illuminating the bats directly, and not opening other roost sites for tourism to limit the disturbance of the bats. ♂♂: Fa: 75.3 ± 2.00 (70 - 80) mm, Weight: 64.6 ± 6.85 (48.0 - 81.5) g; ♀♀: Fa: 73.0 ± 2.04 (66 - 80) mm, Weight: 56.8 ± 6.54 (31.0 - 75.5) g. CONSERVATION ACTIONS: Andriafidison et al. (2008q) [in IUCN (2009)] report that, as a game species under Malagasy law (Durbin, 2007), R. madagascariensis is only protected when it occurs in nature reserves. In a survey of western Madagascar, it was found in six protected areas: Réserve Spéciale d'Ankarana, Réserve Spéciale d'Analamerana, Parc National d'Ankarafantsika, Parc National de Namoroka, Parc National du Tsingy de Bemaraha and Parc National d'Isalo (Goodman et al., 2005a). Roosts within existing protected areas (only three of which are known - Cardiff and Jenkins, 2016: 99) need to receive close attention from park staff to discourage hunting. Other roosts need to be conserved and this might be best achieved through their inclusion within new protected areas and with the cooperation of local communities. MOLECULAR BIOLOGY: DNA - Goodman et al. (2010b) examined Cyt-b from 131 individuals from 17 sites across Madagascar. Andrianaivoarivelo et al. (2008: 1025) isolated 22 nuclear microsatellite loci, which will provide valuable information for population genetic studies. GENERAL DISTRIBUTION: Rousettus madagascariensis is endemic to the island of Madagascar where it is widespread but rare or absent from the central highlands and the arid south-west (MacKinnon et al., 2003; Goodman et al., 2005a). It has also been recorded from the off-shore islands of Nosy Be, Nosy Komba, and Ile Sainte-Marie (Rakotonandrasana and Goodman, 2007: 6). Native: Madagascar (MacKinnon et al., 2003; Goodman et al., 2005a; Rakotonandrasana and Goodman, 2007: 6). BIOGEOGRAPHY: Goodman et al. (2010b: 600) did not find any clear phylogeographic structure for populations of R. madagascariensis on Madagascar. Across the 1,600 km length of Madagascar individuals of R. madagascariensis from extreme ends of the island and across numerous biomes, and nearshore islands up to 13 km from the island, demonstrated complete genetic mixing. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 55) indicate that the baculum has a rounded distal tip and a distinctly narrower proximal base; length: 2.25 ± 0.228 (1.96 - 2.57) mm, width: 0.98 ± 0.179 (0.76 - 1.19) mm. SEXUAL DIMORPHISM: Goodman et al. (2017b: 72) measured 271 adult males and 289 adult females, and found males to be significantly larger in both forearm length and body mass, although with a considerable overlap: Karyotype - 2n = 36 and FN = 66, containing 7 large metacentrics, 4 medium-sized submetacentrics, 5 small metacentrics and one acrocentric pair; X: large submetacentric; Y: smallest chromosome, largely heterochromatic (see Richards et al., 2016d: 187, 189). Protein / allozyme - Unknown. HABITAT: Due to its small size, Andrianaivoarivelo et al. (2011a: 70) suggest that it is potentially an important seed disperser and pollinator inside forests, since the two other Madagascan fruit bat species are rather large and unable to fly within dense foliage. It was also found to forage in relatively intact forest and highly modified habitats close to villages (see Andrianaivoarivelo et al., 2011a: 76), although this contradicts observations by Randrianandriananina et al. (2006), who did not find any R. madagascariensis specimens in relatively intact humid forest. Dammhahn and Goodman (2013: 108) indicate that mid- to upper-canopy in forest and trees growing in open areas form the foraging habitat of this species. HABITS: Using radio tracking and light-tagged individuals, Andrianaivoarivelo et al. (2011a: 69, 73) found that these bats travelled over 8 km (up to 14.2 km for females and 14.8 km for males) from their roost sites to their feeding grounds, and that round trips might be as far as 27 km. Goodman (1999: 253) and Andrianaivoarivelo et al. (2011a: 73) report that they only grasp a ripe fruit in their mouths and then fly off to consume it. ROOST: Rousettus madagascariensis roosts in caves and although it is widespread, relatively few roosts are known (MacKinnon et al., 2003). Even though some roosts are protected from hunters because they occur in protected areas (Kofoky et al., 2007) or sacred sites (Rakotoarivelo and Randrianandrianina, 2007), there are many African Chiroptera Report 2020 reports of hunting in and around caves (Rakotonandrasana and Goodman, 2007; Jenkins et al., 2007b). Ramanantsalama et al. (2019: 115) examined the seasonal behavioral activity in a cave day roost, and found that the behavior changed according to season. During the dry season, bats formed 'tight' groups, and spent more time in 'rest' and 'consume ectoparasites' behaviors and limited 'move' (flight and crawl) and 'groom' activities. During the wet season, 'groom' behavior frequency was higher and individuals formed a more dispersed roosting group configuration. The thresholds for the shift were the average temperature outside the cave, the mean temperature in the cave and the average relative humidity. If these were respectively = 25.3°C, = 24.0°C and = 68.2%, 'tight' groups were formed. When the values were respectively = 27.1°C, = 25.0°C and = 93.4%, the bats changed to a 'loose' group configuration. DIET: Goodman (1999: 253), Andrianaivoarivelo et al. (2007: 3) and Andrianaivoarivelo et al. (2011a: 69) reported that this species extensively feeds on the fruits of banana (Musa paradisiaca, family Musaceae), litchi (Litchi chinensis, family Sapindaceae), and Dimocarpus longan (Sapindaceae). Jenkins et al. (2007b: 213) [referring to Rakotonirainy, 2001] mention these bats visiting Ficus soroceoides, F. pyrifolia, and F. guatteriifolia. However, Andrianaivoarivelo et al. (2012: 19) indicate that in flight cage experiments, R. madagascariensis preferred native or introduced fruit of no commercial value (Ficus polita, Syzygium jambos and S. malaccense (Myrtaceae)) to commercially important fruits (Litchi chinensis and Diospyros kaki). Faecal analysis performed by Nogales et al. (2006: 69) indicated that small amounts of seeds were present, which suggests that these bats either feed on nectar, flowers and leaves or on fruits with seeds too large to swallow. They did find Ficus seeds in the faeces during the months of March and April. Jenkins (2007: 17) reported the following plant species being found in faecal samples: Ficus soroceoides, Ficus botryoïdes, Ficus pyrifolia, Ficus brachyclada and Rubus mollucanus. Seeds less than 2.5 mm in length were swallowed. The seeds that passed through the bat's gut germinated faster than non-ingested seeds. Andrianaivoarivelo et al. (2011a: 75) also indicate that seeds of Ficus rubra, which have passed 179 through the digestive tract of R. madagascariensis germinate faster than those not passing through it. Ramanantsalama et al. (2018: 6) reported that adult R. madagascariensis consumed 37 ± 16 (range 2 - 67) ectoparasites (Streblidae and Nycteribiidae) per day while grooming, whereas juveniles did not ingest any parasites. However, they do admit that this number is an overestimation as they refer to Rajemison et al. (2017b), who reported an average of 6.6 bat flies per adult. PREDATORS: In one locality on Madagascar (near Antonibe), Goodman and Griffiths (2006: 555) found that 81.3% of the total biomass predated by the barn owl (Tyto alba) consisted of R. madagascariensis. POPULATION: Structure and Density:- It roosts in aggregations of up over a thousand individuals (MacKinnon et al., 2003; Rakotoarivelo and Randrianandrianina, 2007), but smaller colonies of a few hundred are also known (Jenkins et al., 2007a; Kofoky et al., 2007; Andrianaivoarivelo et al., 2011b: 80). It can be locally abundant and is often the most commonly trapped species in mist-netting surveys in the west of Madagascar (Kofoky et al., 2007; Rakotoarivelo and Randrianandrianina, 2007). For the "Grotte des Chauves-souris, Ankarana", Noroalintseheno Lalarivoniaina et al. (2017: 72) estimated the population size to be between 675 and 4000 (IC 95 %, between September 2014 and September 2015). For the same population, Noroalintseheno Lalarivoniaina et al. (2018: 2627) found that in adults there was a male bias during the dry season and a female bias during the wet season, whereas this was the inverse for juvenile bats. Adults were also present in higher numbers during the dry season than during the wet season. Trend:- 2008: Decreasing (Andriafidison et al., 2008q; IUCN, 2009; Andrianaivoarivelo et al., 2011b: 80). LIFESPAN: Noroalintseheno Lalarivoniaina et al. (2017: 73) found that the survival rate of males tended to exceed that of females. This was confirmed for adults (♂♂: 0.50 / ♀♀: .047) by Noroalintseheno Lalarivoniaina et al. (2019: 103), who also found that the inverse was the case for subadults (♂♂: 0.55 / ♀♀: .070). REPRODUCTION AND ONTOGENY: Andrianaivoarivelo et al. (2011a: 69, 73) report that most juveniles were captured between March and July, and they suspect that young are weaned 180 ISSN 1990-6471 before 8 weeks of age. Pregnancy (OctoberDecember), lactation (December-January) and weaning occurs in the austral summer, when food is readily available. Noroalintseheno Lalarivoniaina et al. (2017: 69) indicate that mating occurs in the dry season and births follow in the wet season. This was also confirmed by Noroalintseheno Lalarivoniaina et al. (2018: 26), who only found neonates during the wet season, with an unbiased sex ratio. MATING: Goodman et al. (2017b: 74) indicate that this species probably has a diffused polygamous mating system, with male-male competition for access to females. POSTNATAL DEVELOPMENT: Andrianaivoarivelo et al. (2011a: 69) were unable to distinguish juveniles from adults in December, and therefore assume that R. madagascariensis reaches somatic maturity within one year. PARASITES: BACTERIA Gomard et al. (2016: 5) found 15 out of the 49 bats they tested to be positive for Leptospira bacteria. DIPTERA Tortosa et al. (2013: 3), Wilkinson et al. (2016), Rajemison et al. (2017a; 2017c: 805) and Ramasindrazana et al. (2017: Suppl.) report the presence of the Nycteribiid bat fly Eucampsipoda madagascarensis Theodor, 1955. Rajemison et al. (2017b: 72; Rajemison et al. (2017c: 805) reported a second bat fly: Megastrebla wenzeli (Jobling, 1952), belonging to the Streblidae. They found that the number of E. madagascarensis flies per bat (up to 27) varied by season and was lowest during the rainy season, and that the sex ratio for this fly was biased towards males (especially during the dry season). As for the bats, adult males were more infected by E. madagascarensis than females, subadults and neonates. No such variation was observed for M. wenzeli. Szentiványi et al. (2019: Suppl.) indicated that the E. madagascarensis bat flies were found to carry Enterobacteriales and Rickettsia bacteria. Rajemison et al. (2018: 514) studied the reproductive biology of the bat fly Eucampsipoda madagascarensis in northern Madagascar, and found that during the bat mating period (dry season - September), pregnant bat flies did not show any preference for hosts of both sexes that were in reproductive or non-reproductive state, except that males with scrotal testes were more parasitized by gravid flies than non-scrotal males. During the lactation period (wet season - January), gravid flies were excessively present on lactating females. SIPHONAPTERA Hastriter (2016: 15) reported the collection of a male flea (Araeopsylla martialis (Rotschild, 1903)) from a Rousettus madagascariensis, but believes this to be an accidental association. VIRUSES: Astroviridae Two out of 41 by Lebarbenchon et al. (2017: Suppl.) tested bats were positive for Astroviruses. Coronaviridae Joffrin et al. (2020: 7) reported Beta-D coronavirus from these bats. Filoviridae Ebola virus - Brook et al. (2019b: 1004) reported the presence of reliable reactive antibodies to tested antigens in serum from R. madagascariensis. However, they also mention (p. 1010) that some of the seropositives recovered in their study result from cross-reactivity of Malagasy bat antibodies to related, but distinct, antigens from those assayed by them. Paramyxoviridae Henipavirus Cedar (CedPV) - Brook et al. (2019b: 1004) reported the presence of reliable reactive antibodies against CedPV in the serum. Hendra (HeV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) were unable to detect the virus. Brook et al. (2019b: 1004), however, were able to find antibodies against this virus in the serum. Nipah (NiV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) could not detect the virus. Kuzmin et al. (2011a: 5) suggest that their exposure to NiV might be the result of contact with infected Pteropus rufus specimens. Tioman (TiV) - In Madagascar during 2003 - 2004, Iehlé et al. (2007) found an overall contamination of 20.0 %. Rhabdoviridae Lyssavirus Lagos bat virus (LBV) - A sero reaction was found by Mélade et al. (2016a: 6) in seven out of 35 bats tested. Duvenhage lyssavirus (DUVV) - Four out of 33 bats tested positive in the same study. UTILISATION: Golden (2005) reported large annual harvests in the Makira forest and Maroantsetra were unsustainable. See Jenkins et al. (2007b) and Goodman et al. (2008d). African Chiroptera Report 2020 181 Golden and Comaroff (2015: 5) indicate that flying foxes and other fruit bats (including R. madagascariensis) are frequently the subject of food taboos, which they link to the fact that these bats are carriers of virulent zoonotic diseases. ANTHROPOPHILOUS: Cardiff et al. (2012) suggests that a minimum visit distance of 12 m should be maintained and not illuminating the bats directly at tourist caves, as well as not opening other roost sites to tourism, is likely to help to limit disturbance of R. madagascariensis by tourists at Ankarana. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar. Figure 41. Distribution of Rousettus madagascariensis Rousettus obliviosus Kock, 1978 *1978. Rousettus (Rousettus) obliviosus Kock, in: Olembo, Castelino, and Mutere, Proceedings of the 4th International Bat Research Conference Nairobi, 208. Type locality: Comoros: Grand Comoro: Boboni, cave near [ca. 11 35 S 43 20 E, 640 m]. Holotype: ZFMK MAM1958.0207:. Presented/Donated by: ?: Collector Unknown. Holotype: ZMB 58207: ad ♂. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 3 August 1903; original number: 60. See Turni and Kock (2008: 21). Paratype: SMF 47973: ad ♂, skull and alcoholic. Formerly ZMB 58201; see Turni and Kock (2008: 21). Paratype: ZMA MAM.20903: sad ♀, skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 10-11 October 1903; original number: 94. From Anjouan [= Nzwani, Comoro Ids.]. Formerly ZMB 58196, exchanged in 1979, see Turni and Kock (2008: 21), Bergmans (2011: 842). Paratype: ZMA MAM.20906: imm ♀, skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 11 October 1903. Presented/Donated by: ?: Collector Unknown. Paratype: ZMB 4908: skull only. Collected by: Dr. Johann Maria Hildebrandt; collection date: 3 August 1903. From Anjouan [Nzwami, Comoro Isls.]; see Turni and Kock (2008: 21). Paratype: ZMB 58189: skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 10-11 October 1903; original number: 91. From Cercle de Bambao, Anjouan [= Nzwani, Comoro Ids.]; see Turni and Kock (2008: 21). Paratype: ZMB 58197: sad ♀. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 10-11 October 1903; original number: 88 ?. From Anjouan [= Nzwani, Comoro Ids.]; see Turni and Kock (2008: 21). Paratype: ZMB 58198: skin and skull. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 10-11 October 1903. From Cercle de Bambao, Anjouan [= Nzwani, Comoro Ids.]; see Turni and Kock (2008: 21). Paratype: ZMB 58199: sad ♀. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 10-11 October 1903; original number: 88 ?. From Anjouan [= Nzwani, Comoro Ids.]; see Turni and Kock (2008: 21). Paratype: ZMB 58200: skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 10-11 October 1903; original number: 97. From Cercle de Bambao, Anjouan [= Nzwani, Comoro Ids.]; see Turni and Kock (2008: 21). Paratype: ZMB 58202: ad ♀, alcoholic (skull not removed). Collected by: Prof. Dr. Alfred Voeltzkow. From Boboni cave, Grand Comoro; see Turni and Kock (2008: 21). Paratype: ZMB 58203: skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 21 December 1903. From Grand Comoro; see Turni and Kock (2008: 21). Paratype: ZMB 58204: skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 21 December 1903. From Grand Comoro; see Turni and Kock (2008: 21). Paratype: ZMB 58205: skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 21 December 1903. From Grand Comoro; see Turni and Kock (2008: 21). Paratype: ZMB 58206: skull and alcoholic. Collected by: Prof. Dr. Alfred Voeltzkow; collection date: 21 December 1903. From Grand Comoro; see Turni and Kock (2008: 21). ? Rousettus obliviosus: (Name Combination, Current Combination) 182 ISSN 1990-6471 TAXONOMY: Considered a valid species by Meirte (1987), Koopman (1993a: 153), Bergmans (1994: 79), Wilson and Cole (2000: 40), and Simmons (2005: 348), but as a subspecies of madagascariensis by Peterson et al. (1995: 48). Goodman et al., 2010b: 599) shows that R. obliviosus is a sister taxa to R. madagascariensis with a 6.2-8.7 % sequence divergence, and clearly form a reciprocally monophyletic clades. COMMON NAMES: Czech: kaloň Kockův. English: Comoro Rousette. French: Roussette des Comores, la petite roussette des Comores. German: Komoren-Höhlenflughund. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B1ab(iii)+2ab(iii) ver 3.1 (2001)) in view of the species has a limited range; both its extent of occurrence and area of occupancy are less than thresholds of 20,000 km 2 and 2,000 km2, respectively (the minimum convex polygon including all three islands it inhabits is 9,085 km2, and the total area of all three islands it inhabits is 1,783 km 2). It is also known to exist at no more than ten locations (seven consistentlyused roosting caves are known), and it is experiencing a continuing decline in the extent and quality of its habitat (due to increasing human pressures leading to substantial loss of native forests; ongoing loss of trees from plantations, orchards, and agroforestry areas; and increasing human disturbance at roost sites) (Sewall, 2016). Assessment History Global 2016: VU B1ab(iii)+2ab(iii) ver 3.1 (2001) (Sewall, 2016). 2008: VU B1ab(iii)+2ab(iii) ver 3.1 (2001) (Mickleburgh et al., 2008v; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004cr; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Sewall (2016) reported that Rousettus obliviosus has a limited range; its area of occupancy is less than the total area of the three islands (Sewall et al., 2003), which is 1,783 km 2 (Louette, 2004), and its extent of occurrence (including unsuitable ocean areas between the islands) is less than 9,085 kmsuper 2. It is therefore susceptible to single threatening processes, like cyclones, that could simultaneously or rapidly affect its entire range. The major threat to the species within this range appears to be disturbance at roost sites. The bats appear highly susceptible to human disturbance, and infrequent hunting and rock falls in unstable lava tubes have been recorded at roost sites (Middleton, 1998b; 1999; Sewall et al., 2003). Additionally, cave surveys between 2000-2005 revealed the bats were absent from three caves where they apparently formerly occurred (Sewall et al., 2003, W. Masefield, pers. comm. in Sewall (2016)), though it is unclear whether this suggests intermittent use of some roost sites or abandonment after disturbance. Over the last few decades, the Comoros have undergone a sustained, rapid deforestation, resulting in the loss of nearly all its native forests. During the past 20 years alone, the country has lost 75% of its remaining forests, the fastest rate of any country in the world (UNDP 2013). Human pressures on native forests are expected to continue, as the growth rate of the human population remains high at 2.5% (UNDP 2013). Because R. obliviosus is regularly recorded in human-modified landscapes, loss of foraging habitat via deforestation is possibly not a major threat (Bergmans, 1994). However, even humanmodified landscapes containing non-native and remnant native trees are disappearing in the Comoros to make room for ground crops (FAO 2010); thus food resources for this species have probably been declining in recent years. Further, the loss of natural and underplanted forests may reduce its ability to cope with droughts and cyclones, and may limit its access to food resources when fruit from horticultural species is unavailable (Sewall et al., 2003). In addition, the replacement of native forests with agricultural land on Anjouan and Mohéli may render formerly inaccessible roost sites on Anjouan and Mohéli more easily reached, possibly exposing roosts to increased levels of human disturbance. Finally, the loss of native forests has resulted in impermanence or complete drying of nearly all rivers on Anjouan and most on Mohéli (Louette, 2004). R. obliviosus roosts on those islands are typically near waterfalls, and it is unclear whether that is simply where suitable caves and rock shelters occur or if the bats benefit from the adjacent water source. The species could benefit from an adjacent water source if, for instance, local humidity reduces evaporative water loss in the normally hot climate of the Comoros. If so, then it is possible the seasonal or year-round loss of African Chiroptera Report 2020 water flow could affect this species on Anjouan and Mohéli as well. CONSERVATION ACTIONS: Sewall (2016) reports that Rousettus obliviosus receives the highest level of legal protection available in the Union of the Comoros. It is listed as an ‘integrally-protected species’ (list 1 of RFIC 2001), which prohibits the capture or detention of R. obliviosus individuals without a permit. This law also expressly prohibits the killing of members of the Family Pteropodidae (flying foxes); transport, purchase, sale, export or re-export of living or dead individuals or their body parts; all disruption during the period of reproduction and raising of young; and the destruction of roosts (RFIC 2001). The Union of the Comoros also ratified the Convention on Biological Diversity in 1994, and in response has developed a National Biodiversity Conservation Strategy (Roby and Dossar, 2000). This strategy highlights the importance of, threats to, and conservation recommendations for fruit bats of the Union of the Comoros (Sewall and Granek, 2000). In practice, however, there are currently no direct conservation actions in place for R. obliviosus, and no formal terrestrial protected areas in the country. The national Conservation Action Plan for Pteropus livingstonii (Sewall et al., 2007), another threatened pteropodid from the Comoros, includes an appendix for the conservation of Rousettus obliviosus. This appendix addresses current and potential emerging threats to R. obliviosus, and recommends five targeted conservation actions for R. obliviosus. Specifically, it recommends (1) protection of roost caves, especially the caves with the largest known colony on each island (these are now thought to be Panga Chilamouinani near Fassi on Grande Comore, Bakomdrundru near Ndrondroni on Mohéli, and Mangua Mitsano near Limbi on Anjouan); (2) discouraging, through environmental education, the hunting of all fruit bats; (3) conducting a comprehensive field search for more roost sites; (4) providing incentives to individuals owning land near roost sites to conserve caves and bats; and (5) devising a suitable population monitoring protocol and conducting regular visits to all known roost sites to track population changes (Sewall et al., 2007). Other key actions for this species include efforts to prevent further native forest habitat loss and additional studies designed to track the species’ population and inform its management (Sewall et al., 2003). GENERAL DISTRIBUTION: Rousettus obliviosus is restricted to the lower elevations of the islands of Grande Comores, Moheli and Ajouan in the Comoros, western Indian 183 Ocean. The species has not been reported on Mayotte Island (Louette, 1999; Sewall et al., 2003; Goodman et al., 2010c: 124); Goodman et al. (2010c: 125) suggest that the reason for the absence from Mayotte island is that it is geologically the oldest (Emerick and Duncan, 1982) and the former lava tubes - important day roost sites for this species elsewhere in the archipelago - have collapsed (Goodman et al., 2010c: 125). On Grande Comore it has an elevational range from near 30 to 1,750 m, on Anjouan from 200 to 1,350 m, and on Moheli from 10 to 700 m (Sewall et al., 2003; Goodman et al., 2010c: 125). ZMB 3891 refers to "Caffaria", which has been interpreted as South Africa, or southern Africa. This specimen needs to be examined to confirm if it is infact E. obliviosus and not a mainland Epomophorus. Native: Comoros (Mickleburgh et al., 1992; Carroll and Feistner, 1996; Sewall et al., 2003; Goodman et al., 2010c: 124). BIOGEOGRAPHY: Goodman et al. (2010b: 600) did not find any clear phylogeographic structure for populations of R. obliviosus on Grand Comore, Anjouan and Moheli, with open-water distances separating these islands between 40 and 80 km. ECHOLOCATION: Although Reason et al. (1994: 402) did not see flying R. obliviosus before dusk, it is unknown whether this species is able to echolocate. MOLECULAR BIOLOGY: DNA - Goodman et al. (2010b) examined Cyt-b from 44 individuals from three islands in the Comoro island (Grande Comore, Anjouan, and Moheli). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: R. obliviosus has been found roosting in forest caves near waterfalls (Anonymus, 2000: 30) and it feeds in the Comoron rain forests. Forages in agricultural zones, degraded or largely intactforested areas, and understory plantations (Goodman et al., 2010c: 125). A number of cave roost sites are at considerable distances from native forests (Goodman et al., 2010c: 125). HABITS: R. obliviosus is strictly nocturnal, as Goodman et al. (2010c: 125) recorded exit times commenced on the 16 November 2006 at 18h45, after complete 184 ISSN 1990-6471 darkness at a cave roost at Nyamaoui Panga (Grand Comore). ROOST: Roosts in lava tube caves or difficult to access rock crevices (Goodman et al., 2010c: 125). Reason et al. (1994) suggested that it uses tree hollows for day roost sites. However, there is no evidence for this (Goodman et al., 2010c: 125). Wilkinson et al. (2012: 160) indicate that caves are typical roosting sites for this species on the Comoros. Middleton and Hume (2010: 137) found a colony of a few thousand animals in a large overhang 12.4 m high, 15 m deep and about 35 m long, beside a waterfall on Anjouan Island. DIET: Young et al. (1993) [in Carroll and Feistner (1996: 330)] indicate that R. obliviosus and P. livingstonii were captured at the same kapok and fig feeding sites on Anjouan, which would suggest that their feeding niches might overlap. However, they also indicate that there might be a vertical separation between the two species. Based on direct observations and food plants recovered from ejecta, R. obliviosus commonly feeds on several species of introduced fruiting plants (Mickleburgh et al., 1992; Sewall et al., 2003). Goodman et al. (2010c: 125) list Ceiba pentandra [= Kapok tree] (Malvaceae), Musa spp. [= Banana] (Musaceae), Artocarpus integrifolia [= Jackfruit] (Moraceae), and Carica papaya [= Papaya] (Caricaceae). Most of these are introduced plants which are abundant in agricultural areas with degraded forest. Ratrimomanarivo (2007) characterized R. obliviosus as a forest dependent species, which is not supported by Goodman et al. (2010c: 125). POPULATION: Structure and Density:- Sewall (2016) reports that Rousettus obliviosus is rarely observed outside of research studies due to its nocturnal foraging and cryptic roosting habits. It has nonetheless been captured frequently by bat researchers (Reason et al., 1994, Clark et al., 1997, Sewall et al., 2003, Goodman et al., 2010c), and thus the species is considered fairly common (Reason et al., 1994, Louette, 2004, Goodman et al., 2010c). The true population size is unknown. The only abundance estimates available are from roost counts, and very few roost sites are known. Sewall et al. (2003) searched Grande Comore, Anjouan, and Mohéli and found or confirmed five permanent roosts plus a sixth that was intermittently used. They estimated that colony size at roosts ranges from around 100 to several thousand animals, and that the total population of these six sites was between 7,100 and 17,100 bats (Sewall et al., 2003). Since then, two additional roosts have been discovered, one small (~100 bats) and one mid-sized (~2,000 bats) (Hume and Middleton, 2011, I. Saïd and W. Masefield pers. comm. in Sewall (2016)). Thus, the current population at known roosts is estimated to number about 10,000 to 20,000 bats. Estimates of demographic parameters from genetic analyses suggest recent population stability in this species (Goodman et al., 2010b). Further, no genetic population structure is evident among the three islands, despite separation of at least 40 km of open water between adjacent islands; this indicates inter-island dispersal (Goodman et al., 2010b). Reason et al. (1994: 399) reported a female sex ratio bias on Anjouan, which was not statistically significant. Trend:- 2016: Unknown (Sewall, 2016). 2008: Decreasing (Mickleburgh et al., 2008v; IUCN, 2009). REPRODUCTION AND ONTOGENY: Reason et al. (1994: 125) noted that during the months of July and August 1992 on Anjouan, nonreproductive subadults were captured, estimated to be between 3 and 12 months old. Similarly, Carroll and Thorpe (1991) on Anjouan late July 1990 captured four parous adult females and two non-parous adult females; not one showed signs of reproductive activity. On the 16 November, Goodman et al. (2010c: 125) visited Nyamaoui Panga on Grande Comore, where among the 14 free-flying adults captured, seven were male, one of which had scrotal testes measuring 8 x 4 mm with partially convoluted epididymes, and the remaining six were not in reproductive condition. Amongst the seven females, two were pregnant, with single embryos measuring 2 - 31 mm in crown rump length, four lactating, and one nonreproductive. Goodman et al. (2010c: 125) revisited the site on the 6 December 2006 and amongst the six individuals caught, one male had abdominal testes, one was a female that had previously reproduced and showed no sign of active reproduction, and the balance were lactating females with single placental scars. This indicates that individuals at different reproductive stages during the same period, and there also appears to be notable inter-island synchronization in the reproductive season (Goodman et al., 2010c: 125). The birthing period would seem to coincide with the wet season, October to April. Reason et al. (1994: 401) suggest that females can reach sexual maturity in their first year (earlier than males). African Chiroptera Report 2020 PARASITES: R. obliviosus from the Comoros tested positive for Leptospira bacteria (Lagadec et al., 2012: 1696). Dietrich et al. (2018a: 3) identified these further as L. interrogans and L. borgpetersenii. Lei and Olival (2014: Suppl.) mention that the species is also a host for Bartonella bacteria. 185 UTILISATION: Infrequent hunting has been recorded (Sewall, 2016). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, South Africa. Tortosa et al. (2013: 3) and Wilkinson et al. (2016) report the presence of the Nycteribiid bat fly Eucampsipoda theodori. These bat files were found to carry Enterobacteriales and Bartonella bacteria (Szentiványi et al., 2019: Suppl.). Reason et al. (1994: 399) collected Brachytarsia sp. (possibly B. wenzeli [=?Megastrebla wenzeli]). VIRUSES: Paramyxoviridae Wilkinson et al. (2012: 160) tested 26 individuals from the Comoros using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Figure 42. Distribution of Rousettus obliviosus TRIBE Scotonycterini Bergmans, 1997 *1997. Scotonycterini Bergmans, Beaufortia, 47 (2): 11, 69. Publication date: 20June 1997. TAXONOMY: Includes the genera Casinycteris Thomas, 1910 and Scotonycteris Matschi, 1894 (Almeida et al., 2020: 11). Genus Casinycteris Thomas, 1910 *1910. Casinycteris Thomas, Ann. Mag. nat. Hist., ser. 8, 6 (31): 111. Publication date: 1 July 1910 [Goto Description]. - Comments: Type species: Casinycteris argynnis Thomas, 1910. (Current Combination) TAXONOMY: Revised by Bergmans (1991). Included in the Scotonycterini tribe by Bergmans (1997: 69), Almeida et al. (2016: 83) and Hassanin et al. (2020: 5), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter two. Currently (Simmons and Cirranello, 2020) recognized species of Casinycteris: argynnis Thomas, 1910; campomaanensis Hassanin, 2014; ophiodon (Pohle, 1943). COMMON NAMES: Czech: krátkonebí kaloni. English: Short-palate Fruit-bats, Short-palated Fruit-bats. French: Casinyctère. German: Kurzgaumen-Flughunde. Casinycteris argynnis Thomas, 1910 *1910. Casinycteris argynnis Thomas, Ann. Mag. nat. Hist., ser. 8, 6 (31): 111. Publication date: 1 July 1910. Type locality: Cameroon: Ja River: Bitye [03 13 N 12 22 E, 2 000 ft] [Goto 186 ISSN 1990-6471 1943. Description]. Holotype: BMNH 1911.5.5.1: ad ♀, skin and skull. Collected by: George Latimer Bates Esq. Collection date: 19 November 1909; original number: 502. (Current Combination) [Scotonycteris] argynnis: Pohle, Sber. Ges. naturf. Freunde Berlin, 86 (for 1942). (Name Combination) TAXONOMY: See Bergmans (1991) and Simmons (2005: 315 316). COMMON NAMES: Czech: kaloň krátkonebý. English: Golden Shortpalate Fruit-bat, Yellow-winged Short-palate Fruitbat, Thomas's Short-palate Fruit-bat, Shortpalated Fruit Bat, Golden Short-palated Fruit Bat. French: Casinyctère dorée, Roussette à petit palais, Chauve-souris frugivore à palais court, Casinyctère. German: Goldener KurzgaumenFlughund. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008c; IUCN, 2009, Webala et al., 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Webala et al., 2016). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008c; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004k; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Mickleburgh et al. (1992) indicate that there is little information on the threats to this species, but that it may be threatened by deforestation. It is possible that this species eats cultivated fruits, and consequently might be persecuted as a crop pest (Mickleburgh et al., 2008c; IUCN, 2009). CONSERVATION ACTIONS: Current conservation efforts There appear to be no direct conservation measures in place (Mickleburgh et al., 2008c, in IUCN (2009), although in the Democratic Republic of Congo the species is present in some protected areas such as Mbiye island sanctuary, Yangambi Man and Biosphere Reserve, Yoko Forest Reserve and Masako Forest Reserve (Webala et al., 2016). Conservation needs/priorities Webala et al. (2016):Studies are needed on the species’ population sizes, distribution, and extent of occurrence throughout its range. Monitoring of population sizes and locations over time are also important to establish whether these are stable or experiencing trends of decline. The threats to these bats are poorly understood. Studies are needed on the species’ habitat requirements and on the effects of forest loss and degradation on the species’ population sizes/distribution. Research is also needed on the amount of hunting and the level of bushmeat trade, and the effects of that hunting on population sizes and persistence. Effective roost site protection efforts are needed to minimize hunting mortality and disturbance to nontarget individuals. Similar to most threatened flying foxes, local capacity building for conservation managers and education and awareness within local communities are greatly needed to support conservation efforts. Mickleburgh et al. (1992) recommend that surveys are needed to assess the conservation status of this species, to determine if it is present in protected areas, and to learn more about the food plants and general natural history of this species. GENERAL DISTRIBUTION: This lowland species is distributed in Central Africa. It has been recorded from southern Cameroon, the Central African Republic and the Democratic Republic of the Congo. Native: Cameroon; Central African Republic; Congo (The Democratic Republic of the) (Bergmans, 1991; Lunde et al., 2001: 537; Monadjem et al., 2010d: 550); Equatorial Guinea; Gabon (Drexler et al., 2012a: Suppl. Table S1). Presence uncertain: Congo. DETAILED MORPHOLOGY: Chawana et al. (2013: 160) report an average brain mass of 0.83 g (n = 2). FUNCTIONAL MORPHOLOGY: The eye of this bat contains about 130,000 retinal ganglion cells (Coimbra et al., 2016: 191). The minimum angle of resolution was about 0.227° (ca. 8 mm at 1 m distance). African Chiroptera Report 2020 187 HABITAT: This species is generally associated with lowland tropical moist forest, swamp forest, a mosaic of these two forest habitats, and a mosaic of swamp forest and secondary grassland (Bergmans, 1991). of the Congo (Mickleburgh et al., 2008c; IUCN, 2009). ROOST: It is a solitary rooster, with one specimen collected in the dense foliage of a bush Nowak (1999). REPRODUCTION AND ONTOGENY: The female gives birth to a single young (Hayssen et al., 1993). DIET: Gembu Tungaluna (2012: 91 - 93) followed the consumption of fruits by this species year-round in the DRC and reported the following: First heavy rain season (September - November): Uapaca guineensis Müll.Arg., 1864 (Sugar plum, red cedar, false mahogany, rikio) (31%), Ficus leprieuri Miq., 1867 (19%), Ficus wildemanniana Warb. (14%), Pseudospondia microcarpa Engl. (10%), Musanga cecropioides R. Br. & Tedlie (African corkwood tree, umbrella tree) (8%), Myrianthus arboreus P. Beauvois, 1805 (giant yellow mulberry, bush pineapple, corkwoord) (8%), Spondias cytherea Sonnerat, 1782 (Ambarella, otaheite apple, great hog plum) (2%), Dacryoides edulis H.J. Lam (safou, African pear, bush pear, Nsafu, bush butter tree, butterfruit) (2%), Carica papaya L. (papaya) (2%). First mild rain season (December - February): Ficus leprieuri (42%), Musanga cecropioides (27%), Myrianthus arboreus (11%), Spondias cytherea (5%), Uapaca guineensis (5%), Ficus vallis-choudae Delile (5%), Ficus wildemanniana (5%). Second heavy rain season (March - May): Ficus wildemanniana (62%), Ficus leprieuri (25%), Pseudospondias microcarpa (A. Rich.) Engl. (13%). "Dry" season (June - August): Ficus leprieuri (100%). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, three individuals for antibody to SARS-CoV in sera from Bandundu Province, DRC, none were found to be positive (0/3). Trend:- 2008: Unknown (Mickleburgh et al., 2008c; IUCN, 2009). 2004: Unknown (Mickleburgh et al., 2004k; IUCN, 2004). Filoviridae - Filo viruses Marburgvirus - Towner et al. (2007) tested 2 individuals from Gabon using real-time RT-PCR and conventional and nested RT-PCR; neither were found to be positive for Marburg virus specific RNA. Drexler et al. (2012a: Suppl. Table S1) indicated that none of the 21 specimens they examined from Gabon tested positive for Respirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Central African Republic, Congo (Democratic Republic of the). PREDATORS: Blanding's Tree Snake (Toxicodryas blandingii) was found in a mist net eating a traped C. argynnis in the Democratic Republic of the Congo (Nagy et al., 2011). POPULATION: Structure and Density:- It is locally not uncommon, with many specimens (~20) caught in mist nets from a locality in the eastern Democratic Republic Figure 43. Distribution of Casinycteris argynnis Casinycteris campomaanensis Hassanin, 2014 *2014. Casinycteris campomaanensis Hassanin, C. R. Biologies, 337 (2): 134, 137, figs 1B, 3, 4. Publication date: 5 February 2014. Type locality: Cameroon: South Region: CampoMa’an area: Nkoélon-Mvini [02 23.831 N 10 02.691 E, 117 m] [Goto Description]. 188 ISSN 1990-6471 Holotype: MNHN ZM-2011-637: ad ♀, skull and alcoholic. Collected by: Alexandre Hassanin; collection date: 21 November 2007; original number: C07-41. Presented/Donated by: ?: Collector Unknown. - Etymology: The specific epithet refers to the Campo-Ma’an area, where the holotype was collected (see Hassanin, 2014: 137). (Current Combination) COMMON NAMES: English: Campo-Ma'an fruit bat. Casinyctère de Campo-Ma'an. Campomaan-Flughund. French: German: CONSERVATION STATUS: Global Justification This species is listed as Data Deficient (DD ver 3.1 (2001)) because it is only known from three specimens and no other informations are available regarding its demographic trend, threats and ecology (Hassanin, 2017). GENERAL DISTRIBUTION: Native: Cameroon, Nigeria (Tanshi et al., 2019: 14788). POPULATION: Structure and Density:- Nothing is known about population trend and size for the species (Hassanin, 2017). Trend:- 2016: Unknown (Hassanin, 2017). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon. Assessment History Global 2016: DD ver 3.1 (2001) (Hassanin, 2017). Reginal None known. MAJOR THREATS: Threats to this species are currently unknown (Hassanin, 2017). CONSERVATION ACTIONS: Hassanin (2017) report that this species is undoubtedly present in the Campo Ma'an National Park, a protected area located near the type locality. Further studies are however needed to better understand the distribution, natural history and threats of this species. Figure 44. Distribution of Casinycteris campomaanensis Casinycteris ophiodon (Pohle, 1943) *1943. Scotonycteris ophiodon Pohle, Sber. Ges. naturf. Freunde Berlin, 78, figs 1 -2 (for 1942). Type locality: Cameroon: Kribi, Bez: Bipindi [03 06 N 10 30 E] [Goto Description]. Holotype: ZMB 50001: sad ♀, skull only. Collected by: Georg August Zenker; collection date: May 1899. See Turni and Kock (2008: 21). Pohle (1943: 78) mentioned that the skull was removed from the alcoholic specimen. - Etymology: From "ophiodon" meaning snake-toothed. 1946. Scotonycteris ophiodon cansdalei Hayman, Ann. Mag. nat. Hist., ser. 11, 12 (95): 766 (for 1945). Publication date: 19 September 1946. Type locality: Ghana: Central province: Oda district: Oda [05 55 N 00 56 W] [Goto Description]. Holotype: BMNH 1946.229: ad ♀, skin and skull. Collected by: George Soper Cansdale; collection date: 24 December 1945; original number: 420. - Etymology: In honour of Mr. George Soper Cansdale, the collector of the type specimen (see Hayman, 1946: 770). 2014. Casinycteris ophiodon: Hassanin, C. R. Biologies, 337 (2): 141. Publication date: 5 February 2014. (Current Combination) African Chiroptera Report 2020 TAXONOMY: See Simmons (2005: 349). COMMON NAMES: Czech: kaloň hadozubý. English: Snake-toothed Harlequin Fruit-bat, Pohle's Harlequin Fruit-bat, Snake-toothed Tear-drop Fruit-bat, Pohle's Teardrop Fruit-bat, Pohle's Fruit Bat. French: Scotonyctère à dents de serpent, Roussette de Pohle. German: Schlangenzähniger HarlekinFlughund, Pohles Harlekin-Flughund, Poehle's Traenenflughund. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU A2c ver 3.1 (2001)) because it is suspected that a population decline, estimated to be more than 30% would be met over the next 10 years or three generations, inferred from the susceptibility of the specialist species to habitat degradation (Mickleburgh et al., 2008db; IUCN, 2009). Assessment History Global 2008: VU A2c ver 3.1 (2001) (Mickleburgh et al., 2008db; IUCN, 2009). 2004: EN A4c ver 3.1 (2001) (Mickleburgh et al., 2004co; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. 189 GENERAL DISTRIBUTION: Scotonycteris ophiodon is known only from a relatively few scattered records in West Africa and Central Africa. It has been recorded from Liberia, Côte d'Ivoire, Ghana, Cameroon and Congo. It seems to be known from 16 localities only, and has not been recorded from a number of areas where intensive netting has taken place. The species has been recorded at elevations from 120 m (Oda, Ghana) to 1,200 m asl (Mount Nimba). Native: Cameroon; Congo; Côte d'Ivoire (Henry et al., 2004: 24); Ghana; Liberia. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Haiduk et al., 1980; 1981: 226) reported 2n = 34, FN = 62, BA = 32, a subtelocentric X chromosome and an acrocentric Y chromosome for specimens from Cameroon. Protein / allozyme - Unknown. POPULATION: Structure and Density:- This appears to be a naturally rare species (Mickleburgh et al., 2008db; IUCN, 2009). Trend:- Decreasing (Mickleburgh et al., 2008db; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo, Côte d'Ivoire, Ghana, Liberia. MAJOR THREATS: This species is highly impacted by deforestation, and through forest fragmentation and degradation. Threats presumably include logging and mining activities, and the conversion of land to agricultural use (Mickleburgh et al., 2008db; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008db) [in IUCN (2009)] report that this species has been recorded from the Mount Nimba World Heritage Site (Guinea, Liberia and Côte d'Ivoire) and from Tai National Park (Côte d'Ivoire). There is a need to maintain areas of suitable primary forest throughout much of West Africa, especially in areas where this seemingly rare bat has been recorded. Further field surveys are needed to locate any additional populations of this species. Figure 45. Distribution of Casinycteris ophiodon Genus Scotonycteris Matschie, 1894 *1894. Scotonycteris Matschie, Sber. Ges. naturf. Freunde Berlin, 200. - Comments: Type species: Scotonycteris zenkeri Matschie, 1894. - Etymology: From the Greek "σκόοτος", meaning darkness and "νυκτερίς", meaning bat (see Palmer, 1904: 627). (Current Combination) 190 ISSN 1990-6471 TAXONOMY: Reviewed by Bergmans (1991). See also Simmons (2005: 349), Hassanin et al. (2015). Based on cyt b analyses, Hassanin (2014: 141) transferred ophiodon Pohle, 1943 from Scotonycteris to Casinycteris. Further analyses on cyt b and nuclear DNA data led Hassanin et al. (2015: 206) to split up S. zenkeri in four clades of which they recognized three as full species. Included in the Scotonycterini tribe by Bergmans (1997: 69), Almeida et al. (2016: 83) and Hassanin et al. (2020: 5), which was considered part of the Epomophorinae by the former and of the Rousettinae by the latter two. Currently (Simmons and Cirranello, 2020) recognized species of Scotonycteris: bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné, and Bonillo, 2015, occidentalis Hayman, 1947, zenkeri Matschie, 1894. COMMON NAMES: Czech: hadozubí kaloni. English: Harlequin Fruitbats, Tear-drop Fruit-bats, West African fruit bats. French: Scotonyctères. Flughunde. German: Harlekin- MOLECULAR BIOLOGY: Based on analyses of mitochondrial cyt b and nuclear DNA sequences, Hassanin et al. (2015: 205) found support for four clades: Upper Guinea (Liberia and Côte d'Ivoire), Cameroon, western Equatorial Africa (EG, Gabon, RC, and southwestern CAR), and eastern DRC. The mtDNA distances calculated between these four geographic clades are slightly higher than those found between the three species of the genus Casinycteris, which suggest that S. zenkeri might contain four cryptic species. These results are confirmed nuclear DNA data, except for the eastern DRC clade. Hassanin et al. (2015) suggest that females from eastern DRC and western Equatorial Africa have remained isolated since the end of the Pliocene (2.7 Mya), whereas males were able to disperse, at least occasionally during interglacial periods, resulting in malemediated nuclear gene flow between these distant populations, apparently from western Equatorial Africa to eastern DRC. Scotonycteris bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, 2015 *2015. Scotonycteris bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, C. R. Biologies, 338 (3): 206. Publication date: 27 January 2015. Type locality: Central African Republic: Mabaéré-Bodingué National Park: Case of Kpoka [3.90 N 17.16 E, 430 m] [Goto Description]. Holotype: MNHN ZM-2011-713: ad ♀, alcoholic (skull not removed). Collected by: Alexandre Hassanin, Emmanuel Nakouné, Nicolas Nesi and Carine Ngoagouni; collection date: 16 November 2008; original number: R08-119. Presented/Donated by: ?: Collector Unknown. - Etymology: In honour of Dr. Wim Bergmans, a Dutch zoologist, for his outstanding contributions in the fields of taxonomy and biogeography of African fruit bats (see Hassanin et al. (2015: 206). COMMON NAMES: English: Bergmans's fruit bat. French: Scotonyctère de Bergmans. German: Bergmans Harlekin-Flughund. GENERAL DISTRIBUTION: Scotonycteris bergmansi occurs in Equatorial Africa, where it is known from the rainforests of Gabon, Equatorial Guinea, Republic of Congo, southern Central African Republic, and eastern Democratic Republic of the Congo Hassanin et al. (2015: 207). Native: Gabon, Equatorial Guinea, Congo, Central African Republic, Democratic Republic of the Congo. Figure 46. Distribution of Scotonycteris bergmansi African Chiroptera Report 2020 191 Scotonycteris bergmansi bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, 2015 *2015. Scotonycteris bergmansi bergmansi Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, C. R. Biologies, 338 (3): 206. Publication date: 27 January 2015. Type locality: Central African Republic: Mabaéré-Bodingué National Park: Case of Kpoka [3.90 N 17.16 E, 430 m] [Goto Description]. - Etymology: In honour of Dr. Wim Bergmans, a Dutch zoologist, for his outstanding contributions in the fields of taxonomy and biogeography of African fruit bats (see Hassanin et al. (2015: 206). GENERAL DISTRIBUTION: Scotonycteris bergmansi bergmansi is found in the rainforests of western Equatorial Africa in the following countries: Central African Republic (South), Republic of Congo, Gabon, and Equatorial Guinea. Karyotype - Primus et al. (2006) reported a male "S. zenkeri" from Gabon with a 2n = 32. Protein / allozyme - Unknown. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Central African Republic, Equatorial Guinea, Gabon. MOLECULAR BIOLOGY: DNA - Unknown. Scotonycteris bergmansi congoensis Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, 2015 *2015. Scotonycteris bergmansi congoensis Hassanin, Khouider, Gembu, Goodman, Kadjo, Nesi, Pourrut, Nakouné and Bonillo, C. R. Biologies, 338 (3): 207. Publication date: 27 January 2015. Type locality: Congo (Democratic Republic of the): Sukisa [2.31 N 24.98 E, 470 m] [Goto Description]. Holotype: MNHN ZM-2014-564: ad ♀, alcoholic (skull not removed). Collected by: ?: Collector Unknown; original number: K13-48. Presented/Donated by: ?: Collector Unknown. GENERAL DISTRIBUTION: Scotonycteris bergmansi congoensis is found in the rainforests of eastern DRC. Native: Congo (Democratic Republic of the). DIET: Gembu Tungaluna (2012: 110 - 112) reported the following fruits being eaten by "Scotonycteris zenkeri" in the Kisangani area (DRC) during the various seasons: First heavy rain season (September - November): Ficus leprieuri Miq., 1867 (29%), Ficus wildemanniana Warb. (22%), Uapaca guineensis Müll.Arg., 1864 (Sugar plum, red cedar, false mahogany, rikio) (14%), Musanga cecropioides R.Br. & Tedlie (African corkwood tree, umbrella tree) (14%), Spondias cytherea Sonnerat, 1782 (Ambarella, otaheite apple, great hog plum) (7%), Annonidium mannii (Oliv.) Engl. & Diels (Junglesop) (7%), Pseudospondias microcarpa (A. Rich.) Engl. (7%). First mild rain season (December - February): Ficus wildemanniana (33%), Ficus leprieuri (27%), Myrianthus arboreus P. Beauvois, 1805 (giant yellow mulberry, bush pineapple, corkwoord) (20%), Musanga cecropioides (20%). Second heavy rain season (March - May): Annonidium mannii (17%), Musanga cecropioides (17%), Oncoba welwitschii Oliv. (11%), Ficus leprieuri (11%), Ficus vallis-choudae Delile (11%), Ficus sp1 (11%), Parinari excelsa Sabine (Guinea plum) (6%), Carica papaya L. (papaya) (6%), Uapaca guineensis (5%), Spondias cytherea (5%). "Dry" season (June - August): Parinari excelsa (50%), Ficus leprieuri (25%), Musanga cecropioides (25%). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the). Scotonycteris occidentalis Hayman, 1947 1947. Scotonycteris zenkeri occidentalis Hayman, Ann. Mag. nat. Hist., ser. 11, 13 (104): 547 (for 1946). Publication date: 25 June 1947. Type locality: Ghana: Central province: Oda [05 55 N 00 56 W] [Goto Description]. Holotype: BMNH 1946.898: ad ♀, skin and skull. 192 ISSN 1990-6471 ? Collected by: George Soper Cansdale; collection date: 1946. - Etymology: occidentalis refers to the western Africa. Scotonycteris occidentalis: (Current Combination) COMMON NAMES: English: Hayman’s tear-drop fruit bat. French: Scotonyctère de Hayman. German: Haymans Harlekin-Flughund. CONSERVATION ACTIONS: Mickleburgh et al. (2008bm) [in IUCN (2009)] report that although there appear to be no direct conservation measures in place [for "S. zenkeri"], it has been recorded from a number of protected areas (e.g. Tai National Park, Côte d'Ivoire). There is a need to investigate whether this species can adapt to secondary forest habitats. Further studies are needed to fully determine if the species is present in forested areas between known localities. Nigerian populations might belong to S. zenkeri, leaving the possibility that some others might belong to S. occidentalis]. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Côte d'Ivoire, Ghana, Guinea, Liberia. GENERAL DISTRIBUTION: Scotonycteris occidentalis is distributed in West Africa, where it ranges from Liberia and Guinea, into Côte d'Ivoire, Ghana and possibly Nigeria. Native: Côte d'Ivoire (Heim de Balsac, 1934b: 24 as "S. zenkeri); Ghana; Guinea; Liberia; Nigeria [Hassanin et al. (2015: 206) indicate that some Figure 47. Distribution of Scotonycteris occidentalis Scotonycteris zenkeri Matschie, 1894 *1894. Scotonycteris zenkeri Matschie, Sber. Ges. naturf. Freunde Berlin, 202. Type locality: Cameroon: Yaunde [03 52 N 11 31 E]. Holotype: ZMB 66533: ad ♀, skull and alcoholic. Collected by: Georg August Zenker. See Turni and Kock (2008: 21). (Current Combination) 1904. Scotonycteris bedfordi Thomas, Abstr. Proc. Zool. Soc. Lond., 4: 14. Publication date: 8 March 1904. Type locality: Equatorial Guinea: Bioko: Fish Town [10 m] [Goto Description]. Holotype: BMNH 1904.7.1.28: ad ♀, skin and skull. Collected by: E. Seimund; collection date: 2 January 1904; original number: 31. Presented by Fernando Poo Committee: see Andersen (1912b: 568). ? Scotonycteris zenkeri zenkeri: (Name Combination) TAXONOMY: See Simmons (2005: 349). See also Hassanin et al. (2015: 206), who split-off occidentalis and bergmansi. COMMON NAMES: Castilian (Spain): Zorra Voladora de Zenker. Czech: kaloň Zenkerův. English: Zenker's Harlequin Fruit-bat, Zenker's Tear-drop Fruit-bat, Zenker's Fruit Bat. French: Scontonyctère de Zenker, Roussette de Zenker. German: Zenkers Harlekin-Flughund, Zenker's Traenenflughund. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in some protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bm; IUCN, 2009; Bergmans et al., 2017a). Assessment History Global 2016: LC ver 3.1 (2001) (Bergmans et al., 2017a). 2008: LC ver 3.1 (2001) (Mickleburgh et al., African Chiroptera Report 2020 2008bm; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bk; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). This status is for S. zenkeri as perceived prior to the publication of Hassanin et al. (2015) and will need to be re-evaluated in view of the split-up of this species. Regional None known. MAJOR THREATS: The species is threatened by habitat loss, presumably largely through logging and mining operations and the conversion of land to agricultural use (Mickleburgh et al., 2008bm; IUCN, 2009; Bergmans et al., 2017a). CONSERVATION ACTIONS: Bergmans et al. (2017a) report that although there appear to be no direct conservation measures in place, it has been recorded from a number of protected areas (e.g., Tai National Park, Côte d'Ivoire). There is a need to investigate whether this species can adapt to secondary forest habitats. Further studies are needed to fully determine if the species is present in forested areas between known localities. GENERAL DISTRIBUTION: Hassanin et al. (2015: 206) restricted the distribution of S. zenkeri to Cameroon, with the remark that additional data are needed to assess the taxonomic status of some populations from Nigeria. Tentatively, we assigned the Nigerian specimens to this species. 193 MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Primus et al. (2006) reported a male from Gabon with a 2n = 32, but the specimen on which this is based, would currently be assigned to S. bergmansi, so the karyotype for S. zenkeri is still to be confirmed. Protein / allozyme - Unknown. HABITAT: Juste B. and Perez del Val (1995: 144) reported that S. zenkeri was only netted below 400 m, and was found more frequently in cultivated clearings than in rainforests. POPULATION: Structure and Density:- It appears to be a rare, or at least rarely recorded, species (Mickleburgh et al., 2008bm; IUCN, 2009; Bergmans et al., 2017a). Trend:- 2016: Decreasing (Bergmans et al., 2017a). 2008: Decreasing (Mickleburgh et al., 2008bm; IUCN, 2009). The above statements pertain to S. zenkeri s.l. and need to be re-evaluated in view of the split-up proposed by Hassanin et al. (2015: 206). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo, Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Nigeria. Native: Cameroon; Nigeria(?). DETAILED MORPHOLOGY: Chawana et al. (2013: 160) mention that the average brain mass for 2 specimens was 0.64 g. FUNCTIONAL MORPHOLOGY: Sagot and Chaverri (2015: 191) indicated that the retinal area in the eye of S. zenkeri covers about 40 - 50 mm2 and contains about 90,000 retinal ganglion cells. The minimum angle of resolution was about 0.227° (ca. 8 mm at 1 m distance). Figure 48. Distribution of Scotonycteris zenkeri TRIBE Stenonycterini Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, 2012 *2012. Stenonycterini Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, Mol. Phylog. Evol., 66 (1):. Publication date: 10 October 2012. 194 ISSN 1990-6471 TAXONOMY: Includes only the genus Stenonycteris K. Andersen, 1912 (Almeida et al., 2020: 11). Genus Stenonycteris Gray, 1870 1870. Stenonycteris Gray, Catalogue of Monkeys, Lemurs and Fruit-eating bats in the collection of the British Museum London, 127. - Comments: Considered a valid genus by Giannini and Simmons (2003: 504), Lavrenchenko et al. (2004b: 130), but only a synonym according to Bergmans (1994: 79). Sometimes considered a valid subgenus containing lanosus. Allen (1939a: 62), Meester et al. (1986: 28), Bergmans (1994: 80), and Kwiecinski and Griffiths (1999: 1) assigned this name to K. Andersen, Cat. Chiroptera Brit. Mus., 1: 23, 1912. - Etymology: From the Greek stenos meaning narrow and nycteris meaning bat (see Andersen, 1912b: 23). TAXONOMY: Included in the Stenonycterini tribe (Almeida et al., 2016: 83), and in the Epomophorini tribe and Stenonycterina subtribe by Hassanin et al. (2020: 5). Currently (Simmons and Cirranello, 2020) recognized species of the genus Stenonycteris: lanosus (Thomas, 1906). Stenonycteris lanosus (Thomas, 1906) *1906. Rousettus lanosus Thomas, Ann. Mag. nat. Hist., ser. 7, 18 (104): 137. Publication date: 1 August 1906. Type locality: Uganda: Ruwenzori East: Mubuku Valley [00 15 N 30 10 E, 13 000 ft] [Goto Description]. Holotype: BMNH 1906.7.1.2: ad ♂, skull and alcoholic. Collected by: Mr. R.B. Woosnam. Presented by Ruwenzori Exploration Committee: see Andersen (1912b). - Comments: Type locality originally "Ruwenzori East at 13,000 ft", but restricted to Mubuku Valley by Hayman et al. (1966: 30) (see Bergmans, 1994: 109). Etymology: From the Greek/Latin? "lanosus" for woolly. (Current Combination) 1909. Rousettus kempi Thomas, Ann. Mag. nat. Hist., ser. 8, 4 (24): 543. Publication date: 1 December 1909. Type locality: Kenya: Southern slopes Mount Elgon, Kirui's: Twere [ca. 00 46 N 34 37 E, 6 000 ft] [Goto Description]. Holotype: BMNH 1910.4.1.6: sad ♂, skin and skull. Collected by: Robin Kemp; collection date: 16 September 1909; original number: 269. Presented/Donated by: C.D. Rudd Esq. Formerly in Rudd collection. 2012. Stenonycteris lanosus: Nesi, Kadjo, Pourrut, Leroy, Pongombo Shongo, Cruaud and Hassanin, Mol. Phylog. Evol., 66 (1): 135 (for 2013). Publication date: 10 October 2012. (Name Combination, Current Combination) ? Rousettus (Stenonycteris) lanosus: (Name Combination) ? Rousettus lanosus kempi: (Name Combination) ? Rousettus lanosus lanosus: (Name Combination) TAXONOMY: Revised by Bergmans (1994), but see Simmons (2005: 348). Included in the genus Stenonycteris by Almeida et al. (2011a: 5, 7, 9, 10) and Patterson and Webala (2012: 10) since it consistently recovered in another clade allied to Myonycteris, Lissonycteris, and Megaloglossus. Nesi et al. (2012: 126) confirm that Stenonycteris should be considered a separate genus and should even be placed in its own tribe (Stenonycterini), within the subfamily Epomophorinae. COMMON NAMES: Chinese: 狭 齿 果 蝠 . Czech: kaloň vlnatý. English: Mountain Rousette, Long-haired Rousette, Woolly Rousette, Ruwenzori Longhaired Rousette. French: Roussette du Ruwenzori, Roussette à longs poils. German: Gebirgs-Höhlenflughund. Kinande (DRC): Karakonube CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) [as Rousettus lanosus] in view of its wide distribution, presumed large population, it occurs in a number of protected areas, and because the global population is unlikely to be declining fast enough African Chiroptera Report 2020 to qualify for listing in a more threatened category (Mickleburgh et al., 2008dg; IUCN, 2009). Assessment History Global 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008dg; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cx; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The major threats to this species include ongoing deforestation in montane regions, likely disturbance of cave roosting sites (possibly from tourists in parts of its range), and possible overhunting of bats at roost sites for subsistence food by local people (Mickleburgh et al., 2008dg; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008dg) [in IUCN (2009)] report that although there appear to be no direct conservation measures in place, it has been recorded from a number of protected areas (e.g. Mount Elgon National Park, Kenya). There is a need to maintain remaining areas of suitable montane forest and to protect major roosting colonies from disturbance. In areas where this species is identified as being overhunted, there is a need to encourage the sustainable use of susceptible populations. 195 POPULATION: Structure and Density:- This appears to be a little recorded, but locally abundant species (Mickleburgh et al., 1992). It forms colonies of up to a few hundred bats (Mickleburgh et al., 2008dg; IUCN, 2009). Trend:- 2008: Decreasing (Mickleburgh et al., 2008dg; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) report a maximum longevity of 18.7 years. REPRODUCTION AND ONTOGENY: Stanley and Goodman (2011: 40) reported on five pregnant females collected between 19 and 24 July 1993 in the South Pare Mountains (Tanzania). Kingdon (1974) [in Krutzsch (2000: 111)] showed that S. lanosus is one of the species in which pregnant or lactating females are present in the population in all months of the year, and where most adult males have enlarged spermatogenic testes and hypertrophied, secretion-filled accessory glands. This would indicate that while males are fecund population-wide, they enter periods of inactivity individually. PARASITES: HAEMATOSPORIDA The two South Sudanese "Rousettus lanosus" specimens tested by Schaer et al. (2017: 2) were infected by Hepatocystis parasites. GENERAL DISTRIBUTION: Stenonycteris lanosus is distributed in East Africa and marginally in Central Africa. It ranges from Ethiopia and southern Sudan in the north of its range, into Kenya, Uganda, eastern Democratic Republic of the Congo and Rwanda, to Tanzania and northern Malawi. It is largely, but not strictly, a montane species being found between 500 and 4,000 m asl (with most records above 1,000 m, e.g. at 13,000 ft in the Mubuku Valley, E. Ruwenzori [Thomas, 1910b: 486]). DIPTERA Nycteribiidae: Dipseliopoda setosa Theodor, 1955 from Kenya and Tanzania (Haeselbarth et al., 1966: 116). The fly Stylidia scissa Speiser, 1901 was collected on a bat from South Sudan (the same bat where also T. dina was collected from, see below), although Bergmans (1982: 161) indicates that this fly is generally found on Microchiropterans and its find might "be due to straggling between spirit specimens". Native: Congo (The Democratic Republic of the); Ethiopia (Bergmans, 1994: 79); Kenya; Malawi (Bergmans, 1994; Bergmans, 1994: 79; Monadjem et al., 2010d: 555); Rwanda (Bergmans, 1994: 79); Sudan; Tanzania (Stanley and Goodman, 2011: 41); Uganda (Kityo and Kerbis, 1996: 60). SIPHONAPTERA Ischnopsyllidae: Thaumapsylla breviceps Rothschild 1907, where the nominate subspecies T. b. breviceps Hopkins and Rothschild, 1956 is occurring in Kenya and the DRC (Haeselbarth et al., 1966: 186). Thaumapsylla dina Jordan, 1937 in the mountains of the Ruwenzori, Congo and Kenya (Haeselbarth et al., 1966: 186). (Bergmans (1982: 161) reported that this flea was also collected from a "Rousettus lanosus" from South Sudan. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data: Fa: 94.5 mm, mass: 121 g, wing loading: 21.8 N/m 2, aspect ratio: 5.6. 196 ISSN 1990-6471 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burundi, Congo (Democratic Republic of the), Egypt, Ethiopia, Kenya, Malawi, Rwanda, South Sudan, Tanzania, Uganda. Figure 49. Distribution of Stenonycteris lanosus †Genus Turkanycteris Gunnell and Manthi, 2018 *2018. Turkanycteris Gunnell and Manthi, J. Hum. Evol., p. "2", "4". Publication date: 6 April 2018 [Goto Description]. - Etymology: Referring to the Turkana Basin, where Kanapoi (the type locality of the species) is located (Gunnell and Manthi, 2018: "4"). †Turkanycteris harrisi Gunnell and Manthi, 2018 *2018. Turkanycteris harrisi Gunnell and Manthi, J. Hum. Evol., p. "2", "4", fig. 1A. Publication date: 6 April 2018. Type locality: Kenya: Nzube's Mandible Site [Goto Description]. Holotype: NMK KP-68723: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. The holotype is a M1. - Etymology: In honour of John M. Harris for his many contributions to the understanding of African Neogene vertebrates (Gunnell and Manthi, 2018: "4"). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Pliocene (Gunnell and Manthi, 2018: "4".). INFRAORDER RHINOLOPHIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 1866. Histiorrhina Haeckel, Generelle Morphologie der Organismen., 2: clx. - Comments: Proposed as family, including the genera Rhinolophus Lacépède, 1799; Megaderma É. Geoffroy, 1810; and Phyllostoma G. Cuvier, 1800 [= Phyllostomus Lacépède, 1799] (see Jackson and Groves, 2015: 229). 1998. Megachiropteramorpha Simmons and Geisler, Bull. Am. Mus. Nat. Hist., 235: 135, 136. Comments: Originally included the fossil Archaeopteropus Meschinelli, 1903 and the suborder Megachiroptera (Dobson, 1875) (see Jackson and Groves (2015: 229). *2007. RHINOLOPHIFORMACEI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the Infraorder is based on Rhinolophus Lacépède, 1799 (type of RHINOLOPHIDAE Gray, 1825). (Current Combination) African Chiroptera Report 2020 197 Superfamily RHINOLOPHOIDEA J. E. Gray, 1825 1825. Rhinolophina Gray, Ann. Philos., 10 (5): 338. - Comments: Proposed as tribe and originally containing the genera Megaderma É. Geoffroy, 1810; Rhinolophus Lacépède, 1799; Nycteris É. Geoffroy & G. Cuvier, 1795; Mormoops Leach, 1821 and Nyctophilus Leach, 1821. Jackson and Groves (2015: 241, 242) placed this name in the synonymy of both Rhinolophoidea and Rhinolophidae. *1825. Rhinolophoidea Gray. - Comments: The name of the Superfamily is based on Rhinolophus Lacépède, 1799 (type of RHINOLOPHIDAE Gray, 1825). Some publications assigned the name to Bell (1836). (Current Combination) COMMON NAMES: Czech: vrápencovci, vrápenci. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Shi and Rabosky (2015: 1532) [based on Teeling et al. (2005)] mention 55 milion years ago as approximate fossil date for the Rhinolophoidea, and Patterson and Webala (2012: 12) indicate that the separation between the Rhinolophidae and the Hipposideridae took place some 28.7 mya, although Shi and Rabosky (2015: 1532) set this about 8 million years earlier (at 37 mya). Foley et al. (2014: 320) provide another estimate for the African divergence between the Rhinolophidae and the Hipposideridae from their common ancestor: 42 mya. Based on transcriptome RNAseq data, Lei and Dong (2016: 2) placed the split at about 40 mya. Ravel et al. (2016: 358) indicate that the oldest occurrence of the Hipposideridae in northern Africa and the oldest occurrence of the Rhinolophidae in eastern Asia suggest that a common ancestor for these two families might have occurred in Central Asia. Unfortunately, there are no fossil records to substantiate this. BIOGEOGRAPHY: Foley et al. (2014: 320) indicate that the Rhinolophoidea and Pteropodidae diverged approximately 59 Mya. While within the Rhinolophoidae, the families Rhinolophidae and Hipposideridae diverged in Africa from their common ancestor approximately 42 May during the Eocene. The family Rhinonycteridae separated from the other Hipposideridae approximately 39 Mya, also in Africa (Foley et al., 2014: 320). Family HIPPOSIDERIDAE Lydekker, 1891 1860. Phyllorhina Koch, Bericht der Oberhessischen Gesellschaft für Natur- und Heilkunde, 8 (4): 26. - Comments: Type genus: Phyllorrhina Bonaparte, 1837 [1832–1841] [= Hipposideros J. Gray, 1831]. Jackson and Groves (2015: 245) indicate the name is unavailable. 1860. Phyllorrhina Koch. - Comments: Pavlinov et al. (1995: 75) mention Phyllorrhina with a question mark as synonym of Hipposideridae. Lanza et al. (2015: 121) mention "Phyllorhina" Koch as a synonym of the genus Asellia. *1891. Hipposideridae Lydekker, in: Flower and Lydekker, Mammals Living and Extinct, 657. (Current Combination) *1891. Hipposiderinae Lydekker, in: Flower and Lydekker, Mammals Living and Extinct, 657. Comments: Type genus: Hipposideros Gray, 1831. Originally included the genera Hipposiderus [sic] J. Gray, 1834 [= Hipposideros J. Gray, 1831]; Anthops Thomas, 1888; Rhinonicteris J. Gray, 1847; Triaenops Dobson, 1871; and Coelops Blyth, 1848 (see Jackson and Groves, 2015: 244). (Current Combination) 1907. Hipposideridae Miller, Bull. U.S. natl. Mus., 57: viii, 109. Publication date: 29 June 1907. - Comments: Type genus: Hipposideros J. Gray, 1831. Originally included the genera Hipposideros J. Gray, 1831; Asellia J. Gray, 1838; Anthops Thomas, 1888; Coelops Blyth, 1848; Cloeotis Thomas, 1901; Rhinonicteris J. Gray, 1847; and Triaenops Dobson, 1871 (see Jackson and Groves, 2015: 245). 1941. Coelopinae Tate, Am. Mus. Novit., 1140: 11. Publication date: 20 August 1941. Comments: Type genus: Coelops Blyth, 1848. 1970. Hipposiderini Koopman and J.K. Jones Jr., in: Slaughter, Classification of bats, 25. Comments: Type genus: Hipposideros J. Gray, 1831. Proposed as tribe and originally 198 ISSN 1990-6471 1994. 1995. 2014. 2014. 2015. 2018. included the genera Hipposideros J. Gray, 1831; Aselliscus Tate, 1941; Cloeotis Thomas, 1901; Rhinonicteris J. Gray, 1847 and Triaenops Dobson, 1871 (see Jackson and Groves, 2015: 245). Hipposiderina Koopman, in: Niethammer et al., Chiroptera: Systematics, 60. - Comments: Type genus: Hipposideros J. Gray, 1831. Proposed as subtribe and originally included the genera Hipposideros J. Gray, 1831; Anthops Thomas, 1888 and Asellia J. Gray, 1838 (see Jackson and Groves, 2015: 245). Coelopsinae: Pavlinov, Borissenko, Kruskop and Jahonton, Archives of the Zoological Museum, Moscow State University,, 133: 84. - Comments: Mentioned as a synonym of the Hipposideridae by Pavlinov et al. (1995: 84). (Lapsus) Hippossideridae: Ndara R., IJISR, 12 (1): 253. Publication date: November 2014. (Lapsus) Hippossyderidae: Ndara R., IJISR, 12 (1): 252. Publication date: November 2014. (Lapsus) Hypposideridae: Lanza, Funaioli and Riccucci, The bats of Somalia and neighbouring areas, 118. (Lapsus) Hipossidaridae: Malekani, Musaba, Gembu, Bugentho, Toengaho, Badjedjea, Ngabu, Mutombo, Laudisoit, Ewango, Van Cakenberghe, Verheyen, Asimonyo, Masudi, Bongo and Ngbolua, Nat. Conserv. Res., 3 (1): 68. Publication date: January 2018. (Lapsus) TAXONOMY: Considered a subfamily of the Rhinolophidae by Koopman (1993a: 169), Schutt and Simmons (1998: 28), but see Harrison and Bates (1991: 51), Corbet and Hill (1992: 104), Peterson et al. (1995: 66), Hutson et al. (2001), Simmons (2005: 365), who consider this to be a valid family. Hoofer and Van Den Bussche (2003: 10 - 12) suggest that the Hipposideridae might not be monophyletic, with Triaenops being different, although they state this might be the result of inadequate sampling. McKenna and Bell (1997) used the name Rhinonycterinae Gray, 1866 for this group, but this has not been accepted by other authors (Simmons, 2005: 365). Although Rhinonycteridae (= Rhinonycterina Gray, 1866) has priority over Hipposideridae as a family-group name, nobody other than Gray (1866b) used the former name until it was resurrected by McKenna and Bell (1997). Miller (1907) used the name Hipposideridae for this group because Hipposideros Gray, 1831 has priority over Rhinonycteris Gray, 1866 (= Rhinonicteris Gray, 1847). Subsequent authors have followed Miller (1907) usage of Hipposideridae/inae, and Simmons (2005: 365) believes that there is little to be gained by replacing it with an unknown name. Simmons (2005: 365) therefore retains Hipposideridae for this group pending action by the International Commission on Zoological Nomenclature. Due to the recent split off of Cloetis, Paratriaenops, Rhinonicteris and Triaenops into a separate family, the above naming issue has resolved itself. McKenna and Bell (1997) proposed a tribal classification for hipposiderids (which they treated as a subfamily), but many of the groups they defined have subsequently been shown to be paraphyletic (Bogdanowicz and Owen, 1998, Hand and Kirsch, 1998); accordingly, Simmons (2005: 365) did not recognize subfamilies or tribes within Hipposideridae. Currently (Simmons and Cirranello, 2020) recognized genera of Hipposideridae: Anthops Thomas, 1888, Asellia Gray, 1838, Aselliscus Tate, 1941, Coelops Blyth, 1848, Doryrhina Peters, 1871, Hipposideros Gray, 1831, Macronycteris Gray, 1866, Additionally, there are also the extinct genera †Miophyllorhina Hand, 1997; †Palaeophyllophora Revilliod, 1917; †Riversleigha Hand, 1998, †Vaylatsia Sigé, 1990. COMMON NAMES: Czech: pavrápencovití, listonosovití. Dutch: Bladneusvleermuizen. English: Old-World leafnosed bat, Trident bats, Leaf-nosed bats. French: Hipposidéridés. German: Rundblattnasen. Italian: Ipposidèridi. Norwegian: Rundbladneser, falske hesteskoneser. Russian: Листоносы, Подковогубы, Ложные подковоносы. Ukrainian: Звичайні листоноси [= Zvychayni lystonosy]. Vietnamese: Họ dơi mũi. SIMILAR SPECIES: Key characters for the genera: Anthops - Head and body is about 50 mm, the tail is very short (less than half as long as the femur), and the forearm length is about 48 - 51 mm. The horseshoe-shape part of the nose leaf has two lateral leaflets, the inner one being quite small and the outer one large. Traces of a small frontal sac are present (Nowak, 1994: 115). African Chiroptera Report 2020 199 Asellia - Head and body length is about 46 - 62 mm, tail length is 18 - 27 mm (tail protruding up to 7 mm beyond interfemoral membrane), and forearm length is 45 - 52 mm. Posterior component of noseleaf with three short, triangular projections. Only one upper premolar on each side, M3 greatly reduced (two ridges) (Nowak, 1994: 114, Happold, 2013al: 357). Aselliscus - Head and body length is about 38 - 45 mm, tail length is 20 - 40 mm (tail extends beyond the membrane, as in Asellia), and forearm length is 35 - 45 mm. The upper margin of the transverse noseleaf is divided into three points, and two lateral leaflets margin the horseshoe (Nowak, 1994: 116). Coelops - Head and body length is 28 - 50 mm, the tail is extremely short or absent, and forearm length is 33 - 47 mm. Lateral leaflets are lacking on the noseleaf, or they are obscured by dense, stiff hairs (Nowak, 1994: 119). Hipposideros - Head and body is about 35 - 100 mm, tail length is 18 - 70 mm, and forearm length is about 33 - 105 mm. Posterior component of noseleaf roughly elliptical in outline (although the outline of upper margin varies from almost flat to low-arched or even subtriangular), without three projections (Nowak, 1994: 111, Happold, 2013al: 357). Paracoelops - Head and body length, 45 mm; tail absent, forearm length is about 42 mm. The nose leaf is horseshoe-shaped and surmounted by a round leaf with radial striations (Nowak, 1994: 119). Manthi (2008: 47) reported on the presence of a large and small species of Hipposideros at Kanapoi (Kenya) between 4.17 and 4.07 mya. They suggested these bones might have been accumulated by either the barn owl (Tyto alba) or the giant eagle owl (Bubo lacteus). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The Hipposideridae have been found from the Miocene of Songhor (Butler, 1969) and from the Mio-Pliocene of Morocco (Lavocat, 1961). The family goes back to the middle Eocene in Europe (Butler, 1978), where the genus Palaeophyllophora is considered to be the oldest representative (Ravel et al., 2016: 412). Early Oligocene to recent in Arabia, early Miocene to recent in Africa, Pleistocene to recent in Asia, late Oligocene to recent in Australia (Hand and Kirsch, 1998). Because they are predominantly cavedwelling bats, their remains are frequently preserved in Cenozoic limestone sediments (Happold, 2013al: 357). The earliest known Hipposideros is from the early Miocene, and the subgenus Syndesmotis (into which H. megalotis is sometimes placed) appears to have separated from Hipposideros in the middle Miocene (Happold, 2013al: 357). The fossils of Hipposideridae from the late Eocene and early Oligocene of France were already distinct from the Rhinolophidae that occurred in the same deposits (Lekagul and McNeely, 1977). DENTAL FORMULA: 1123/ 2123 = 30. Except in Asellia and Hipposideros megalotis, which have a dental formula of 1113/2123 = 28. Based on phylogenetic analyses, Bogdanowicz and Owen (1998) suggest that the Hipposideridae probably originated somewhere in the Oriental region. In contrast, based on evidence from fossils, Hand and Kirsch (1998) suggest that the family originated in Australia. Shi and Rabosky (2015: 1537) calculated the family-level stem and crown ages to be respectively 49.9 and 49.3 Mya. BIOGEOGRAPHY: The Hipposideridae diverged from the Rhinolophidae from a common ancestor in Africa approximately 42 Mya during the Eocene (Foley et al., 2014: 320). While the family Rhinonycteridae separated from the other Hipposideridae approximately 39 Mya, also in Africa (Foley et al., 2014: 320). The monophyletic Hipposideros clade diverged from other Hipposideridae approximately 31 Mya during the Oligocene in Africa (Foley et al., 2014: 320). During the Miocene at about 15 Mya, Hipposideros diversified into distinct clades based on geography: these being the African, India and South East Asia clades (Foley et al., 2014: 320). DETAILED MORPHOLOGY: In their comparative study on bat brains, Bhatnagar et al. (2016) found that Hipposiderids stand apart from all other examined bats due to their relatively large adenohypophysis MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) mention the 2n within the family varies between 24 and 52. HABITS: Hipposiderids are often fly-catchers and gleaners but they also forage by slow-hawking, some to a greater extent than others (Happold, 2013al: 358). Unlike most rhinolophids, many hipposiderids forage in small groups (Happold, 2013al: 358). Hipposiderids are unable to scuttle over the ground, and apparently cannot climb (Happold, 2013al: 358). 200 ISSN 1990-6471 DIET: Hipposiderids are insectivorous: the taxa and size of prey varies according to the size of the bat and the robustness of its teeth (Happold, 2013al: 358). PARASITES: Fain (1994: 1280) reports that the Hipposideridae are hosts for four genera of Myobiidae (Acari): Hipposiderobia with 12 species (parasiting on five genera of bats), Metabinuncus also with 12 species (occurring only on the genus Hipposideros), Binunculoides with one species on Hipposideros, and Triaenomyobia, with one species on Triaenops. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, Namibia. Genus Asellia Gray, 1838 *1838. Asellia Gray, Mag. Zool. Bot., Edinburgh, 2 (12): 493. Publication date: 1 February 1838. - Comments: Type species: Rhinolophus tridens E. Geoffroy Saint-Hilaire, 1813. Etymology: From the Greek "α-" "alpha privative" and the Greek feminine substantive "σέλλα" (sélla), meaning "saddle", referring to the absence of a sella in the noseleaf (see Lanza et al., 2015: 121). Palmer (1904: 124), however, mentions that Asellia is an adjective used as a noun, from the Latin "asellus", a little ass - probably in allusion to the long, pointed ears. (Current Combination) 1957. Aseltia: Harrison, Mammalia, 21 (1): 8. Publication date: March 1957. (Lapsus) 2008. Ascidia: Selim, Nahla and Shelfeh, Tishreen Univ. J. Res. Sci. Stud. - Biol. Sci. Ser., 30 (1): 253. (Lapsus) TAXONOMY: Allen (1939a: 78) mentioned that Asellia was originally described as subgenus of Hipposideros. Currently (Simmons and Cirranello, 2020) recognized species of the genus Asellia: arabica Benda, Vallo and Reiter, 2011; italosomalica de Beaux, 1931; patrizii de Beaux, 1931; tridens (E. Geoffroy St.-Hilaire, 1813). Additionally, there is the extinct †vetus Lavocat, 1961. COMMON NAMES: Czech: trojzubcoví pavrápenci. English: Lesser Trident Bats, Trident Bats, Trident Leaf-nosed Bats. French: Tridents, Asellia, Chauve-souris à 3 endentures. German: Dreizackblattnasen. Italian: Asèllie. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Egypt, Sudan. Asellia italosomalica de Beaux, 1931 *1931. Asellia tridens italo-somalica de Beaux, Ann. Mus. civ. Stor. nat. Genova, 55: 190. Publication date: 6 June 1931. Type locality: Somalia: Oddur. Holotype: MSNG 30942: ♀, skin and skull. Collected by: N. Mosconi Brozzi; collection date: 1929. Presented/Donated by: ?: Collector Unknown. See Benda et al. (2011f: 266). Etymology: From the feminine scientific Latin adjective italosomalica, meaning "inhabiting Italian Somalia, referring to country where the type specimens were collected (see Lanza et al., 2015: 123).. ? Asellia italosomalica: (Current Combination) ? Asellia tridens italosomalica: (Alternate Spelling) TAXONOMY: Based on morphometric analyses and a K2P genetic distance of 12.3 to 13.7 % with other Asellia populations, Benda et al. (2011f: 262) distinguish italosomalica specimens from Socotra and Somalia as a different, well-defined taxon, which is best considered to be a separate species. COMMON NAMES: English: De Beaux's Trident Bat, Somali Trident Leaf-nosed Bat (Armstrong et al., 2016: 116). German: Somalische Dreizackblattnase. Italian: Asèllia italosòmala. CONSERVATION STATUS: Global Justification This is a newly established species without sufficient data on distribution and ecology to be able to assess it beyond Data Deficient at present (Benda, 2017b). Assessment History African Chiroptera Report 2020 Global 2016: DD ver. 3.1 (2001) (Benda, 2017b). Regional None known. MAJOR THREATS: Human disturbance in roost sites and pesticide use against insects are the main threats (Benda, 2017b). CONSERVATION ACTIONS: Benda (2017b) report that no specific measures are known to be in place for this species. 201 GENERAL DISTRIBUTION: Socotra, Somalia POPULATION: Structure and Density:- Benda (2017b) report that no data is available about population size and trend for this species. Trend:- 2016: Unknown (Benda, 2017b). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Somalia. Asellia patrizii de Beaux, 1931 *1931. Asellia patrizii de Beaux, Ann. Mus. civ. Stor. nat. Genova, 55: 186. Publication date: 6 June 1931. Type locality: Eritrea: Dancalia: Gaare [13 13 N 42 07 E]. Holotype: MSNG 13313: ♂, skull and alcoholic. Collected by: Marquis Saverio Patrizi Montoro; collection date: December 1912. Presented/Donated by: ?: Collector Unknown. See Benda et al. (2011f: 267). Paratype: MSNG 13314: ♀, skull and alcoholic. Collected by: Marquis Saverio Patrizi Montoro; collection date: December 1929. Presented/Donated by: ?: Collector Unknown. See Benda et al. (2011f: 267), who mention this specimen along with the holotype, but not explicity as "paratype". They do mention three further paratypes from the series: MSNG 33203, 33204 [S+B], 31315a, 31315b, collected at Attab (Eritrea), in July 1893 by G. Pestalozza and October 1906 by P. Felter. - Etymology: In honour of the Marquis Saverio Patrizi Nero Montoro (1902 - 1978), eminent Italian entomologist, biospeleologist and explorer, collector of zoological material especially in central and eastern Africa, Sardinia and eastern Mediterranean countries (see Lanza et al., 2015: 128). (Current Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: pavrápenec eritrejský. English: Patrizi's Lesser Trident Bat, Patrizi's Trident Leaf-nosed Bat. French: Trident de Patrizi, Asellia de Patrizi. German: Patrizis Dreizackblattnase. Italian: Asèllia di Patrìzi. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008a; IUCN, 2009; Monadjem et al., 2017e). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017e). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008a; IUCN, 2009). 2004: VU D2 ver 3.1 (2001) (Mickleburgh et al., 2004ad; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There is very little information, but roost disturbance is a possible threat. However, the species is associated with habitats that are not under significant threat, and has been found in buildings, and so can adapt to human presence (Mickleburgh et al., 2008a; IUCN, 2009; Monadjem et al., 2017e). CONSERVATION ACTIONS: Mickleburgh et al. (2008a) and Monadjem et al. (2017e) suggest surveys are needed to determine the distribution of this species, both in northeastern Africa and in the Arabian Peninsula. Has been recorded from Awash National Park (Hill and Morris, 1971). GENERAL DISTRIBUTION: This species has a restricted known range in northeastern Africa, from central Ethiopia north to Eritrea (Yalden et al., 1996), and also from islands in the Red Sea including Farasan Island and Segid Island (=As-Saqid) of southwestern Saudi Arabia 202 ISSN 1990-6471 (Moeschler et al., 1990; Harrison and Bates, 1991: 316; Horácek et al., 2000). It probably occurs more widely in this poorly explored region (Mickleburgh et al., 2008a). Native: Eritrea (Yalden et al., 1996); Ethiopia (Yalden et al., 1996); Saudi Arabia (Moeschler et al., 1990; Harrison and Bates, 1991: 316; Horácek et al., 2000). 2009). 2004: Decreasing (Mickleburgh et al., 2004ad; IUCN, 2004). ACTIVITY AND BEHAVIOUR: It flies close to the ground catching insects (Mickleburgh et al., 2008a; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Eritrea, Ethiopia. HABITAT: Dry open terrain such as the semi-desert grassland of the Awash National Park (Hill and Morris, 1971). ROOST: It roosts in caves and buildings and in dry open terrain such as the semi-desert grassland of the Awash National Park (Hill and Morris, 1971). POPULATION: Structure and Density:- Appears to be rare with small roosts. There is a possibility that it will be found in larger roosts, but to date none have been found (Mickleburgh et al., 2008a; IUCN, 2009; Monadjem et al., 2017e). Trend:- 2016: Unknown (Monadjem et al., 2017e). 2008: Unknown (Mickleburgh et al., 2008a; IUCN, Figure 50. Distribution of Asellia patrizii Asellia tridens (E. Geoffroy St.-Hilaire, 1813) *1813. Rhinolophus tridens E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 20: 260, pl. 5. Publication date: 1813. Type locality: Egypt: Qena province: Luxor, near [ca. 25 41 N 32 39 E] [Goto Description]. Syntype: MNHN A.235: ad ♂, alcoholic (skull not removed). Number 187 in Rode (1941: 240), who mentions this specimen as holotype. Furthermore, he remarks that the skull is not removed, so the skull in the atlas, pl. IV, fig. 2, 2', 2" does not correspond to any of the skins in the collection. See also Qumsiyeh (1985: 41). Benda et al. (2011f: 267) mention MNHN 1986-1068 as lectotype and indicate this is collected by E. Geoffroy Saint-Hilaire. Syntype: MNHN A.237: ad ♀. Presented/Donated by: ?: Collector Unknown. - Comments: Grubb et al. (1998: 82) mention p. 265. Kock (1969a: 122), Corbet and Hill (1992: 116) and Horácek et al. (2000: 98) only mention "Egypt" as type locality. Kock (1969a: 122) also indicates that E. Geoffroy Saint-Hilaire (1818: 127) restricted the type locality to "Grab der Könige (=Theben, Luxor)". 1855. Ph[yllorhina] tridens: Giebel, Die Säugethiere., 987. (Name Combination) 1881. Phyllorhina tridens murraiana Anderson, Catalogue of the Mammals in the Indian Museum, vol. I: 113. Type locality: Pakistan: Sind: Karachi [24 51 N 67 02 E]. 1918. Asellia tridens diluta K. Andersen, Ann. Mag. nat. Hist., ser. 9, 2 (10): 375. Publication date: 1 October 1918. Type locality: Algeria: Algerian Sahara: El Golea [30 35 N 02 53 E] [Goto Description]. Holotype: BMNH 1912.11.14.2: ♀. Collected by: Dr. Ernst Johnann Otto Hartert; collection date: 16 May 1912; original number: 42. Presented/Donated by: Nathanial Charles Rothschild. See Andersen (1918: 379). 1937. Asellia tridens pallida Laurent, Mammalia, 1: 111, fig. 8. Type locality: Morocco: Anti Atlas: Oued Tata [=Tata] [29 44 N 07 56 W] [Goto Description]. 2008. Ascidia tridens: Selim, Nahla and Shelfeh, Tishreen Univ. J. Res. Sci. Stud. - Biol. Sci. Ser., 30 (1): 253. (Name Combination, Lapsus) ? Asellia tridens murraiana: (Name Combination) ? Asellia tridens tridens: (Name Combination) ? Asellia tridens: (Current Combination) African Chiroptera Report 2020 TAXONOMY: Simmons (2005: 366) recognises three subspecies A. t. diluta Andersen, 1918 [see discussion below]; A. t. italosomalica De Beaux, 1931; and A. t. murraiana Anderson, 1881. Simmons (2005: 366) assigned A. tridens diluta to J. Anderson (1881), which would make this form Asian. However, no description is found in J. Anderson's publication, but a description is found in Andersen (1918: 379) as is confirmed by Owen and Qumsiyeh (1987 334), who mention "Andersen, 1918" as author. Owen and Qumsiyeh (1987: 335) did not examine any specimens from A. t. italosomalica De Beaux, 1931, but Benda et al. (2011f: 261) consider italosomalica to be a separate species, a view that is followed here. COMMON NAMES: Arabian: Khaffash. Czech: pavrápenec trojzubcový, wrápenec trojzeykowý, listonos trojzubcový. Dutch: Drietandbladneusvleermuis. English: Geoffroy's Lesser Tident Bat, Geoffroy's Trident Leaf-nosed Bat, Trident Leaf-nosed Bat, Trident bat, Trident Rhinolophe. French: Trident du désert, Asellia à trois endentures, Chauvesouris trident, Trident, Rhinolophe trident. German: Dreizackblattnase, Geoffroys Dreizackblattnase, Dreyzahnige Kammnase. Hebrew: ‫פרספון‬, pasafon, Parsafon. ETYMOLOGY OF COMMON NAME: So named because of the three-pronged process at the top of the nose-leaves and noticeably short ears (Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Kock et al., 2008b; IUCN, 2009; Monadjem et al., 2017m). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017m). 2008: LC ver 3.1 (2001) (Kock et al., 2008b; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004l; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. 203 MAJOR THREATS: The main threat is the widespread use of pesticides against locusts, while human disturbance in caves and old buildings is affecting some populations (Kock et al., 2008b; IUCN, 2009; Monadjem et al., 2017m). Besides water stress and their aerial hawking feeding style, roosting in caves and trees is an important risk factor linked with climatic change (Sherwin et al., 2012: 174). CONSERVATION ACTIONS: Kock et al. (2008b) [in IUCN (2009)] and Monadjem et al. (2017m) suggest underground roost management is needed in some places and a study on the impacts of pesticides is required, especially ways in which the impact might be minimised. It presumably occurs in several protected areas (Kock et al., 2008b; IUCN, 2009; Monadjem et al., 2017m). GENERAL DISTRIBUTION: Asellia tridens ranges widely in the Sahara, through the Arabian Peninsula and the Middle East, to Afghanistan, Pakistan, and India. It is absent from the northern parts of Morocco and Algeria, and Tunisia, and occurs south to Ethiopia and Somalia. Native: Afghanistan; Algeria (Kowalski and Rzebik-Kowalska, 1991: 64); Burkina Faso (Kangoyé et al., 2015a: 607); Chad; Djibouti; Egypt; Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 147); Gambia (Kock et al., 2002: 86); India (Senacha and Dookia, 2013: 21); Iran, Islamic Republic of; Israel; Libyan Arab Jamahiriya; Mali; Mauritania (Brito et al., 2010: 453, Allegrini et al., 2011: 2); Morocco (Aulagnier and Thevenot, 1986: 38; Benda et al., 2010a: 157; El Ibrahimi and Rguibi Idrissi, 2015: 360); Niger; Oman; Pakistan; Saudi Arabia; Senegal; Sudan; Syrian Arab Republic; Tunisia (Horácek et al., 2000: 99); Western Sahara. Presence uncertain: Iraq; Kuwait; Lebanon; Qatar; United Arab Emirates. A record from Zanzibar (Peters, 1871) has never been confirmed (though often repeated) and may be a MNHN mislabelled specimen. Records from Socotra and Somalia are now considered to belong to A. italosomalica (see Benda et al., 2011f). 204 ISSN 1990-6471 DETAILED MORPHOLOGY: Brain: Adult hippocampal neurogenesis was studied by Chawana R. et al. (2016: 1551). Gastro-intestinal tract (GIT): Aylward et al. (2019: 1108) reported for six specimens an average weight of 10.67 ± 0.98 g, and a GIT of 0.52 ± 0.11 g (4.83 ± 0.84 %). ECHOLOCATION: Search-phase call-shape: CF/FM. CF component is sometimes of very low amplitude, but is always present (Pye, 1972). For 11 Egyptian bats, the CF-frequency varied from 115 120 kHz; each individual emitted a narrow range of frequencies, and different individuals emitted different ranges of frequencies (ibid.). For 18 Tunisian bats, resting, CF-frequency: 111 - 124 kHz (the CF-frequency varied from bat to bat, individuals kept the frequency constant); bandwidth of terminal FM component: 19-21 kHz; callduration 7 - 10 ms; minimum intercall interval 27 215 ms (depending on degree of alertness) (Gustafson and Schnitzler, 1979). Flying bats lowered their emission frequency to compensate for Doppler shifts caused by the flight movement; call-duration: 9 - 11 ms; minimum intercall interval: 26 ms. For bats in a roost in Gambia containing thousands of individuals, CF-frequency: 108 - 122 kHz (frequency found to be negatively correlated with FA length within sexes); lowest frequency of the terminal FM sweep: 97 kHz: call-duration: 7.5 - 9 ms, inter-call interval: 10 - 20 ms (Jones et al., 1993). In the UAE, Davis (2007: 5) reports a CFfrequency with an FM-ending (117 - 124 kHz). Benda et al. (2012a: 182) provide two sets of data for Iranian A. tridens: 1: Fstart: 118.4 ± 0.6 (117.5 119.0) kHz, Fend: 117.1 ± 0.6 (116.3 - 118.0) kHz, Fpeak: 117.9 ± 0.6 (117.0 - 118.5) kHz, duration: 10.3 ± 2.1 (6.8 - 13.1) msec, and interpulse interval: 32.3 ± 15.7 (13.4 - 55.9) msec; 2 (with bat in resting position): Fstart: 122.3 ± 0.4 (121.7 122.8) kHz, Fend: 119.7 ± 4.1 (112.5 - 123.0) kHz, Fpeak: 122.0 ± 0.4 (121.3 - 122.6) kHz, duration: 9.0 ± 0.4 (8.4 - 9.3) msec, and interpulse interval: 28.0 ± 7.7 (19.3 - 38.4) msec. From Israel, the following values were reported by Hackett et al. (2016: 223) for 56 calls: Pulse duration: 7.97 ± 0.29 msec, Fstart: 117.74 ± 12.58 (114.2 - 121) kHz, Fend: 106.96 ± 10.44 (96.6 101.91) kHz, Fpeak: 116.60 ± 10.69 (107.4 - 120.1) kHz. Luo et al. (2019a: Supp.) reported the following data (Free flying bats, 2 calls): F peak: 117.9, 116.6 kHz, Fstart: 118.4, 117.74 kHz, F end: 117.1, 106.96 kHz, and duration: 10.3, 7.97 msec. Jones and Siemers (2010: 449 - 450) indicate that females emit pulses with a higher frequency than males, and also that juveniles emit lower frequencies than adults (see also Jones et al., 1993; Kunz and Hood, 2000: 420). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Baker et al. (1974) reported 2n = 50, FN = 62, a submetacentric X chromosome and an acrocentric Y chromosome. Baker et al. (1975) reported BA = 14. Protein / allozyme - Unknown. Bray and Benda (2016: 283) studied concatenated fragments of Cytochrome-b, NADH Dehydrogenase 2, and Cytochrome Oxidase 1 genes of primarily Saudi-Arabian bats, but also covered African representatives. HABITAT: Arid and semi-desert habitats. HABITS: This is a gregarious and colonial species (Kock et al., 2008b). ROOST: It roosts in temples, caves, mines, open-wells, underground irrigation tunnels and old tombs and buildings and in the desert and semi desert vegetation zones it occurs in crevices or in cliffs (Kock et al., 2008b). From Israel, Amichai et al. (2012: 96 - 97) report they use abandoned manmade structures (such as military bunkers), where the sexes are initially roosting separately. Later on (October), when their young become independent, this sexual segregation weakens. MIGRATION: In the Dakhla Oasis in Egypt, Benda et al. (2014b: 55) found A. tridens to be the most common bat during the summer months until the beginning of October, with at least 5,000 individuals divided over several roosts. However, from November to the beginning of April, these roosts were empty (except for a few single animals). In that period, only small colonies were found (70 - 120 ind.), and no animals were observed during the night. Benda et al. (2014b) were unable to determine whether the bats disappeared to the Nile Valley (long-distance migration) or whether they moved over short distances to other roosts in the neighbourhood, although there is some evidence that the latter is the case. DIET: Amichai et al. (2012: 101) report its diet in Israel to consist primarily of Coleoptera and Heteroptera, whereas Lepidoptera, Hymenoptera, Diptera were African Chiroptera Report 2020 important additions in different parts of the country. In Iran, Benda et al. (2012a: 265) report that Coleoptera - mostly Scarabaeidae, but also Cerambycidae and Curculionidae were dominatingly present, but another faecal sample was dominated by Blattodea. The most frequent representatives of Heteroptera in the diet samples collected on both occasions were bugs of the family Tingidae. In Libya, Benda et al. (2014c: 37) examined the digestive tract of one bat and found it contain 80 % (by volume) Coleoptera (Scarabeidae) and 20 % of Hymenoptera (Formicoidea). Benarbia and Fentrouci (2017: 44) found the following items in droppings of A. tridens in Algeria: Arachnida: Aranea: Araneidae: Aranea sp. Ind1; Insecta: Coleoptera: Carabidae: Megacephala australis; Chrisomilidae: Chrisomelida sp.; Curculionidae: Curculionida sp., Phyllognatnus sp.; Anthicidae: Anthicidae sp. / Anthicus sp.; Carabidae: Cicindella sp.; Tenebrionidae: Akis sp.; Aphodiidae: Aphodius sp.; Blatidae: Blatida sp.; Hymenoptera: Formicidae: Messor sp., Cataglyphis sp., Myrmecina sp., Formica sp.; Vespoidae sp.; Orthoptera: Gryllidae: Gryllus sp.; Ensiphera sp.; Orthoptera sp.; Odonatoptera; Diptera: Tabanidae sp; Ephemeroptera sp., and Plantea: Planta: Arecaceae: Phoenix dactilifera. The year-round analysis of droppings in Algeria by Loumassine et al. (2019: 354) revealed the following composition: Coleoptera (31.07 % Carabidae, Curculionidae, Chrysomelidae, Tenebrionidae, Aphodidae, Scarabaeidae, and Histeridae), Hymenoptera (21.31 % - Formicidae, Myrmicidae and Vespidae), Diptera (13.78 % Culicidae and Tabanidae), Orthoptera (11.40 % Acrididae and Gryllidae), Mantodea, Hemiptera, Isoptera, Ephemeroptera, Blattodea and Odonata. During the autum and winter, Odonata, Scarabaeidae, Blattodea, Acrididae and Myrmicidae were present in large amounts, whereas during the spring and summer Chrysomelidae, Vespidae, Tenebrionidae, Aphodidae and Araneae were more prominent. POPULATION: Structure Unknown. Density It is a very common species, found in colonies of up to several hundred in North Africa, and in its Asian distribution it has been found in groups up to 5,000 animals (Kock et al., 2008b; IUCN, 2009; Monadjem et al., 2017m). 205 Trend 2016: Stable (Monadjem et al., 2017m). 2008: Stable (Kock et al., 2008b; IUCN, 2009). 2004: Unknown (Mickleburgh et al., 2004l; IUCN, 2004). A. tridens populations within the Asia Minor and Levant region are predicted to have a stable population trend, under climate change scenarios (Bilgin et al., 2012: 433). ACTIVITY AND BEHAVIOUR: Forages over desert and semi-desert vegetation zones, mainly in oases (Kock et al., 2008b). Forages by slow hawking, has been observed foraging around palm trees and buildings, and over water (Kock et al., 2008b). REPRODUCTION AND ONTOGENY: Amichai et al. (2012: 96) report that almost all females arrive in their summer roosts in Israel at an advanced stage of pregnancy, and that births take place between the end of June and mid July. There is one young, which is born hairless, with closed eyes and weighting up to 30 % (2.5 - 3 g) of its mother's weight. In the United Arab Emirates, Davis (2007: 5) found females to be pregnant in April-May, and giving birth in June after a gestation period of about 9 to 10 weeks. The lactation period lasts for 40 days. Intrauterine ultrasonography of pregnant females performed by Amichai et al. (2019: 1) showed that the development of the horseshoe is completed (100%) in-utero, whereas skull (70%) and forearm (40%) development continues after birth. Neonates already begin emitting precursor echolocation calls two days after birth, and they are able to fly three weeks after birth. PARASITES: ACARI: Trombiculidae: Neoschoengastia elegans (Vercammen-Grandjean, Rohde and Mesghali, 1970) from Iran. Myotrombicula aselliae Vercammen-Grandjean, 1963, Riedlinia platypygia Vercammen-Grandjean, 1963, and R. afghanensis Vercammen-Grandjean, 1963, all three from Afghanistan (Benda et al., 2012a: 266267). Myobiidae: Ugandobia euthrix Fain, 1972 from Saudi Arabia (Fain 1972 [in Benda et al., 2012a: 267]) and Hipposiderobia heteronycha (Berlese et Trouessart, 1889) from Saudi Arabia and various parts of Africa (Radford 1949, Fain 1972, 1978 [in Benda et al., 2012a: 267]). Argasidae: Argas boueti Roubaud et ColasBelcour, 1933 from various countries in Africa (Roubaud & Colas-Belcour 1933, Hoogstraal 1955 [in Benda et al., 2012a: 267]). 206 ISSN 1990-6471 DIPTERA: Streblidae: Raymondia huberi Frauenfeld, 1856 from Tanzania (Haeselbarth et al., 1966: 102), and Iran (Benda et al., 2012a: 266) (see also Shapiro et al., 2016: 254). Raymondia setosa Jobling, 1930 from Israel (Haeselbarth et al., 1966: 104). Nycteriibidae: Nycteribia schmidlii Schiner, 1853 from the Levant (Benda et al., 2012a: 266). Nycteribia (Listropoda) schmidlii schmidlii Schiner, 1853 and Penicillidia (Penicillidia) dufourii Westwood, 1835 were reported by Bendjeddou et al. (2017: 15) from Algeria. Hippoboscidae: Penicillidia dufourii Westwood, 1834 from the Levant (Benda et al., 2012a: 266). Brachytarsina flavipennis Macquart, 1851 from the Levant (Benda et al., 2012a: 266). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Burkina Faso, Chad, Congo (Democratic Republic of the), Egypt, Ethiopia, Libya, Mali, Mauritania, Morocco, Niger, Saudi Arabia, Senegal, Somalia, Sudan, Tanzania, The Gambia, Tunisia. SIPHONAPTERA: Ischnopsyllidae: Chiropteropsylla brockmani Rothschild, 1915 from Iran and Iraq (Benda et al., 2012a: 266). Rhinolophopsylla unipectinata (Taschenberg, 1880) from Iraq (Benda et al., 2012a: 266). Figure 51. Distribution of Asellia tridens Genus Doryrhina Peters, 1871 *1871. Doryrhina Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 314. Publication date: 1871 [Goto Description]. - Comments: Grubb et al. (1998: 79) mention p. 324. Type species: Phyllorrhina cyclops Temminck, 1853, by monotypy (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 246).. - Etymology: From the Greek "δόρυ", meaning spear and "ρινός", meaning nose, referring to the club-shaped process which is directed forward from the base of the sella, or to the slender and somewhat longer vertical process which projects upward from the margin of the transverse erect nose leaf (see Palmer, 1904: 244). TAXONOMY: Generally considered a synonym of the genus Hipposideros, but based on Cyt-b analyses, Foley et al. (2017: 12) considered cyclops to be that different from the other members of this genus to be placed in a separate genus. Due to the generally recognized close relation between cyclops and camerunensis, Musila et al. (2018a: 21) provisionally placed the latter in the genus Doryrhina, but they also indicated that this still needs to be confirmed. MOLECULAR BIOLOGY: Karyology The genus Hipposideros s.l. shows an extreme conservatism concerning the 2n complement. Almost all species have 32 chromosomes. The only exceptions are the species transferred by Foley et al. (2017) to the genus Macronycteris, which have 52, and cyclops, which has 2n = 36. Doryrhina camerunensis (Eisentraut, 1956) *1956. Hipposideros camerunensis Eisentraut, Zool. Jb. Syst., 84 (3): 526, fig. 6 - 8. Publication date: 27 December 1956. Type locality: Cameroon: Buea, near [ca. 04 09 N 09 13 E] [Goto Description]. Holotype: SMNS 5194: ad ♂, skin and skull. Collected by: Prof. Dr. Martin Eisentraut; collection date: 29 April 1954; original number: 505. Presented/Donated by: ?: Collector Unknown. See Dieterlen et al. (2013: 294). Paratype: SMNS 5195: ad ♂, alcoholic (skull not removed). Collected by: Prof. Dr. Martin Eisentraut; collection date: 29 April 1954; original number: 510. Presented/Donated by: African Chiroptera Report 2020 2015. 2018. 2020. 207 ?: Collector Unknown. See Hutterer (1984: 29 - 30), Dieterlen et al. (2013: 294). Paratype: SMNS 5197: ad ♀, skull and alcoholic. Collected by: Prof. Dr. Martin Eisentraut; collection date: 29 April 1954; original number: 511. Presented/Donated by: ?: Collector Unknown. See Hutterer (1984: 29 - 30), Dieterlen et al. (2013: 294). Paratype: ZFMK MAM-1979.0138: ad ♀. Collected by: Prof. Dr. Martin Eisentraut; collection date: 29 April 1954; original number: 506. Exchanged from SMNS in March 1979; SMNS catalog number 5196 (see Hutterer, 1984: 29 - 30; Hutterer and Peters, 2010: 12), Dieterlen et al., 2013: 294). - Etymology: Referring to the country where the type specimen was collected (Cameroon). (Current Combination) H[ipposideros] camarunensis: Gunnell, Winkler, Miller, Head, El-Barkooky, Gawad, Sanders and Gingerich, Hist. Bio., 28 (1 - 2): 163. Publication date: 1 October 2015. (Lapsus) Doryrhina camerunensis: Musila, Monadjem, Webala, Patterson, Hutterer, De Jong, Butynski, Mwangi, Chen and Jiang, Zool. Res. (China), 40 (1): 21 (for 2019). Publication date: 17 October 2018. - Comments: Due to the view that camerunensis and cyclops are closely related, Musila et al. (2018a: 21) provisionally placed this taxon in the genus Doryrhina. (Name Combination) Doryrhina cf. camerunensis: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) TAXONOMY: Simmons (2005: 369) as part of the cyclops species group. Mickleburgh et al. (2008l) in IUCN (2009) report that there is debate as to whether all the records comprise a single species, or whether there might be two or three different species involved. The records from the Democratic Republic of Congo and Kenya could refer to other, possibly undescribed, species. Musila et al. (2018a: 21) provisionally placed camerunensis in the genus Doryrhina (instead of Hipposideros) due to the assumed close relationship of this taxon with D. cyclops. They also mentioned that this new assignment needs to be confirmed. Webala et al. (2019a: 264; 2019b: 10) mention this taxon in Doryrhina without any reservation. COMMON NAMES: Chinese: 喀 麦 隆 蹄 蝠 . Czech: pavrápenec kamerunský. English: Cameroon Leaf-nosed Bat, Greater Roundleaf Bat, Greater Cyclops, Eisentraut's Leaf-nosed Bat. French: Phyllorine du Cameroun, Phyllorine d'Eisentraut. German: Kamerun-Rundblattnase, Kamerun-Blattnase. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of continuing uncertainties as to its taxonomic status, extent of occurrence and ecological requirements (Mickleburgh et al., 2008l; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008l; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004ac; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: If it is a forest-dependent species, then it is probably suffering from ongoing forest loss. If future taxonomic studies show it to be restricted to Mount Cameroon, then it could be seriously threatened by habitat destruction (Mickleburgh et al., 2008l; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008l) [in IUCN (2009)] report that no conservation measures are currently in place. It is not known from any protected areas. Taxonomic studies are urgently needed to determine whether or not the three recorded localities all refer to the same species. GENERAL DISTRIBUTION: Doryrhina camerunensis is known only from three localities: Buea in Cameroon (type locality); Shabunda in eastern Democratic Republic of Congo; and at the Kakamega Forest in western Kenya close to Lake Victoria. If it is a single species, then it is likely to be found in future in intervening locations. Its elevational range is from sea-level to 500 m, and probably higher. 208 ISSN 1990-6471 Native: Cameroon (Eisentraut, 1956b, 1963, 1973); Congo (The Democratic Republic of the); Kenya (Schlitter et al., 1986). ECHOLOCATION: Webala et al. (2019b: 15) reported a F value of 50.8 kHz for a male and 51.1 kHz for a female the Kakamega Forest (Kenya). The duration of the calls was 24.5 to 27.7 msec for the male's call and 23.1 to 27.1 msec for the female's. Trend:- 2008: Unknown (Mickleburgh et al., 2008l; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burundi, Cameroon, Congo (Democratic Republic of the), Kenya, Tanzania, Uganda. HABITAT: The Kakamega specimen was captured in 'Intermediate Evergreen Forest' (Schlitter et al., 1986). ROOST: The type specimens were taken from a hollow tree (Eisentraut, 1963). POPULATION: Structure and Density:- Very little known about the species. It is known form only three records in a general region that is not widely surveyed (Mickleburgh et al., 2008l; IUCN, 2009). Figure 52. Distribution of Doryrhina camerunensis Doryrhina cyclops (Temminck, 1853) *1853. Phyllorrhina cyclops Temminck, Esquisses zoologiques sur la Côte de Guiné. 1e partie, les Mammifères, 75. Publication date: 1853. Type locality: Ghana: Boutry River [=Butre river] [ca. 04 49 N 01 55 W] [Goto Description]. - Comments: Type locality originally: "sur la rivière Boutry, côte de Guiné". - Etymology: Refers to the one-eyed giants of Greek mythology and alludes to the orifice in the center of the forehead of these bats (see Rosevear, 1965 in Decher and Fahr, 2005: 4). 1897. Rhinolophus micaceus de Winton, Ann. Mag. nat. Hist., ser. 6, 20 (120): 524. Publication date: 1 December 1897. Type locality: Gabon: 75 mi (112.5 km) from Gaboon: Como River [Goto Description]. Holotype: BMNH 1897.12.1.7: ♀. Collected by: George Latimer Bates Esq. Collection date: 6 July 1897; original number: GLB 230. Note: Smoked out of hollow tree. 1899. Hipposideros cyclops: de Winton, Ann. Mag. nat. Hist., ser. 7, 4 (23): 354. Publication date: 1 November 1899. (Name Combination) 1917. Hipposideros langi J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 434, text-fig. 4 - 6. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Avakubi [01 18 N 27 32 E] [Goto Description]. Holotype: AMNH 49098: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 24 January 1914; original number: 2481. 1922. H[ipposideros]. cyclops micaceus: J.A. Allen, Bull. Am. Mus. Nat. Hist., 47: 2, footnote. (Name Combination) 1922. Hipposideros cyclops langi: J.A. Allen, Bull. Am. Mus. Nat. Hist., 47: 2, footnote. (Name Combination) 2017. Doryrhina cyclops: Foley, Goodman, Whelan, Puechmaille and Teeling, Acta Chiropt., 19 (1): 12. Publication date: June 2017. (Current Combination) 2020. Doryrhina cyclops 1: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) 2020. Doryrhina cyclops 2: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) African Chiroptera Report 2020 TAXONOMY: Simmons (2005: 370) includes this taxon in the Hipposideros cyclops species group. See Decher and Fahr (2005, Mammalian Species, 763), but see also Foley et al. (2017: 12), who transferred it to the genus Doryrhina. COMMON NAMES: Chinese: 大眼蹄蝠. Czech: pavrápenec Kyklop. English: Cyclops Leaf-nosed Bat, Cyclops Roundleaf Bat, Cyclops Bat. Fang: angon. French: Phyllorine des Cyclopes, Phyllorine cyclope. German: Zyklopen-Rundblattnase, Zyklossen-Blattnase. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bw; IUCN, 2009). Assessment History Global 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bw; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cn; IUCN, 2004). 1994: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Although not threatened throughout its range, the species may disappear from many areas that are threatened by deforestation, especially on the extreme limits of its distribution (Decher and Fahr, 2005). CONSERVATION ACTIONS: Mickleburgh et al. (2008bw) [in IUCN (2009)] report that the roosting habits of this bat requires very large hollow trees, so conservation of these must be a priority. It is found in some protected areas, for example Kwamgumi Forest Reserve in Tanzania, and the Udzungwa Mountains National Park of Tanzania. GENERAL DISTRIBUTION: Doryrhina cyclops has been recorded widely over much of West Africa and Central Africa, with a few scattered records from East Africa. It ranges from Senegal and The Gambia in the west, through most countries in West Africa, to Cameroon, Equatorial Guinea (Rio Muni and Bioko), Gabon, Congo (The Democratic Republic of the) Congo, Central African Republic and Rwanda, being recorded from East Africa in southern Sudan, 209 Uganda and western Kenya, with an isolated population in southeastern Kenya and northeastern Tanzania in coastal forests and Eastern Arc Mountains (Decher and Fahr, 2005: 3). It has been collected at Tongwe Forest near Tanga and Kwamgumi Forest Reserve in Tanzania (K. Howell pers. comm. 2004). There is a dubious record from the border area of Chad and the Central African Republic. It occurs from sealevel up to at least 1,000 m. Native: Benin; Burkina Faso (Kangoyé et al., 2012: 6024; 2015a: 608); Cameroon (Hill, 1968; Eisentraut, 1973); Central African Republic (Lunde et al., 2001); Congo (Bates et al., 2013: 335); Congo (The Democratic Republic of the); Côte d'Ivoire; Equatorial Guinea (Bioko); Gabon; Gambia; Ghana (Decher and Fahr, 2007: 17); Guinea; Guinea-Bissau (Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 52); Kenya (Aggundey and Schlitter, 1984); Liberia; Nigeria (Happold, 1987); Rwanda; Senegal; Sierra Leone; Sudan; Tanzania (Decher and Fahr, 2005: 3; Stanley et al., 2005b), United Republic of; Togo; Uganda. Presence uncertain: Burundi; Chad. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data for 6 specimens: Fa: 70.63 ± 1.942 mm, mass: 37.3 ± 3.27 g, wing loading: 11.5 ± 0.53 N/m 2, aspect ratio: 5.5 ± 0.46. DETAILED MORPHOLOGY: Baculum - Unknown Brain - Amrein et al. (2007) reported from specimens collected in Bénin that proliferating cells, detected with Ki-67 and MCM2, in the subgranular layer of the dentate gyrus were found to be absent. They also reported moderate to ample proliferating cells (Ki-67 positive cells) and migrating young neurons (DCX positive cells) in the rostral migratory stream. No NeuroD was detected in the hippocampal granule cells (Amrein et al., 2007). SEXUAL DIMORPHISM: Decher and Fahr (2005) [in Kangoyé et al. (2015a: 608)] indicate that a pronounced sexual dimorphism exists, with females being larger than males. ECHOLOCATION: Monadjem et al. (2013b: 351) report a Fmax of 55.5 (55 - 56) kHz for two specimens from Mount Nimba. From western Uganda, Monadjem et al. (2011: 32) report the following values for 6 calls: Fmin: 51.24 ± 0.670 kHz, Fmax: 51.54 ± 0.387 kHz, Fchar: 51.37 ± 210 ISSN 1990-6471 0.459 kHz, Fknee: 51.51 ± 0.395 kHz, duration: 14.5 ± 2.02 msec. In Sierra Leone, Weber et al. (2019: 21) reported on two females calling at 54.5 (54.0 - 55.0) kHz. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Koubínová et al. (2010b) report for one female from Senegal, 2n= 36, Fna= 62 (as "Hipposideros cyclops") , while the chromosomal complement comprised 15 metacentric or submetacentric, and three small acrocentric pairs. Protein / allozyme - Unknown. HABITAT: Monadjem et al. (2016y: 365) recorded it from two forested sites in the Mount Nimba area, between 480 and 730 m. ROOST: Killick-Kendrick (1973: 641) reported on pygmy flying squirrels (Idiurus macrotis Miller, 1898) collected from hollow trees, which were associated with "Hipposideros cyclops". POPULATION: Structure and Density:- It is a reasonably common species in some areas (Mickleburgh et al., 2008bw; IUCN, 2009). Macronycteris cyclops is the only known host species for this parasite. Lutz et al. (2016: 9) examined three M. cyclops specimens from East Africa and found all of them infected with Nycteria sp. Perkins and Schaer (2016: Suppl.) also mention Dionisia bunoi Landau, Chabaud, Miltgen and Baccam, 1980 from bats from Gabon, Nycteria sp. from Uganda, Plasmodium cyclopsi from Gabon and Liberia. Landau et al. (1980a: 272), furthermore, mentioned a third atypical hematozoon: Trypanosoma lizae Miltgen and Landau, 1979. DIPTERA: Streblidae: Raymondia brachyphysa Jobling, 1956 from Abidjan, Côte d'Ivoire (Haeselbarth et al., 1966: 102; Shapiro et al., 2016: 254). Raymondia intermedia Jobling, 1936 in Côte d'Ivoire (Haeselbarth et al., 1966: 102). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea, Kenya, Liberia, Nigeria, Senegal, Sierra Leone, South Sudan, Tanzania, The Gambia, Togo, Uganda. Trend:- 2008: Decreasing (Mickleburgh et al., 2008bw; IUCN, 2009). REPRODUCTION AND ONTOGENY: One female, carrying a furred female infant, was captured in December 2005 at Uzungwa Scarp Forest Reserve, Tanzania (Trentin and Rovero, 2011: 50). Two lactating females were reported on 26 and 28 May from Sierra Leone by Weber et al. (2019: 24). PARASITES: HAEMOSPORIDA: Schaer et al. (2013a: 17416) report the presence of the hemosporidian parasite Plasmodium cyclopsi Landau and Chabaud, 1978 in three out of four investigated bats. Up until now, Figure 53. Distribution of Doryrhina cyclops Genus Hipposideros Gray, 1831 *1831. Hipposideros Gray, Zool. Misc., 1: 37. Publication date: 1831. - Comments: Type species: Vespertilio speoris Schneider, 1800, by subsequent designation by Sclater (1901: 166) (see Meester et al., 1986: 41; Jackson and Groves, 2015: 245). - Etymology: From the Greek "ίπποξ", meaning horse and "σίδηρος", meaning iron (=horse-iron, horseshoe), an allusion to the shape of the noseleaf (see Palmer, 1904: 327). (Current Combination) 1831. Phyllorrhina Bonaparte, Icon. Fauna Ital.. - Comments: Nomen nudum (see Palmer, 1904: 535). African Chiroptera Report 2020 1834. 1837. 1837. 1866. 1866. 1866. 1866. 1871. 1871. 1871. 1871. 1871. 1871. 211 Hipposiderus Gray, Proc. zool. Soc. Lond., 1834, II (xviii): 53. Publication date: 26 September 1834. - Comments: Unjustified emendation (see Meester et al., 1986: 41; Pavlinov et al., 1995: 84; Decher and Fahr, 2005: 1; Jackson and Groves (2015: 245). Blanford (1888: 638) claims the name Hipposiderus was published by Gray (1831: 37), but this is in error. (Emendation) Phyllorhina: Bonaparte, Iconografia Fauna italiana, 1, fasc. 21: 3. Publication date: 1837. - Comments: Type species: Rhinolophus diadema E. Geoffroy Saint Hillaire, 1813. Preoccupied by Phyllorhina Leach, 1816 (see Pavlinov et al., 1995: 84; although they do not explicitly mention its preoccupation), although also see Fenton and Rautenbach (1998: 535), who refers to Bonaparte (1837) to refute this. Fenton and Rautenbach (1998: 535) and Allen (1939a: 79) indicated that it was described as a subgenus of Rhinolophus. Etymology: From the Greek "φύλλον", meaning leaf and "ρίς" or "ρινος", meaning nose, referring to the nose-leaf (see Palmer, 1904: 535). Phyllorrhina Bonaparte, Icon. Fauna Ital., 1, fasc. 21: 3. - Comments: Type species: Rhinolophus diadema É. Geoffroy, 1813, by subsequent designation by Sclater (1901: 116) (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 245). Chrysonycteris Gray, Proc. zool. Soc. Lond., 1866, I (vi): 82. Publication date: May 1866. - Comments: Type species: Hipposideros fulvus Gray, 1838, by monotypy (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 246). - Etymology: From the Greek "χρυδός", meaning gold and "νυκτερις", meaning bat, referring to the briljant golden yellow fur (see Palmer, 1904: 189). Gloionycteris Gray, Proc. zool. Soc. Lond., 1866, I (vi): 82. Publication date: May 1866. Comments: Type species: Rhinolophus armiger Hodgson, 1835, by monotypy (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 246). - Etymology: From the Greek "γλοιός", meaning gum and "νυκτερις", meaning bat, referring to the large glandular elevations on the sides of the forehead (see Palmer, 1904: 297). Rhinophylla Gray, Proc. zool. Soc. Lond., 1866, I (vi): 82. Publication date: May 1866. Comments: Type species: Phyllorrhina labuanensis Tomes, 1859, by monotypy (see Jackson and Groves, 2015: 246). Preoccupied by Rhinophylla Peters, 1865 (Chiroptera, Phyllostomidae) (see Pavlinov et al., 1995: 84). Speorifera Gray, Proc. zool. Soc. Lond., 1866, I (vi): 82. Publication date: May 1866. Comments: Type species: Rhinolophus vulgaris Horsfield, 1823 (=Rhinolophus larvatus Horsfield). Cyclorhina Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 326. Publication date: 1871. - Comments: Type species: Phyllorrhina obscura Peters, 1861 and Phyllorrhina doriae Peters, 1871. Meester et al. (1986: 42) retain only Phyllorhina obscura Peters as type species, by subsequent designation by Moreau et al. (1946: 208), Tate (1941a: 354), Corbet and Hill (1992: 105) (see Jackson and Groves, 2015: 246). Not Cyclorhina J. Hall and Clarke, 1894 (Brachiopoda, Rhynchonellata, Rhynchonellida, Machaerariidae) (see Jackson and Groves, 2015: 246). - Etymology: From the Greek "κύκλος", meaning circle, and "ρις" or "ρινός", meaning nose (see Palmer, 1904: 208). Ptychorhina Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 325. Publication date: 1871. - Comments: Type species: Rhinolophus caffer Sundevall, 1846, by monotypy (according to Decher and Fahr, 2005: 1) or by subsequent designation by Palmer (1904: 597) (according to Jackson and Groves, 2015: 246). Sideroderma Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 324. Publication date: 1871. - Comments: Type species: Phyllorrhina fuliginosa Temminck, 1853, by monotypy (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 246). - Etymology: From the Greek "σίδηρος", meaning iron and "δέρμα", meaning skin, referring to the dark brown or reddish colour of the fur (see Palmer, 1904: 630). Sybdesmotus Peters. Syndesmotis Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 329. Publication date: 1871. - Comments: Type species: Phyllorhina megalotis Heuglin, 1862, by monotypy (see Jackson and Groves, 2015: 247). Thyreorhina Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 327. Publication date: 1871. - Comments: Type species: Phyllorrhina coronata Peters, 1871, by monotypy (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 246). - Etymology: From the Greek "θυρεός", meaning a large, oblong shield and "ρις" or "ρινός", meaning nose, referring to the upper noseleaf with thickened edge (see Palmer, 1904: 678). 212 ISSN 1990-6471 1902. 2015. 2016. 2017. 2019. ? Syndesmotus C.O. Waterhouse, Index Zoologicus. - Comments: Objective synonym of Syndesmotis Peters, 1871 (see Simmons, 2005) or incorrect subsequent spelling of Syndesmotis Peters, 1871 (see Jackson and Groves, 2015: 247). - Etymology: From the Greek "σύνδεσμος", meaning bond or fastening and "ούς" or "ώτός", meaning ear (see Palmer, 1904: 656). Hypposideros: Sylla, Pourrut, Diatta, Diop, Ndiaye and Gonzalez, Afr. J. Microbio. Res., 9 (22): 1460. Publication date: 3 June 2015. (Lapsus) Hipposieros: Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 42. Publication date: 9 May 2016. (Lapsus) Hipposidereos: Hammond, Tan and Johnson, Protein Sci., 26: 1727. Publication date: 5 June 2017. (Lapsus) Hipposederos: Zeghbib, Herczeg, Kemenesi, Zana, Kurucz, Urbán, Madai, Földes, Papp, Somogyi and Jakab, Sci. Reps., 9 (15706): 2. Publication date: 31 October 2019. (Lapsus) Hipposideros sp.: TAXONOMY: For review of the genus see Hill (1963a), on which the above synonymy and the treatment that follows are largely based, followed by Meester et al. (1986); Koopman (1993a: 170). Hill (1963a) recognizes seven species groups, five are recognized in Africa speoris, bicolor, cyclops, commersoni, and megalotis (Simmons, 2005: 367 - 379). However, see also Bogdanowicz and Owen (1998: 38), who indicate that recent species are not expressed accurately by current systematic arrangements based on the supraspecific groupings proposed by Hill (1963a), and that the genus Hippoderos might be a paraphyletic taxon. This is confirmed by Foley et al. (2014: 324), who indicate that H. commersoni and H. vittatus pose a problem here. Without these species, the monophyly of the remaining Hipposideros species is supported by all of their analyses. Foley et al. (2017: 2) furthermore indicate that the species groups, currently recognized in the genus, have yet to be thoroughly evaluated using molecular data. One of the most controversial groups in the African Hipposideros representatives is the caffer/ruber complex. Vallo et al. (2009: 196) subdivided this in four main groups, of which three are again subdivided in two lineages, and which were tentatively called: A1, A2, B1, B2, C1, C2, and D. These lineages would represent at least five species. Monadjem et al. (2013b: 345) separated two further lineages in southern African H. caffer: A1a and A2b, and two in Senegalese H. cf. ruber: D1 and D2; and one in Ghanian (E2) and Liberian and Côte d'Ivorian H. cf. ruber: (E1). Currently (Simmons and Cirranello, 2020) recognized species of the genus Hipposideros: abae J.A. Allen, 1917; alongensis Bourret, 1942 – Vietnam; armiger (Hodgson, 1835) – Northern India, Nepal, Burma, south and southeastern China, Vietnam, Laos, Cambodia, Thailand, Malay Peninsula, Taiwan (Simmons, 2005: 367); ater Templeton, 1848 – Sri Lanka; India to western Malaysia, through Philippines, Indonesia, and New Guinea to northern Queensland, north Northern Territory, and north Western Australia (Australia) (Simmons, 2005: 367); atrox K. Andersen, 1918 – Thailand, Malaysia, to Sumatra, and Teratau and Tioman Islands; beatus K. Andersen, 1906; bicolor (Temminck, 1834) – Laos, Vietnam, southern Thailand and Malaysia to Borneo and the Philippines, Java, Sumbawa, Seralu, Sumba, Savu, Roti and Timor Isls (Indonesia) and adjacent small islands (Simmons, 2005: 368); boeadii Bates, Rossiter, Suyanto and Kingston, 2007 – SE Sulawesi (Indonesia); breviceps Tate, 1941 – Mentawai Isls (Indonesia) (Simmons, 2005: 368); caffer (Sundevall, 1846); calcaratus (Dobson, 1877) – central New Guinea (Simmons, 2005: 368); cameruensis Eisentraut, 1956; cervinus (Gould, 1854) – western Malaysia, Sumatra, and Mindanao (Philippines) to the Mollucca Isls, Vanuatu, and northeastern Australia (Simmons, 2005: 369); cineraceus Blyth, 1853 – Pakistan and India to Burma, Thailand, Laos, Vietnam, Sumatra and Borneo, adjacent small islands including Kangean Isls (Indonesia) (Simmons, 2005: 369); coronatus (Peters, 1871) – northeastern Mindanao (Philippines) (Simmons, 2005: 369); corynophyllus Hill, 1985 – central New Guinea (Simmons, 2005: 370); coxi Shelford, 1901 – Sarawak (Borneo, Malaysian part) (Simmons, 2005: 370); crumeniferus (Lesueur and Petit, 1807) – Timor (Indonesia) (Simmons, 2005: 370); curtus G.M. Allen, 1921; demissus K. Andersen, 1909 – San Cristobal Isl (Solomon Isls) (Simmons, 2005: 370); diadema (E. Geoffroy Saint Hilaire, 1813) – Burma and Vietnam through Thailand, Laos, western Malaysia and Indonesia (including Sumatra, Borneo and Bali) to New Guinea, Bismarck Arch., Solomon Isls and northeastern Australia, Philippines, Nicobar Isls (Simmons, 2005: 370); dinops K. Andersen, 1905 – Solomon Isls; Bougainville Isl (Paua New Guinea) (Simmons, African Chiroptera Report 2020 2005: 370); doriae (Peters, 1871) – western Malaysia, Sarawak and Sabah (Malaysia), Borneo and Sumatra (Indonesia) (Simmons, 2005: 371); durgadasi Khajuria, 1970 – central India (Simmons, 2005: 371); dyacorum Thomas, 1902 – Borneo (including Sarawak, Malaysia), Peninsular Thailand (Simmons, 2005: 371); edwardshilli Flannery and Colgan, 1993 – northwestern Papua New Guinea (Simmons, 2005: 371); einnaythu Douangboubpha, Bumrungsri, Satasook, Soisook, Hla Bu, Aul, Harrison, Pearch, Thomas and Bates, 2011 – Myanmar; fuliginosus (Temminck, 1853); fulvus Gray, 1838 – Afganistan, India, Sri Lanka, Pakistan to Vietnam (Simmons, 2005: 371); galeritus Cantor, 1846 – Sri Lanka and India through southeast Asia (including Burma, Thailand and Peninsular Malaysia) to Java and Borneo, Sanana Isl (Sula Group, Moluccas Isls) (Simmons, 2005: 372); griffini Thong, Puechmaille, Denzinger, Dietz, Csorba, Bates, Teeling, and Schnitzler, 2012 – Vietnam; halophyllus Hill and Yenbutra, 1984 – Thailand (Simmons, 2005: 372); hypophyllus Kock and Bhat, 1994 – southern India (Simmons, 2005: 372); inexpectatus Laurie and Hill, 1954 – northern Sulawesi (Indonesia) (Simmons, 2005: 372); inornatus McKean, 1970 – Northern Territory (Australia) (Simmons, 2005: 373); jonesi Hayman, 1947; khaokhouayensis Guillén-Servent and Francis, 2006 – Laos, Vietnam; lamottei Brosset, 1985; lankadiva Kelaart, 1850 – Sri Lanka, southern and central India (Simmons, 2005: 373); larvatus (Horsfield, 1823) – northern and eastern India and Bangladesh; Yunnan, Kwangsi and Hainan (China), Burma, Thailand, Cambodia, Laos and Vietnam, western Malaysia to Sumatra, Java, Borneo and adjacent small islands including Kangean Isls (Indonesia) (Simmons, 2005: 373); lekaguli Thonglongya and Hill, 1974 – Thailand, peninsular Malaysia, Luzon (Philippines) (Simmons, 2005: 373); lylei Thomas, 1913 – Burma, Vietnam, Thailand, western Malaysia (Simmons, 2005: 374); macrobullatus Tate, 1941 – Sulawesi, Seram (Molucca Isls) and Kangean Isls (Indonesia) (Simmons, 2005: 374); madurae Kitchener and Maryanto, 1993 – Madura Isl, central Java (Indonesia) (Simmons, 2005: 374); maggietaylorae J.D. Smith and Hill, 1981 – New Guinea, Bismarck Arch (Simmons, 2005: 374); marisae Aellen, 1954; megalotis (Heuglin, 1861); muscinus (Thomas and Doria, 1886) – New Guinea (Simmons, 2005: 374); nequam K. Andersen, 1918 – Malaysia (Simmons, 2005: 375); nicobarulae Miller, 1902 – Nicobar islands; obscurus (Peters, 1861) – Philippines except Palawan region (Simmons, 2005: 375); orbiculus Francis, Kock, and Habersetzer, 1999 – Sumatra (Indonesia), Peninsular Malaysia (Simmons, 2005: 375); papua (Thomas and Doria, 1886) – Biak and Numfoor Isls, western New Guinea and northern 213 Molucca Isls (Simmons, 2005: 375); pelingensis Shamel, 1940 – Peleng Isl and Sulawesi (Indonesia) (Simmons, 2005: 375); pendlebury Chasen 1936 – S Thailand; pomona K. Andersen, 1918 – Bangladesh and India to Burma, Thailand, Laos, Cambodia, Vietnam, southern China and western Malaysia (Simmons, 2005: 375); pratti (Thomas, 1913) – S China, Burma [=Myanmar], Thailand (?), Vietnam, W Malaysia, Laos (?); pygmaeus (G.R. Waterhouse, 1843) – Philippines except Palawan region (Simmons, 2005: 376); ridleyi Robinson and Kloss, 1911 – Peninsular Malaysia, Singapore, northern Borneo (Simmons, 2005: 376); rotalis Francis, Kock, and Habersetzer, 1999 – Laos (Simmons, 2005: 376); ruber (Noack, 1893); scutinares Robinson, Jenkins, Francis, and Fulford, 2003 – Laos, Vietnam (Simmons, 2005: 376); semoni Matschie, 1903 – northern Queensland (Australia), eastern New Guinea (Simmons, 2005: 376); sorenseni Kitchener and Maryanto, 1993 – central and western Java (Indonesia) (Simmons, 2005: 376); speoris (Schneider, 1800) – India, Sri Lanka (Simmons, 2005: 377); stenotis Thomas, 1913 – Northern Territory, north Western Australia and northern Queensland (Australia) (Simmons, 2005: 377); sumbae Oei, 1960 – Sumba, Roti, Sumbawa, Flores, Semau and Savu Isls (Indonesia) (Simmons, 2005: 377); tephrus (Cabrera, 1906); turpis Bangs, 1901 – Peninsular Thailand and Vietnam; Ryukyu Isls (Japan) (Simmons, 2005: 377); wollastoni Thomas, 1913 – western and central New Guinea (Simmons, 2005: 378). Additionally, there are extinct species such as the African †besaoka Samonds 2007 and †kaumbului Wesselman 1984. COMMON NAMES: Czech: praví pavrápenci. English: Leaf-nosed Bats, Leafnosed Bats, Roundleaf Bats. French: Phyllorhines. German: Altwelt-Rundblattnasen. Italian: Ipposidèri. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The approximate fossil date for crown Hipposideros is set by Shi and Rabosky (2015: 1532) to 40.4 Mya. At Songhor, Butler (1969, 1978) found Hipposideros sp. from Early Miocene deposits. BIOGEOGRAPHY: During the Miocene at about 15 Mya, Hipposideros diversified into distinct clades based on geography: these being the African, Indian and South East Asian clades (Foley et al., 2014: 320). 214 ISSN 1990-6471 PARASITES: HAEMATOSPORIDA Schaer et al. (2017: 2) examined 23 Hipposideros specimens from South Sudan, of which 1 (4 %) was infected by Hepatocystis. VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - During Febuary 2008 in Ghana, Pfefferle et al. (2009) tested fecal samples of various Hipposideros species of the caffer/ruber complex, where the specific identification needs to be confirmed, and therefore is presented here Pfefferle et al. (2009) as Hipposideros caffer ruber tested 40 fecal samples, 10 tested positive for coronavirus (CoV) RNA, while 4 were positive for group 1 CoV and 6 for group 2 CoV. Pfefferle et al. (2009) as Hipposideros cf. ruber tested 8 fecal samples which tested negative for coronavirus (CoV) RNA, from Booyem A cave. While from Booyem B cave [identified as H cf. ruber two morphotypes], in Ghana 11 fecal samples, 2 samples tested positive for coronavirus (CoV) RNA, while the one was positive for group 1 CoV and the other for group 2 CoV (Pfefferle et al., 2009). Drexler et al. (2013a: 50) and Hu et al. (2015: 5) indicate that Hipposideros betacoronaviruses detected in Africa (Ghana, Kenya, and Nigeria) and Asia are clearly distinct from SARS-related coronaviruses. 27.6 % (?; 16 out of 68) of the Kenyan Hipposideros specimens tested by Tao et al. (2017: 3, Suppl.) were positive for CoV. Filoviridae Marburgvirus Markotter et al. (2020: 6) mention this virus from non-specified Hipposideros spp. Paramyxoviridae Mortlock et al. (2015: 1841) reported that one out of eight examined Kenyan Hipposideros sp. specimens tested positive for Paramyxovirus sequences. This was also the case for one out of 39 specimens from Cameroon, and one out of three Nigerian specimens. Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 16 Tanzanian Hipposideros sp. specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Gabon, Guinea, Kenya, Liberia, Madagascar, Namibia, Niger, Nigeria, South Sudan, Tanzania, Uganda, Zambia. †Hipposideros africanum Ravel, 2016 *2016. Hipposideros (Pseudorhinolophus) africanum Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 372, figs 8 - 10. Publication date: 30 September 2016. Type locality: Tunisia: Kassérine province: Djebel Chambi National Park: Chambi [35 14 03 N 08 45 29 E, 630 m] [Goto Description]. - Etymology: The name refers to the African continent and underlines the geographic importance as the genus was herethereto only reported from Europe (see Ravel et al., 372). 2019. Hipposideros africanuum: Brown, Cashmore, Simmons and Butler, Palaeontology, 62 (5): Suppl.. Publication date: 25 March 2019. (Lapsus) GENERAL COMMENTS: This fossil species is generally assigned to the subgenus Pseudorhinolophus Schlosser, 1887. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. †Hipposideros amenhotepos Gunnell, Winkler, Miller, Head, El-Barkooky, Gawad, Sanders and Gingerich, 2015 *2015. Hipposideros (Pseudorhinolophus) amenhotepos Gunnell, Winkler, Miller, Head, El-Barkooky, Gawad, Sanders and Gingerich, Hist. Bio., 28 (1-2): 162, figs 4, 5, 6 (for 2016). Publication date: 1 October 2015. Type locality: Egypt: Eastern Desert: Khasm El-Raqaba [28.451 N 31.834 E] [Goto Description]. Holotype: CGM 83763: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Right M1. - Comments: The paper was published online on 1 October 2015, and in print on 17 February 2016. - Etymology: Named after Amenhotep IV (= Akhenaten), an Egyptian 18th Dynasty pharaoh, whose capital city of Amarna was located in the Eastern Desert not far from the type locality of this species (see Gunnell et al., 2015b: 162). African Chiroptera Report 2020 PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: 215 Pliocene (Tortonian - Brown et al., 2019: Suppl.). †Hipposideros kaumbului Wesselman, 1984 *1984. Hipposideros kaumbului Wesselman, Contributions to vertebrate evolution, 7: 59. Type locality: Ethiopia: Locality 28, lower Member S, Shungura Formation. - Etymology: This taxon is named in honour of John Kaumbulu Kimau, a Wakamba tribesman from Kikoko, Kenya, who worked with Henry ("Hank") Barnard Wesselman as his field assistant following the 1972 and 1973 field seasons and helped with the task of handpicking almost 20 tons of screenwashed residues (Wesselman, 1984: 59). (Current Combination) TAXONOMY: Wesselman (1984: 59) suggests that this taxon may be ancestral to "Hipposideros commersoni" (=Macronycteris commersoni. Rakotoarivelo et al. (2019: 14), however, indicate that many different taxa included in the commersoni) group are morphologically very similar to each other, yet genetically quite distinct PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late Pliocene (2.08 Mya) (Wesselman, 1984: 59; Brown et al., 2019: Suppl.). GENERAL DISTRIBUTION: Ethiopia, Kenya. †Hipposideros vetus (Lavocat, 1961) *1961. Asellia vetus Lavocat, Notes Mém. Serv. Mines Carte geol. Maroc, 155: ?. - Comments: Marandat (1994: 61) mentions that the holotype consists of a left maxillary fragment with P 4-M3. 1982. Hipposideros (Syndesmotis) vetus: Legendre, J. Vert. Paleont., 2 (3): 379. Publication date: December 1982. (Name Combination) 2013. Asellia/Hipposideros vetus: Stoetzel, Palaeog., Palaeoclim., Palaeoecol., 392: 363. Publication date: 29 September 2013. (Name Combination) ? Hipposideros vetus: (Current Combination) TAXONOMY: Originally described in the genus Asellia, the generic assignment of this taxon was unclear. Some authors (e.g. Legendre, 1982: 372; Gunnell et al., 2011: 71; Hugueney et al., 2015) assign it to the genus Hipposideros, whereas others (e.g. Aulagnier, 2013a: 360) assign it to the genus Asellia. Stoetzel, 2013: 363) furthermore mentions it as Asellia/Hipposideros vetus. Legendre (1982: 382) examined the teeth and dental characters and found that although the absence of a P 2 and the degree of reduction of M2 as well als M3 are similar to those of other fossil Asellia, there are characters that are incompatible with this genus: upper canines with a separate secondary cusp on the distal edge, mandible with a long and slender horizontal ramus; low coronoid process; long and outward deflected angular process. These characters are much more similar to those of Hipposideros. Legendre (1982: 383) assigned vetus to the subgenus Syndesmotis, together with H. megalotis. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Miocene - Pliocene (Langhian, Serravallian - Brown et al., 2019: Suppl.). Hipposideros abae J.A. Allen, 1917 *1917. Hipposideros abæ J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 432. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Aba [03 53 N 30 17 E] [Goto Description]. Holotype: AMNH 49123: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 13 December 1911; original number: 1715. Etymology: Referring to the locality where the type specimen was collected (Aba). (Current Combination) 216 ISSN 1990-6471 ? Hipposideros abae: (Current Spelling) TAXONOMY: Simmons (2005: 367) places it in the speoris species group. Patterson et al. (2020: 121) place it in the ruber subgroup of the bicolor group. COMMON NAMES: Czech: pavrápenec abánský. English: Aba Roundleaf Bat, Aba Leaf-nosed Bat. French: Phyllorhine d'Aba. German: Aba-Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008at; IUCN, 2009; Monadjem et al., 2017aq). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017aq). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008at; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bq; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There are no known major threats to this species. It appears to have taken advantage of forest clearance by moving into formerly forested land (Mickleburgh et al., 2008at; IUCN, 2009; Monadjem et al., 2017aq). CONSERVATION ACTIONS: Mickleburgh et al. (2008at) [in IUCN (2009)] and Monadjem et al. (2017aq) report that no specific conservation measures are in place, but it occurs in Comoe National Park in Côte d'Ivoire, and presumably also in other protected areas across its wide range. GENERAL DISTRIBUTION: Hipposideros abae is patchily distributed in a range which extends from Guinea-Bissau in West Africa (also the most northerly record) to northern Uganda (also the most eastern and southerly record) (Simmons, 2005: 367). It is a lowland species found below 1,000 m. Native: Burkina Faso (Koopman et al., 1978: 4; Kangoyé et al., 2015a: 608); Cameroon; Central African Republic; Congo (The Democratic Republic of the); Côte d'Ivoire; Ethiopia (Kruskop et al., 2016: 57); Ghana; Guinea (Weber and Fahr, 2007); Guinea-Bissau (Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 56); Liberia; Nigeria; Sierra Leone; Sudan; Togo; Uganda (Kityo and Kerbis, 1996: 61). Presence uncertain: Benin; Chad; Mali. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Young (1975: 76) indicates that two distinct colour phases can be found: rufous (orange) and grey. ECHOLOCATION: In Sierra Leone, Weber et al. (2019: 21) recorded the call of one female with CF at 103.0 kHz. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Unknown. Protein / allozyme - Unknown. ROOST: It roosts in caves and rocky crevices or on the undersides of boulders, but apparently not in houses (Happold, 1987). For Guinea, Weber and Fahr (2006: 4) indicate that H. abae largely or exclusively depends on the availability of caves as day roosts. POPULATION: Structure and Density: Is thought to be sparsely distributed throughout its range, and to roost in small colonies (approximate group size in the dozens, not hundreds) (Mickleburgh et al., 2008at; IUCN, 2009; Monadjem et al., 2017aq). Trend:- 2016: Unknown (Monadjem et al., 2017aq). 2008: Unknown (Mickleburgh et al., 2008at; IUCN, 2009). REPRODUCTION AND ONTOGENY: One female captured on 24 May in Sierra Leone was lactating (Weber et al., 2019: 23). PARASITES: ACARI Fain (1959c: 239) described Psorergates (Psorergatoides) hipposideros (Acari: Psoridatidae) from a H. abae captured near Bunia, Ituri, DRC. DIPTERA Streblidae: Ascodipteron jonesi Jobling 1952 from the wing at Njala, Sierra Leone (Haeselbarth et al., 1966: 106). African Chiroptera Report 2020 VIRUSES: Coronaviridae Alphacoronavirus: SARS-CoV (subgenus Duvinacovirus) - During Febuary 2008 in Ghana, 16 fecal samples for coronavirus (CoV) RNA from two localities tested negative Pfefferle et al., 2009). Corman et al. (2015: 11859) tested 242 Ghanese bats and found 19 (7.8 %) to test positive for 229E-related coronaviruses. 217 de Jong et al. (2011: 11), Luis et al. (2013: Suppl.), and Willoughby et al. (2017: Suppl.) report the occurrence of Rift Valley fever virus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Ghana, Nigeria, Senegal, Sierra Leone, South Sudan, Togo. Hepeviridae Drexler et al. (2012b: 9137) reported two positive feacal samples out of a total of 57 examined (3.51 %) from Ghana. Johne et al. (2014: 223, 224) suggested to place the Bat hepatitis E virus found in H. abae, as well as Hepevirus-species occuring in other bat families into a separate genus: Chiropteranhepevirus. Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) reported that none of the 80 specimens they examined from Ghana tested positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. Phenuiviridae Phlebovirus Figure 54. Distribution of Hipposideros abae Hipposideros beatus (K. Andersen, 1906) *1906. Hipposiderus beatus K. Andersen, Ann. Mag. nat. Hist., ser. 7, 17 (99): 275, 279. Publication date: 1 March 1906. Type locality: Equatorial Guinea: Rio Muni: Benito River, 15 mi (24 km) from [01 31 N 09 52 E] [Goto Description]. Holotype: BMNH 1900.2.5.45: ad ♀, alcoholic (skull not removed). Collected by: George Latimer Bates Esq. Collection date: February 1899. (Current Combination) 1957. Hipposideros beatus maximus Verschuren, Exploration du Parc national du Garamba, 7: 365. Type locality: Congo (Democratic Republic of the): Garamba National Park: Pidigala-Nord, Source area of [04 38 N 29 42 E]. Holotype: RBINS 13768: ad ♂. Collected by: Jacques Verschuren; collection date: 23 April 1952; original number: 4565. Presented/Donated by: ?: Collector Unknown. Holotype: RBINS 4037: ad ♂. Collected by: Gaston-François de Witte et al.; collection date: 23 April 1952; original number: 4565. Presented/Donated by: ?: Collector Unknown. 2020. Hipposideros beatus 1: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) 2020. Hipposideros beatus 2: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) ? Hipposideros beatus beatus: (Name Combination) ? Hipposideros beatus: (Current Spelling) TAXONOMY: Simmons (2005: 368) places it in the bicolor species group, and Patterson et al. (2020: 121) in the ruber subgroup of the bicolor group. COMMON NAMES: Castilian (Spain): Murciélago de Nariz de Hoja. Czech: pavrápenec guinejský. English: Benito Roundleaf Bat, Benito Leaf-nosed Bat, Dwarf Leafnosed Bat. French: Phyllorine naine, Phyllorhine 218 de Benito. German: Benito-Blattnase. ISSN 1990-6471 Benito-Rundblattnase, CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bz; IUCN, 2009; Monadjem et al., 2017bm). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bm). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bz; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ck; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Habitat loss is the major threat to this species (Mickleburgh et al., 2008bz; IUCN, 2009; Monadjem et al., 2017bm). CONSERVATION ACTIONS: Mickleburgh et al. (2008bz) [in IUCN (2009)] and Monadjem et al. (2017bm) report that it occurs in the Mount Allen National Park (Equatorial Guinea), and Garamba National Park (Democratic Republic of Congo), and presumably in several other protected areas. GENERAL DISTRIBUTION: Hipposideros beatus has been recorded from Sierra Leone eastwards across the forest zone west and central Africa, as far east as the Sudan/Democratic Republic of Congo border. There are records from Sierra Leone, Guinea, Liberia, Côte d'Ivoire, Ghana, Togo, Nigeria, Cameroon, Equatorial Guinea, Congo, Democratic Republic of Congo and Sudan. A previous report of this species in Central African Republic is in error, though it might occur in the south of that country. Native: Cameroon (Eisentraut, 1973); Congo (The Democratic Republic of the); Côte d'Ivoire; Equatorial Guinea; Gabon; Ghana (Decher and Fahr, 2007: 16); Guinea; Guinea-Bissau (Rainho and Ranco, 2001: 59); Kenya (Webala et al., 2019b: 13); Liberia; Nigeria (Happold, 1987); Sierra Leone; Sudan; Togo. Presence uncertain: Central African Republic. Bates et al. (2013: 338) reject the presence of H. beatus in the Republic of Congo, because previous authors (e.g. Malbrant and Maclatchy, 1949; Dowsett et al., 1991) indicated it would be 'likely' to occur in that country. ECHOLOCATION: Monadjem et al. (2013b: 351) report the Fmax to be 129.0 kHz at Mount Nimba. Heller and Volleth (2016: 4) report different bandwidths for H. beatus specimens from various parts of Africa: Côte d'Ivoire: 139 - 147 kHz (Happold, 2013ap), Mount Nimba: 129 kHz (Monadjem et al., 2013b), Lake Tumba, DRC: 108 and 129 kHz (2 specimens; see Novick, 1958), and Irangi, DRC: 109.2 kHz (their study). This 20 kHz difference makes them suggest that two cryptic species might be present. In Kenya, Webala et al. (2019b: 15) report Fmax: 114.8 - 123.7 kHz, duration: 7.0 - 9.1 msec for three females and Fmax: 124.0 kHz, duration: 8.0 8.4 msec for two males. HABITAT: In Nigeria, it is widely distributed in the rainforest zone (Happold, 1987). Monadjem et al. (2016y: 365) also report its wide distribution in forested habitats in the Mount Nimba area, on altitudes between 420 and 700 m. POPULATION: Structure and Density:- It has a patchy distribution, typically found in small familial groups, but can be found in groups of up to 20 individuals. There is no information on overall abundance (Mickleburgh et al., 2008bz; IUCN, 2009; Monadjem et al., 2017bm). Trend:- 2016: Decreasing (Monadjem et al., 2017bm). 2008: Decreasing (Mickleburgh et al., 2008bz; IUCN, 2009). VIRUSES: Poxviridae Orthopoxvirus Mande (2019: 7) reported a seropositive case of Mokeypoxvirus in a H. beatus from the DRC. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea, Kenya, Liberia, Nigeria, Senegal, Tanzania, Uganda. African Chiroptera Report 2020 219 Figure 55. Distribution of Hipposideros beatus Hipposideros caffer (Sundevall, 1846) *1846. Rhinolophus caffer Sundevall, Öfvers. kongl. Sv. Vet.-Akad. Förhandl., 3 (4): 118. Publication date: 1846. Type locality: South Africa: KwaZulu-Natal: Port Natal [=Durban], near [ca. 29 52 S 31 00 E] [Goto Description]. Syntype: BMNH 1848.6.2.16: skull only. "Cotype" from Stockholm Museum, labeled "Hipposideros caffer typicus" locality Natal. Syntype: BMNH 1849.11.22.11: skin only. - Comments: Type locality originally mentioned as "Circa Port-Natal inventus". - Etymology: From the masculine scientific Latin adjective càffer "Kafir", referring to the southern African locality where the type specimen was collected (see Lanza et al., 2015: 139). 1852. Phyllorrhina caffra Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 39, pl. 8. Publication date: 1852. Type locality: Mozambique: Ibo island. (Name Combination, Alternate Spelling) 1852. Phyllorrhina gracilis Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 36, pl. 7, fig. 1 - 4; pl. 13, fig. 14 - 15. Publication date: 1852. Type locality: Mozambique: Lower Zambesi River: Tete [16 10 S 33 35 E, 230 m] [Goto Description]. Holotype: ZMB 364/85624: ♂. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847; original number: An 16311. See Turni and Kock (2008: 38) [skin & skull: ZMB 364 + skeleton: ZMB 85624]. 1852. Phyllorrhina patellifera Peters, Naturw. Reise Mossamb., Säugethiere, p. 39. Type locality: Mozambique: Ibo island. - Comments: Manuscript name, cited by Peters (1852: 39, 40). 1898. Phyllorhina angolensis Seabra, J. Sci. mat. phys. nat., ser. 2, 5: 256. Publication date: December 1898. Type locality: Angola: Rio Coroca [=Curoca river] [15 43 S 11 55 E]. Syntype: BMNH 1906.1.3.2:. - Comments: Meester et al. (1986: 43) indicate "Benguela, Angola, restricted by Hill and Carter (1941: 40), who point out that the name is based on four specimens from Benguela, housed in the Museu Bocage, Lisbon, and one from Rio Coroca, Angola, in the British Museum (Natural History). Koopman (in litt.) points out that, as the Museu Bocage had been destroyed by fire, the Rio Coroca specimen may be the only remaining of the type series.". 1917. Hipposideros nanus J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 434. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Faradje [03 44 N 29 43 E] [Goto Description]. Holotype: AMNH 49426: ad ♀, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 24 September 1912; original number: 1689. Allen (1917: 434) erroneously mentioned 24 October 1912 as collection date, see Goodwin (1953: 245). 1924. Hipposideros caffer aurantiaca de Beaux, Atti Soc. Lig. Sci. Nat. Geogr, 3: 155. Type locality: Somalia: Basso Giuba [=Lower Juba river]: Belet Mamu. - Comments: Lanza et al. (2015: 139) indicate that the correct spelling shoud actually be "aurantiacus" as the generic name ("Hipposideros") is a masculine name. 220 ISSN 1990-6471 2015. 2016. 2020. 2020. 2020. 2020. 2020. 2020. 2020. 2020. ? ? ? ? ? H[ipposideros] cafer: Gunnell, Winkler, Miller, Head, El-Barkooky, Gawad, Sanders and Gingerich, Hist. Bio., 28 (1 - 2): 163. Publication date: 1 October 2015. (Lapsus) Hipposideros cf. caffer Monadjem, Richards and Denys, Acta Chiropt., 18 (2): 365. Publication date: December 2016. - Comments: . Hipposideros caffer 1: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicate that this is the same form as the Hipposideros caffer A2 lineage mentioned by Vallo et al. (2009) and Hipposideros caffer tephrus mentioned by Monadjem et al. (2013b). (Name Combination) Hipposideros caffer 2: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) Hipposideros caffer 3: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) Hipposideros caffer 4: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicate that this is the same form as the Hipposideros caffer A1 lineage mentioned by Vallo et al. (2009) and the A1a and A1b lineages mentioned by Monadjem et al. (2013b). (Name Combination) Hipposideros caffer 5: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicate that this is the same form as the Hipposideros caffer B lineage mentioned by Vallo et al. (2009). (Name Combination) Hipposideros caffer 6: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) Hipposideros caffer 7: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicates that this is the same form as the Hipposideros ruber B2 lineage mentioned by Monadjem et al. (2013b). (Name Combination) Hipposideros caffer 8: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) Hipposideros caffer angolensis: (Name Combination) Hipposideros caffer caffer: Hipposideros caffer centralis: Hipposideros caffer nanus: (Name Combination) Hipposideros caffer: (Name Combination, Current Combination) TAXONOMY: 1978; Corbet and Hill, 1980, Koopman, 1982). Although Hill (1963a) and Koopman (1975 regard ruber as only subspecifically distinct from caffer. Simmons (2005: 368) places H. caffer within the bicolor species group. Vallo et al. (2007: 257) consider tephrus to be a separate species, based on cytochrome b analysis. Reviewed by Vallo et al. (2009). Patterson et al. (2020: 121) place it in the ruber subgroup of the bicolor group. Figure 56. Hipposideros caffer (yellow form). Meester et al. (1986) state that further extralimital [to Southern Africa] subspecies may include centralis Andersen, 1906, tephrus Cabrera, 1906, nanus J.A. Allen, 1917 (Koopman, 1975), and also guineensis Andersen, 1906, and niapu J.A. Allen, 1917 (Hill, 1963a). Most recent authors appear to regard ruber Noack, 1893, as a separate species (Kock, 1969a; Hayman and Hill, 1971; Ansell, COMMON NAMES: Afrikaans: Sundevall se bladneusvlermuis, Sundevall-bladneusvlermuis, Sundevallse blaarneusvlermuis, Kaapse Blaarneusvlermuis. Azande (DRC): Nkulo. Chinese: 松 氏 蹄 蝠 . Czech: pavrápenec natalský, listonos africký, pavrápenec africký. English: Sundevall's Leafnosed Bat, Common African Leaf-nosed Bat, Lesser Leaf-nosed bat, Sundevall's Roundleaf Bat, Cape Leaf-nosed bat, Sundevall's African Leaf-nosed Bat, South African Lesser Leaf-nosed African Chiroptera Report 2020 Bat. French: Phyllorhine de Cafrerie, Phyllorhine de Sundevall, Rhinolophe de Cafrerie. German: Sundevalls Rundblattnase, Sundevall's Blattnase, Gewöhnliche Rundblattnase. Italian: Ipposìdero càfro. Kiluba (DRC): Kasusu. Nyungwe (Malawi): Kalemawalema (not specific). Portuguese: Morcego de nariz enfolhado da cafraria. Ukrainian: Листоніс південноафриканський [= Lystonis pivdennoafrykans'kyy]. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category (Kock et al., 2008c; IUCN, 2009). Assessment History Global 2008: LC ver 3.1 (2001) (Kock et al., 2008c; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ae; IUCN, 2004). 1994: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016t). 2004: DD ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Human disturbance to roosting sites (caves) could have a negative effect (Kock et al., 2008c; IUCN, 2009). Sherwin et al. (2012: 174) consider aerial hawking and water stress to be the major risk factors linked to climatic change. CONSERVATION ACTIONS: Kock et al. (2008c) [in IUCN (2009)] report that no specific conservation measures are in place. Occurs in protected areas throughout its range. GENERAL DISTRIBUTION: Hipposideros caffer is a very wide ranging species, occurring from the south-western Arabian Peninsula (including Yemen) and across most of sub-Saharan Africa (except for central forested regions) (Simmons, 2005: 368). This species has also been recorded in southern Algeria, central Niger, eastern Chad, the Senegal/Mauritania border. Elevation ranges from sea level to 2,500 m in this area. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,581,888 km 2. In South Africa, its 221 distribution is stongly associated with the mean temperature of the coldest quarter (Babiker Salata, 2012: 49). Specimens from Morocco need verification as these may be attributed to H. tephrus (Cabrera, 1906). A supposed record from southern Algeria requires confirmation. Specimens identified by Fahr et al. (2006a) as H. caffer from Mount Nimba might need to be reassigned, as this species is restricted to southern Africa (Monadjem et al., 2016y: 365). Native: Angola (Crawford-Cabral, 1989; Cotterill, 2004a: 261; Monadjem et al., 2010d: 536); Benin; Botswana (Monadjem et al., 2010d: 536); Burkina Faso; Burundi; Cameroon; Chad; Congo (Bates et al., 2013: 335); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 536); Côte d'Ivoire; Equatorial Guinea; Eritrea; Ethiopia; Gabon (Brosset, 1966c); Gambia; Ghana; Guinea; Guinea-Bissau (Seabra, 1900c; Monard, 1939;; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 50); Kenya; Malawi (Happold et al., 1988; Monadjem et al., 2010d:536); Mali; Mauritania; Morocco (El Ibrahimi and Rguibi Idrissi, 2015: 360); Mozambique (Lopes and Crawford-Cabral, 1992; Monadjem et al., 2010d: 536; Monadjem et al., 2010c: 380); Namibia (Monadjem et al., 2010d: 536); Niger; Nigeria (Menzies, 1973); Rwanda; Saudi Arabia; Senegal; Sierra Leone; Somalia; South Africa (Taylor, 2000: 140; Monadjem et al., 2010d: 537); Sudan; Swaziland (Monadjem et al., 2010d: 537); Tanzania (including the islands of Pemba, Unguja, and Mafia [see O'Brien, 2011: 287]); Togo; Uganda (Kityo and Kerbis, 1996: 61); Yemen; Zambia (Ansell, 1969; Ansell, 1978; Ansell, 1986; Cotterill, 2002b: 5; Monadjem et al., 2010d: 537); Zimbabwe (Monadjem et al., 2010d: 537). Presence uncertain: Algeria; Liberia. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Menzies (1973) [in Young (1975: 76) indicates that, in some colonies in northern Nigeria, rufous bats get a grey fur during the molting season, and subsequently change back to rufous. Young (1975: 76) also mentions that the majority of H. caffer specimens in Angola and south-west Africa (Namibia) is grey, whereas in eastern Africa, the rufous phase is more common. From western Uganda, Monadjem et al. (2011: 30) reported the following data for 5 H. cf. caffer specimens: Fa: 51.50 ± 1.398 mm, mass: 10.2 ± 1.10 g, wing loading: 5.2 ± 0.27 N/m 2, aspect ratio: 5.8 ± 0.10. 222 ISSN 1990-6471 DETAILED MORPHOLOGY: Baculum - Unknown Brain - Amrein et al. (2007), from specimens collected from Bénin, reported moderate proliferating cell activity, detected with Ki-67 and MCM2, in the subgranular layer of the dentate gyrus, with doublecortin (DCX) they detected highest levels of young migrating neuron in the subgrannular layer of the hippocampus. Amrein et al. (2007) also reported moderate to ample proliferating cells (Ki-67 positive cells) and migrating young neurons (DCX positive cells) in the rostral migratory stream. No NeuroD was detected in the hippocampal granule cells (Amrein et al. (2007). ECHOLOCATION: At Irangi (Kivu province, DRC), Heller (1992: 77) was able to distinguish H. caffer from H. ruber as the prior emitted calls at a much higher frequency: 137 - 160 kHz (versus 130 - 140 kHz in ruber). In Guinea, Fahr and Ebigbo (2003) reported a constant frequency component of 160.2 - 164.1 kHz (these may not represent H. caffer see Vallo et al., 2009). In Kenya, Taylor et al. (2005) reported a maximum frequency of 157 (± 0.2) kHz, using a Pettersson D980 detector. From the same country, Eisenring et al. (2016: SI 2.) reported the following values: PF: 144.0, HF: 149.4, LF: 110.9, DT: 0.1, DF: 38.5, and IPI: 0.0. At Sengwa in Zimbabwe, Fenton (1986a) reported a maximum frequency of 141.5 (± 2.7) kHz, whereas previously Jacobs (1996) and Fenton and Bell (1981) had reported a maximum frequency of 138 kHz from the same locality. In South Africa, Fenton (1986a) reported a maximum frequency of 145.4 (± 2.5) kHz from Levuvhu, while Taylor (1999b) using a Pettersson D980 detector reported a similar maximum frequency 145 (± 0.2) kHz from Jozini Dam. Monadjem et al. (2007a), using an Anabat detector, reported a maximum frequency of 143.0 (± 1.43) kHz from various localities in Swaziland. See also Pye (1972, O'Shea and Vaughan, 1980). Monadjem et al. (2010c: 381) reported from Mozambique that the peak echolocation frequencies of two individuals were recorded at 145 kHz (ANABAT). Monadjem et al. (2007a) also reported sexual dimorphism in the call with males emitting a higher call than females (males 145.0 (± 0.76) kHz, females 142.9 (± 1.39) kHz). For 13 calls from Swaziland, Taylor et al. (2013b: 19) reported the following parameters: Fmax: 142.9 ± 0.96 (140.4 - 144.1) kHz, Fmin: 123.2 ± 6.40 (108.9 - 130.3) kHz, Fknee: 141.5 ± 0.95 (139.3 142.9) kHz, Fchar: 142.0 ± 1.00 (139.8 - 143.5) kHz, duration: 4.2 ± 0.28 (3.7 - 4.7) msec. Monadjem et al. (2017c: 180) indicated that calls from this species could not be detected at a distance of more than 0.5 m, using an Anabat detector. They also reported the following values: Fmin: 121.2 ± 5.88 (109.1 - 127.3) kHz, Fknee: 141.4 ± 1.97 (138.7 - 144.3) kHz, Fc: 141.7 ± 1.69 (139.5 - 144.0) kHz, duration: 4.8 ± 1.08 (3.7 - 6.7) msec. Monadjem et al. (2011: 32) reported on 3 calls from Hipposideros cf. caffer from western Uganda: Fmin: 144.11 ± 6.373 kHz, Fmax: 151.73 ± 1.661 kHz, Fchar: 146.88 ± 3.909 kHz, Fknee: 151.13 ± 1.650 kHz, duration: 2.8 ± 1.18 msec. In the Mapungubwe National Park (RSA), Parker and Bernard (2018: 57) recorded the following parameters: Fchar: 143.00 ± 1.64 kHz, Fmax: 143.28 ± 1.63 kHz, Fmin: 124.99 ± 5.92 kHz, F>sub knee: 143.14 ± 1.64 kHz, duration: 7.41 ± 0.91 msec, with 12.29 ± 7.81 calls/sec. In Kenya, Webala et al. (2019b: 15) reported echolocation data from nine localities, where F max varied between 148.9 and 153.6 for seven unsexed bats, 145.0 and 155.8 for eight males, and 145.5 and 150.9 for ; 5.0 - 9.7 for six males. The call durations varied respectively between 6.7 and 9.7, 5.4 and 10.6, and 5.0 and 9.7 msec. Luo et al. (2019a: Supp.) reported the following data (Free flying bats): Fpeak: 145.4 kHz and duration: 8 msec. MOLECULAR BIOLOGY: DNA - Vallo et al. (2009). Karyotype - Ðulic and Mutere (1974), Peterson and Nagorsen (1975), Rautenbach et al. (1993) and Porter et al. (2010) all reported 2n = 32, FN = 60, and BA = 30. Peterson and Nagorsen (1975) and Rautenbach et al. (1993) also agreed on an acrocentric Y chromosome. However, Ðulic and Mutere (1974) reported a metacentric X chromosome, Peterson and Nagorsen (1975) reported a subtelocentric X chromosome, and Rautenbach et al. (1993) reported a submetacentric X chromosome. Protein / allozyme - Unknown. HABITAT: This species occurs in savanna, bushveld and coastal forest, and is usually associated with rivers and other water resources (Taylor, 2000: 140). ROOST: Weber and Fahr (2006: 4) indicate that H. caffer, in Guinea, largely or exclusively depends on the availability of caves as day roosts. In the Durban area, Taylor et al. (1999: 70) reported it roosting in a tunnel of the Songweni Dam. African Chiroptera Report 2020 DIET: In their study of H. caffer bats from Skukuza (Kruger National Park, RSA), Dunning and Krüger (1996: 708) found that they overwhelmingly fed upon tympanate Lepidoptera, whereby relative numbers of noctuid, pyralid, and arctiid moths taken by the bats were proportional to the representation of these families in the general population of moths in the area. They also took much less moths belonging to the Geometridae, and arctiid moths that were able to produce clicking sounds. Taylor et al. (2017b: 245, 252 - 254) reported the diet from bats from the Limpopo province (RSA) to include: one unidentified beetle and seven genera or species of moths belonging to four families: Sena sp., Mimallo amilia DHJ01, Xenopseustis sp., Pericyma atrifusa, Tycomarptes sp., Maurilia arcuata. From Sudwala (RSA), Jacobs (2000: 201) reported the following proportions of preys: Lepidoptera (65.6 ± 46.3), Coleoptera (19.1 ± 40.0), Isoptera (14.5 ± 35.6), Hemiptera (0.5 ± 1.2) and unknwon (0.3 ± 0.6). PREDATORS: Mikula et al. (2016: Supplemental data) mention H. caffer as a possible prey of the Lanner falcon (Falco biarmicus Temminck, 1825), Peregrine falcon (Falco peregrinus Tunstall, 1771) and Bat hawk (Macheiramphus alcinus Bonaparte, 1850). POPULATION: Structure and Density:- Are extremely gregarious and capable of forming huge colonies where there is adequate roosting space. In Nigeria, at Shagunu (Menzies, 1973), a colony of about 1,000 individuals inhabited a cave five meters wide, 1-2 m high and 10-15 m deep. In Gabon they live in colonies of 200-1,000 and one exceptionally large cave contained an estimated 500,000 bats (Brosset, 1966c; Happold, 1987). Trend:- 2008: Decreasing (Kock et al., 2008c; IUCN, 2009). REPRODUCTION AND ONTOGENY: For Uganda, Mutere (1968) reported pregnancies between December and March and births in April. Bernard and Meester (1982: 131) studied the reproductive cycle in H. caffer in Natal and found that follicular development began in February, with copulation and ovulation taking place from late April onwards. The duration of the gestation period was about 220 days (zygote: 21 days, embryo: 123 days and foetus: 76 days). Parturition took place in December, after which 223 there was a lactation period until January, during which the bats were anoestrus. Menzies (1973) claimed it can take up to two months before a blastocyst gets implanted, but this was revoked by Racey (1982: 83), who indicated that some of Menzies' illustrations proved otherwise. In NE Gabon, Brosset and Saint Girons (1980: 226) found that H. caffer was reproducing with a boreal cycle (giving birth in March) in some caves, whereas in others they were following an austral cycle (giving birth in October). A line separating both groups seemed to run just north of the equator. For the northern group, mating occurred in November-December. In both groups, the single young was weaned after three and a half months. In the caves, where the females were following a boreal cycle, the males were in full spermatocytogenesis at the beginning of October. The spermiogenesis was starting in that period and this phase ended between 25 and 30 November. In the other caves, the males were in sexual rest between August and January. Meinig (2010: 391) indicates that the gestation period differs according with latitude. In bats from South Africa it would last for seven months, whereas for tropical bats, it would take only four months. PARASITES: Adam and Landau (1973a: 5) report that H. caffer was rarely infected by a protozoan parasite of the genus Polychromophilus (Haemoproteidae). The bats were also found to carry low numbers of the the nycteribiid flies Penicillidia fulvida Bigot, 1885 and Nycteribia schmidlii scotti Falcoz, 1923, of which the prior was a major carrier of Polychromophilus sporozoites. Perkins and Schaer (2016: Suppl.) reported the presence of Polychromophilus murinus (Dionisi, 1899). Wünschmann et al. (2010: 178) refer to Boulard (1975), who reported the first case of renal coccidiosis in a bat (on H. caffer and Rhinolophus sp.) from Sierra Leone and the Congo, and caused by Klossiella killicki Boulard, 1975. Teixeira and Camargo [in Lima et al. (2013: 13)] described Trypanosoma livingstonei from two bats from Mozambique: Rhinolophus landeri (type host) and H. caffer (additional host) [see also Barbosa et al., 2016: 215 and Cai et al., 2019: Suppl.]. Vercammen-Grandjean and Fain (1958: 10, 12, 14, 20, 22, 30, 33) described Trombigastia (Ascoschongastoïdes) ascoschongastoïdes, Trombigastia (Ascoschongastoïdes) berghei, 224 ISSN 1990-6471 Trombigastia (Scapularia) scapularia, Trombigastia (Trombigastia) laarmani, Trombigastia (Trombigastia) vinckei, Trombigastia (Trombigastia) minor, and Myotrombicula bidentipalpis (Acari: Trombiculidae) from 77 H. caffer centralis specimens collected at Irangi, DRC. Fain (1959c: 239) described Psorergates (Psorergatoides) hipposideros (Acari: Psoridatidae) from a H. caffer centralis captured in Kakontwe cave, Jadotville (=Likasi), DRC. Reviewing African chiggers (Acari: Trombiculidae), Stekolnikov (2018a: 26; 2018b: 268) indicated that Gahrliepia nana (Oudemans, 1910) was described from H. caffer from Durban, where it was also the type host for Austracarus polydiscum (Oudemans, 1910) (Stekolnikov, 2018a: 43). Stekolnikov (2018a: 49, 50, 117, 119, 121, 154, 160, 179) also reported Whartonia atracheata Taufflieb and Mouchet, 1959, Whartonia oweni Vercammen-Grandjean and Brennan, 1957, Trombigastia ascoschoengastoides Vercammen-Grandjean and Fain, 1958, Trombigastia berghei VercammenGrandjean and Fain, 1958, Trombigastia laarmani Vercammen-Grandjean and Fain, 1958, Trombigastia minor Vercammen-Grandjean and Fain, 1958, Trombigastia scapularia VercammenGrandjean and Fain, 1958, Trombigastia vinckei Vercammen-Grandjean and Fain, 1958, Microtrombicula irangiensis VercammenGrandjean, 1965; Microtrombicula minutissima (Oudemans, 1910), and Myotrombicula bidentipalpis Vercammen-Grandjean and Fain, 1958. Periglischrus moucheti Till, 1958 (Acari: Spinturnicidae) was reported by McKechnie et al. (2013: 112). Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100), Brachytarsina alluaudi (Falcoz, 1923) in Botswana (Haeselbarth et al., 1966: 101), Gabon (Obame-Nkoghe et al., 2016: 5), Brachytarsina longiarista (Jobling, 1949) in Sierra Leone (Haeselbarth et al., 1966: 101), Raymondia alulata Speiser, 1908 (Shapiro et al., 2016: 253), Raymondia aspera Maa, 1968 in Mozambique (Shapiro et al., 2016: 254), Raymondia hardyi in South Africa (Shapiro et al., 2016: 254). Raymondia huberi Frauenfeld, 1856 may be associated but remains to be confirmed (Jobling, 1954 [in Haeselbarth et al. (1966: 102)]). Raymondia huberi Frauenfeld, 1856 group in Gabon (Obame-Nkoghe et al., 2016: 5). Raymondia intermedia Jobling, 1936 (Shapiro et al., 2016: 255). Raymondia seminuda Jobling, 1954 from Sierra Leone, the Congo, Tanzania, Zimbabwe, South Africa (Haeselbarth et al., 1966: 103; Shapiro et al., 2016: 255). Raymondia setiloba Jobling, 1954 from Katanga (Haeselbarth et al., 1966: 104; Shapiro et al., 2016: 256 [host as H. caffer-ruber complex]). Raymondia waterstoni Jobling, 1931 (Haeselbarth et al., 1966: 104; Shapiro et al., 2016: 256 [host as H. caffer-ruber complex]). Raymondioides leleupi Jobling, 1954 from Duékoué, Côte d'Ivoire (Haeselbarth et al., 1966: 104). Ascodipteron jonesi Jobling, 1952 from Sudan, the Congo, Uganda and Tanzania (Haeselbarth et al., 1966: 106). Ascodipteron megastigmatos Jobling, 1956 from Duékoué, Côte d'Ivoire (Haeselbarth et al., 1966: 106). Ascodipteron semirasum Maa, 1965 from Botswana, but Maa (1965) doubts the correctness of the host-identification (Haeselbarth et al., 1966: 106). A Brachytarsina allaudi captured on a Gabonese "H. caffer complex" bat was carrying a Polychromophilus melanipherus blood parasite (Szentiványi et al., 2019: Suppl.). Theodor (1968) described Raymondia allisoni Theodor, 1968 from a H. caffer from Ghana, but Shapiro et al. (2016: 253) indicate that the bat species is probably not correct and they assigned it to the H. caffer-ruber complex. Shapiro et al. (2016: 255) also mention Raymondia pagodarum Speiser, 1900 from the species complex in Uganda (but here the original host was referred to as H. ruber), as well as Raymondia simplex Jobling, 1955 (from the DRC). Shapiro et al. (2016: 256) furthermore mention an unnamed Raymondia species (sp. A) from Congo. Szentiványi et al. (2019: Suppl.) indicated that a Gabonese "H. caffer complex" bat was parasitized by a Raymondia huberi group bat fly, which was carrying the blood parasite Polychromophilus melanipherus. Nycteribiidae: Nycteribia schmidlii Schiner, 1853 (Haeselbarth et al., 1966: 108; Duron et al., 2014: 2108; Obame-Nkoghe et al., 2016: 5). Nycteribia scissa sudanica Theodor, 1957 from Ethiopia, the Sudan and the Congo (Haeselbarth et al., 1966: 108). Nycteribia integra Theodor and Moscona, 1954 (Haeselbarth et al., 1966: 109). Nycteribia ovalis Theodor, 1957 from Sierra Leone (Haeselbarth et al., 1966: 110). Nycteribia tecta Theodor, 1957 from Zimbabwe and South Africa, but Haeselbarth et al. (1966: 110) suggests these may represent a distinct subspecies. Penicillidia pachymela Speiser, 1901, from Somalia, Sudan, Kenya, Tanzania, the Congo, Guinea and the Cameroon Mts (Haeselbarth et al., 1966: 114), and Zambia (Blackwell, 1980: 145 - itself infected by the fungus Arthrorhynchus nycteribiae (Peyritsch) Thaxter). Eucampsipoda africana Theodor, 1955 and Penicilidia fulvida Bigot, 1885, found both on bats from the H. caffer complex in Gabon (ObameNkoghe et al., 2016: 5). African Chiroptera Report 2020 Haelewaters et al. (2018: 794) also mention the Nycteribiid Penicillidia pachymela Speiser, 1900, which might be the host for the fungus Arthrorhynchus nycteribiae (Peyr.) Thaxt. VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Hipposideros caffer: Astroviruses, Coronaviruses and Paramyxoviruses. Willoughby et al. (2017: Suppl.) report the following viruses: Chikungunya virus, Human coronavirus 229E, Rift Valley fever phlebovirus. Astroviridae 10 of 57 bats from Mozambique, tested by Hoarau et al. (2018: 2) were positive for Astroviruses. Coronaviridae - Coronaviruses Alphacoronavirus: SARS-CoV (subgenus Duvinacovirus) - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in five individuals from Limpopo Province, South Africa. None were tested positive (0/5), neither did nine individuals from Oriental Province, DRC (0/9). See also note under "viruses" for the genus as possiblitiy. Pfefferle et al. (2009) report the detection of several alpha- and betacoronaviruses from H. cf. caffer/ruber. The alphacoronaviruses were related to the human coronavirus HCoV-229E, and Porcine coronaviruses. Joffrin et al. (2020: 6) reported alphacoronaviruses from bats from Mozambique. 225 Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that one of the 80 H. cf. caffer specimens they examined from Ghana tested positive for Morbillivirus. Of the 337 specimens from Gabon they identified as belonging to H. cf. caffer/ruber, two tested positive for Morbillivirus and one for Rubulavirus. Mortlock et al. (2015: 1841) reported that two out of six examined South African H. caffer specimens tested positive for Paramyxovirus sequences. Markotter et al. (2020: 6) reported the presence of Human orthorubulavirus-related viruses of the genus Orthorubulavirus. In H. caffer bats from Zimbabwbe, Bourgarel et al. (2018: 255) identified viruses belonging to the genus Jeilongvirus. Phenuiviridae Phlebovirus de Jong et al. (2011: 11) and Luis et al. (2013: suppl.) report the occurrence of Rift Valley fever virus (RVFV), either by virus isolation or molecular evidence (see Fagre and Kading, 2019: 5). Togaviridae Alphavirus de Jong et al. (2011: 10) and Luis et al. (2013: suppl.) reported Chikungunya virus occurring on H. caffer. Betacoronavirus: Anthony et al. (2017b: Suppl.) mention the betacoronaviruses Predict_CoV_43 and Predict_CoV_44 (see also Nziza et al., 2019: 157). Both alpha- and betacoranoviruses were reported from Zimbabwe by Bourgarel et al. (2018: 255). Filoviridae Marburgvirus This virus was reported by Nieto-Rabiela et al. (2019: Suppl.). Hepadnaviridae Roundleaf bat HBV – detected in the lung and liver of H. cf. caffer/ruber from Gabon. Nairoviridae Orthonairovirus Only three of the 36 H. cf. caffer specimens from Gabon tested by Müller et al. (2016: 3) were positive for Crimean Congo hemorrhagic fever virus (CCHFV). Figure 57. Distribution of Hipposideros caffer DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Eswatini, Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Malawi, Mali, Mauritania, Mozambique, Namibia, Niger, Nigeria, Rwanda, São Tomé and Principé, 226 ISSN 1990-6471 Senegal, Sierra Leone, Somalia, South Africa, South Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. Hipposideros cf. centralis Andersen, 1906 2016. Hipposideros cf. centralis: Kruskop, Benda, Vasenkov and Lavrenchenko, Lynx, 47: 56. TAXONOMY: Generally, this taxon is considered to be a subspecies of H. ruber, but Kruskop et al. (2016: 57) consider the large-sized NE African individuals from the H. caffer group to represent a separate species, although they indicate that a genetic comparison with specimens from Uganda and NE DRC is still required. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia. Figure 58. Distribution of Hipposideros cf. centralis Hipposideros curtus G.M. Allen, 1921 *1921. Hipposideros curtus G.M. Allen, Rev. Zool. afr., 9 (2): 194. Publication date: December 1921. Type locality: Cameroon: Littoral province: on the Sanaga River: Sakbayeme [04 02 N 10 35 E] [Goto Description]. Holotype: MCZ 19305: ad ♀, alcoholic (skull not removed). Collected by: Rév. George W. Schwab; collection date: 1920. See Helgen and McFadden (2001: 142). - Comments: The cover of issue 9 (2) mentions 15 October 1921 as publication date, but the actual date might be December 1921. (Current Combination) 1937. Hipposideros sandersoni Sanderson, Animal Treasure, 290 (fig.), p. 296. Publication date: September 1937. Type locality: Cameroon: near Mamfe [05 47 N 09 18 E]. Comments: ("bluish-grey and small"). The type locality (Mamfe) was originally situated in Nigeria, but is currently located in Cameroon. TAXONOMY: Simmons (2005: 370) placed it as part of the bicolor species group. Includes sandersoni (Hill, 1963a: 60). The taxonomic relationship between Hipposideros curtus and H. vittatus is unclear and needs further investigation (Mickleburgh et al., 2008dj; IUCN, 2009). COMMON NAMES: Czech: pavrápenec krátkoocasý. English: Shorttailed Leaf-nosed Bat, Short-tailed Roundleaf Bat. French: Phyllorine à queue courte. German: Kurzschwanz-Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B2ab(iii) ver 3.1 (2001)) because its area of occupancy (cave roosting localities) is probably less than 2,000 km ², its distribution is severely fragmented, and there is continuing decline in the extent and quality of its habitat (Mickleburgh et al., 2008dj; IUCN, 2009). Assessment History Global 2008: VU B2ab(iii) ver 3.1 (2001) (Mickleburgh et al., 2008dj; IUCN, 2009). 2004: VU A3c ver 3.1 (2001) (Mickleburgh et al., 2004da; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). African Chiroptera Report 2020 Regional None known. MAJOR THREATS: Threats to this species include ongoing forest loss, and disturbance of cave roosting sites. A number of the few known roosts have already been deserted presumably because of disturbance (Mickleburgh et al., 2008dj; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008dj) [in IUCN (2009)] report that this species has not been recorded from any protected areas. There is an urgent need to protect any important roosting site, and to maintain suitable foraging habitat in the vicinity of these. Further field surveys are needed to locate additional populations of this species, and to determine if the species is present in Nigeria. 227 POPULATION: Structure and Density:- Has been rarely recorded and is known only from a few locations (Mickleburgh et al., 2008dj; IUCN, 2009). Trend:- 2008: Decreasing (Mickleburgh et al., 2008dj; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon. GENERAL DISTRIBUTION: Hipposideros curtus is restricted to southwestern Cameroon and Equatorial Guinea (both Bioko island and Rio Muni). The species might be present in Nigeria, however, this needs confirmation. It is a lowland species found at elevations from sea-level to 500 m. Native: Cameroon (Eisentraut, 1973); Equatorial Guinea (Bioko) (Juste B. and Ibáñez, 1994b). Presence uncertain: Nigeria. Figure 59. Distribution of Hipposideros curtus Hipposideros fuliginosus (Temminck, 1853) *1853. Phyllorrhina fuliginosa Temminck, Esquisses zoologiques sur la Côte de Guiné. 1e partie, les Mammifères, 77. Publication date: 1853. Type locality: Ghana: "Ghana" [Goto Description]. - Comments: Temminck (1853: 78), Allen (1939a: 81) and Grubb et al. (1998: 79) mentioned the type locality as "La côte de Guiné", whereas Jentink (1888b: 167) gave it as 'Cöte d'Or, Pays des Aschantes.'). Temminck (1853: viii) also indicated that M. Pel went to the Ashanti county and visited its capital Coumassie [=Kumasi]. However, on page 88 (discussing Felis celidogaster), Temminck mentions: "Patrie. La côte de Guiné, où on la voit rôder de temps en temps; mais elle est plus abondante dans le pays des Aschantes." This indicates that both are different places. On page 135 (Sciurus leucostigma) he mentions "Patrie. La Guiné, sur les bords de la rivière Boutry et·dans les forêts de la côte jusqu'aux confins du pays des Aschantes." The Butre river is situated in southwestern Ghana, which suggests that the type locality for fuliginosus is on the coast of Ghana (probably somewhere between Butre and Kumasi). 1895. Phyllorhina fuliginosa: Bocage, J. Sci. mat. phys. nat., ser. 2, 4 (8): 4. Publication date: December 1895. (Name Combination) 2014. Hipposideros fuliganosas: Chawana, Alagaili, Patzke, Spocter, Mohammed, Kaswera, Gilissen, Bennett, Ihunwo and Manger, Neuroscience, 277: 725. Publication date: 26 September 2014. (Lapsus) 2016. Hipposideros fulginosus: Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 42. Publication date: 9 May 2016. (Lapsus) ? Hipposideros fuliginosus: (Name Combination, Current Combination) 228 ISSN 1990-6471 TAXONOMY: Simmons (2005: 371) as part of the bicolor species group. Patterson et al. (2020: 121) include it in the ruber subgroup of the bicolor group. Populations from central Africa may well represent a different species from those in western central (west of about 15°) and the Ethiopian records represent another, possibly undescribed, species (Mickleburgh et al., 2008da in IUCN, 2009) based on J. Fahr pers. comm.). Herkt et al. (2017: Appendix S9) indicate that genetic tests evaluating the distinct species status are still lacking, and no clear major geographic gap between both putative species ranges can be discerned. COMMON NAMES: Czech: pavrápenec sazový. English: Sooty Leafnosed Bat, Sooty Roundleaf Bat, Dusky leaf-nosed bat. French: Phyllorine sombre, Phyllorine fuligineuse, Phyllorhine fuligineux. German: Temmincks Rundblattnase, Rußfarbene Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008da; IUCN, 2009; Monadjem et al., 2017bn). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bn). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008da; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004cp; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The major threat to this species is deforestation or degradation of its habitat. Presumably this is largely a result of logging operations and the conversion of land to agricultural use (Mickleburgh et al., 2008da; IUCN, 2009; Monadjem et al., 2017bn). CONSERVATION ACTIONS: Mickleburgh et al. (2008da) [in IUCN (2009)] and Monadjem et al. (2017bn) report that although there are no specific conservation measures in place for this bat, it is presumably present in several protected areas. There is a specific need to maintain large trees as roosting sites for the species. Further studies are needed into the taxonomy, distribution, natural history and possible additional threats to this species. GENERAL DISTRIBUTION: Hipposideros fuliginosus has been recorded in West Africa and Central Africa. It ranges from Guinea and Sierra Leone in the west, and has been recorded from Liberia, Côte d'Ivoire, Ghana and Nigeria. In Central Africa it is distributed in southern Cameroon, with scattered records from Gabon, Central African Republic, northern parts of Democratic Republic of the Congo and western Uganda. It is generally a lowland species, being recorded up to 500 m asl. Native: Cameroon (Eisentraut, 1973); Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 538); Côte d'Ivoire; Gabon; Ghana (Decher and Fahr, 2007: 16); Guinea (Fahr and Ebigbo, 2003: 128; Decher et al., 2016: 265); Guinea-Bissau (Rainho and Ranco, 2001: 58); Liberia (Koopman, 1989b; Koopman et al., 1995); Nigeria; Sierra Leone; Uganda. Presence uncertain: Central African Republic; Congo; Equatorial Guinea. DETAILED MORPHOLOGY: Brain: Adult hippocampal neurogenesis was studied by Chawana R. et al. (2016: 1551). ECHOLOCATION: In Guinea Fahr and Ebigbo (2003) reported the frequency of the CF-component at 122.5 ± 1.3 (120.0 - 123.3) kHz. POPULATION: Structure and Density:- There is little information on its overall abundance. It is usually found in small groups, but sometimes in congregations of up two hundred bats (Mickleburgh et al., 2008da; IUCN, 2009; Monadjem et al., 2017bn). Trend:- 2016: Decreasing (Monadjem et al., 2017bn). 2008: Decreasing (Mickleburgh et al., 2008da; IUCN, 2009). PARASITES: DIPTERA: Streblidae: Raymondia huberi huberi Frauenfeld, 1855 (Shapiro et al., 2016: 254). Raymondia seminuda Jobling, 1954 in southern Nigeria (Haeselbarth et al., 1966: 103). VIRUSES: Paramyxoviridae Mortlock et al. (2015: 1841) reported that three out of 21 examined H. fuliginosus specimens from the DRC tested positive for Paramyxovirus sequences. African Chiroptera Report 2020 229 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Gabon, Ghana, Guinea, Liberia, São Tomé and Principé, Senegal, Sierra Leone, Tanzania. Figure 60. Distribution of Hipposideros fuliginosus Hipposideros jonesi Hayman, 1947 *1947. Hipposideros jonesi Hayman, Ann. Mag. nat. Hist., ser. 11, 14 (109): 71, fig. 1. Publication date: 19 November 1947. Type locality: Sierra Leone: Makeni [08 57 N 12 02 W] [Goto Description]. Holotype: BMNH 1947.629: ad ♀, skull and alcoholic. Collected by: Theo Simpson Jones; collection date: 12 December 1946; original number: 42. Etymology: In honour of Mr. T.S. Jones, the collector of the type specimen. (Current Combination) TAXONOMY: Simmons (2005: 373) as part of the bicolor species group. Patterson et al. (2020: 121) place it in the speoris group. COMMON NAMES: Czech: pavrápenec Jonesův. English: Jones' Leaf-nosed Bat, Jones' Roundleaf Bat, Jones's Roundleaf Bat, Western Africa leaf-nosed bat. French: Phyllorine de Jones, Phyllorine d'Afrique occidentale. German: Jones' Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Near Threatened (NT ver 3.1 (2001)) since the species depends on caves for roost sites, and the area of occupancy might not be much greater than 2,000 km2. The quality of its limited roosting habitat is likely to be declining through disturbance of caves, thus making the species close to qualifying for Vulnerable under criterion B2 (Mickleburgh et al., 2008au; IUCN, 2009). Assessment History Global 2008: NT ver 3.1 (2001) Mickleburgh et al., 2008au; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004br; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is generally threatened by the disturbance of roosting caves and might additionally be threatened by subsistence hunting although this needs to be confirmed (Mickleburgh et al., 2008au; IUCN, 2009). Nkrumah et al. (2018: 755) indicate that H. jonesi is a generalist in the Kwamang agro-ecosystem of Ghana, and as such, might not be sensitive to increased disturbance. However, they also mention that their results need to be cautiously interpreted. CONSERVATION ACTIONS: Mickleburgh et al. (2008au) [in IUCN (2009)] report that there appear to be no direct conservation measures in place for this species. It is not known if the species is present within any protected areas. There is a need to conserve important roosts for this species. Additional studies are needed into the distribution, natural history and threats to this species. GENERAL DISTRIBUTION: Hipposideros jonesi is a West African endemic, closely associated with caves. It has been recorded from Sierra Leone and Guinea to Mali, 230 ISSN 1990-6471 Burkina Faso, Ghana and Nigeria. between sea level and 1,400 m asl. It is found Native: Burkina Faso (Kangoyé et al., 2015a: 608); Côte d'Ivoire; Ghana; Guinea (Decher et al., 2016: 265); Liberia (Fahr, 2007a: 104); Mali; Nigeria (Happold, 1987); Sierra Leone (Hayman, 1947d; Grubb et al., 1998). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Young (1975: 76) indicates that two distinct colour phases can be found: rufous (orange) and grey. MOLECULAR BIOLOGY: DNA - Unknown Karyotype - Koubínová et al. (2010b) report for on a single male from Senegal, 2n= 32, Fna= 60 and FN= 64, these included 30 biarmed autosomes, including four metacentric, eight submetacentric and three subtelocentric, eight submetacentric and three subtelocentric pairs. A secondary constriction situated near the centromere was observed in one pair of the medium-sized submetacentric autosomes (approximately the 9th largest pair). The X chromosome was identified as a large metacentric element, the Y chromosome as a medium submetacentric. Protein / allozyme - Unknown. DIET: In the Kwamang agro-ecosystem in Ghana, E.K. Badu (in Nkrumah et al., 2018: 755) found that the diet of H. jonesi consists of nearly 100 % Lepidoptera. POPULATION: Structure and Density:- This species forms roosts of only a few animals, usually in the dozens rather than hundreds. There is a pronounced geographic morphological variation between populations, suggesting that there may be little movement of animals between populations (Mickleburgh et al., 2008au; IUCN, 2009). Trend:- 2008: Decreasing (Mickleburgh et al., 2008au; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Burkina Faso, Cameroon, Côte d'Ivoire, Ghana, Guinea, Liberia, Mali, Nigeria, Senegal, Sierra Leone. HABITS: Lawer and Darkoh (2016: 429) found that this bat's activity declined with increasing lunar illumination, indicating that bright moonlight suppresses the nocturnal activities of both insectivorous bats and insects. They also found a negative association between wind speed and overal insectivorour bat activity (p. 430). ROOST: Weber and Fahr (2006: 4) indicate that in Guinea H. jonesi largely or exclusively depends on the availability of caves as day roosts. Figure 61. Distribution of Hipposideros jonesi Hipposideros lamottei Brosset, 1985 *1985. Hipposideros lamottei Brosset, Mammalia, 48 (4): 548 (for 1984). Publication date: 1985. Type locality: Guinea: Mount Nimba: Pierré Richaud [07 41 N 08 22 W] [Goto Description]. Holotype: MNHN ZM-MO-1984-487: ad ♂. Collected by: André Brosset; collection date: December 1983. Paratype: MHNG ad ♂. Collected by: André Brosset; collection date: December 1983. Paratype: MNHN ZM-MO-1984-488: ad ♂. Collected by: André Brosset; collection date: December 1983. Paratype: MNHN ZM-MO-1984-489: ad ♂. Collected by: André Brosset; collection date: December 1983. (Current Combination) 2020. Hipposideros cf. lamottei: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicate that this is the same form as the Hipposideros ruber B1 lineage mentioned by Monadjem et al. (2013b). (Name Combination) African Chiroptera Report 2020 TAXONOMY: Simmons (2005: 373) bicolor species group. Patterson et al. (2020: 121) place it in the ruber subgroup of the bicolor group. Distinction from H. ruber is not entirely clear (Koopman, 1993a: 173; Koopman et al., 1995; Simmons, 2005: 373). Monadjem et al. (2013b: 341) state that, morphologically, H. lamottei falls within the caffer/ruber group. COMMON NAMES: Czech: pavrápenec nimbánský. English: Lamotte's Leaf-nosed Bat, Lamotte's Roundleaf Bat, Mount Nimba Leaf-nosed Bat. French: Phyllorine de Lamotte, Phyllorine du Mont Nimba. German: Mount Nimba-Rundblattnase, Lamottes Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Critically Endangered (CE B1ab(I,ii,iii,iv,v)+2ab(I,ii,iii,iv,v) ver 3.1 (2001)) because its extent of occurrence is probably less than 100 km2 and its area of occupancy (cave roosting sites) is likely to be less than 10 km 2, all individuals are in a single location (Mount Nimba), and there is continuing decline in the extent and quality of its habitat (Mickleburgh et al., 2008av; IUCN, 2009). Kouame et al. (2012: 2748) indicate that H. lamottei is a highly threatened species, which only occurs in one single KBA [= Key Biodiversity Area] in the Upper Guinea Forest: Mount Nimba. Collen et al. (2011: 2616) rank it on position 62 (as only African mainland Chiropteran) out of 100 mammal species requiring urgent conservation attention. Assessment History Global 2008: CE B1ab(I,ii,iii,iv,v)+2ab(I,ii,iii,iv,v) ver 3.1 (2001) (Mickleburgh et al., 2008av; IUCN, 2009). 2004: CR B1ab(iii) ver 3.1 (2001) (Mickleburgh et al., 2004bs; IUCN, 2004). 1996: DD (Baillie and Groombridge, 1996). CONSERVATION ACTIONS: Mickleburgh et al. (2008av) [in IUCN (2009)] report that it is present within the Mount Nimba Strict Nature Reserve World Heritage Site. There is a need to enforce the protection of this area. Additional surveys are needed to learn more about the distribution, natural history and threats to this species. GENERAL DISTRIBUTION: Known only from Mt. Nimba on Guinea-Liberia border, but probably more widespread. However, Fahr (2007a: 104) and Decher and Fahr (2007: 5) indicate that specimens from Liberia, Ghana, Sierra Leone and Cameroon (mentioned by Koopman et al., 1995 and Grubb et al., 1998) were re-identified as belonging to H. caffer. They restrict the distribution of lamottei to the Guinean side of Mt. Nimba. It has been recorded between 500 and 1,400 m asl. Native: Guinea (Brosset, 1985; Denys et al., 2013: 284). ECHOLOCATION: Monadjem et al. (2013b: 351) report a Fmax of 119.0 (118 - 120) kHz for two specimens from Mount Nimba. POPULATION: Structure and Density:- Only six specimens are currently known from two localities (Mickleburgh et al., 2008av; IUCN, 2009). Trend:- 2008: Decreasing (Mickleburgh et al., 2008av; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Guinea, Liberia, Senegal. Regional None known. MAJOR THREATS: The species is believed to be highly threatened because of extensive iron ore mining activities underway, and planned, within its limited range on Mount Nimba. It is probably additionally threatened by general deforestation in parts of its range. Indiscriminate subsistence hunting of bats for food occurs in caves on Mount Nimba and likely impacts this species (Mickleburgh et al., 2008av; IUCN, 2009). 231 Figure 62. Distribution of Hipposideros lamottei 232 ISSN 1990-6471 Hipposideros marisae Aellen, 1954 *1954. Hipposideros marisae Aellen, Rev. suisse Zool., 61 (24): 474. Type locality: Côte d'Ivoire: Duékoué: White Leopard Rock [Goto Description]. Holotype: [Unknown] ad ♂. Collection date: 13 May 1953; original number: 400. See Aellen (1954: 474). Comments: Type locality original: roche de la Panthère Blanche. - Etymology: In honour of Mrs. Aellen (Aellen, 1954: 474). (Current Combination) TAXONOMY: Simmons (2005: 374) as part of the bicolor species group. Monadjem et al. (2013b: 346) indicate that H. marisae is a sistergroup to H. jonesi, which Patterson et al. (2020: 121) confirm by placing both in the speoris group. COMMON NAMES: Czech: pavrápenec Aellenův. English: Marisa's Leaf-nosed Bat, Aellen's Leaf-nosed Bat, Aellen's Roundleaf Bat. French: Phyllorine d'Aellen. German: Marisas Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B2b(ii,iii,iv,v) ver 3.1 (2001)) because its area of occupancy is probably less than 2,000 km 2, its distribution is possibly severely fragmented (roosting sites), there is continuing decline in the quality of its cave habitats (through disturbance), and there is likely to be a corresponding decline in subpopulations and individuals (with over hunting likely at some colonies) (Mickleburgh et al., 2008aw; IUCN, 2009). Assessment History Global 2008: VU B2b(ii,iii,iv,v) ver 3.1 (2001) (Mickleburgh et al., 2008aw; IUCN, 2009). 2004: EN A4c ver 3.1 (2001) (Mickleburgh et al., 2004bt; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is threatened by general deforestation, mining operations on Mount Nimba and the disturbance of roosting caves. It is likely harvested for subsistence use as food (Mickleburgh et al., 2008aw; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008aw) [in IUCN (2009)] report that it is present within the Mount Nimba Strict Nature Reserve World Heritage Site. There is a need to protect important roost sites and areas of suitable forest habitat. Further studies are needed into the distribution of this species. GENERAL DISTRIBUTION: This West African species has been recorded from seven localities in Guinea, Liberia (central and western uplands), and Côte d'Ivoire, and two localities in Sierra Leone. It may range into Ghana, although there is no record from this country (Grubb et al., 1998). It is a lowland species recorded between sea level and 650 m asl. Native: Côte d'Ivoire (Aellen, 1954); Guinea; Liberia (Kuhn, 1965; Hill, 1982a; Wolton et al., 1982; Koopman et al., 1995; Monadjem and Fahr, 2007: 53); Sierra Leone (Weber et al., 2019: 22 first record). Presence uncertain: Ghana. ECHOLOCATION: Monadjem et al. (2013b: 351) report Fmax to be 146.0 kHz at Mount Nimba. In Sierra Leone, Weber et al. (2019: 21) recorded two females calling at 142.5 (141.0 - 144.0) kHz and two males at 146.5 (146.0 - 147.0) kHz. HABITAT: Weber et al. (2019: 22) indicate that this is an extremely rare species, which might depend on suitable caves as day roost and forest habitats. POPULATION: Structure and Density:- It is generally considered to be a very rare species (Mickleburgh et al., 2008aw; IUCN, 2009). Monadjem et al. (2013b: 342) state that this species has not been captured nor seen since 1990. Trend:- 2008: Decreasing (Mickleburgh et al., 2008aw; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Côte d'Ivoire, Guinea, Liberia, Sierra Leone. African Chiroptera Report 2020 233 Figure 63. Distribution of Hipposideros marisae Hipposideros megalotis (Heuglin, 1861) *1861. Phyllorrhina megalotis Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 8. Publication date: 1861. Type locality: Eritrea: Bogos country [=Keren region] [ca. 15 45 N 38 20 E] [Goto Description]. Holotype: SMNS 984: ad ♂, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. See: Dieterlen et al. (2013: 294). - Comments: Type specimen: SMNS, ???. Horácek et al. (2000: 98) mention "Monatsber. Königl. Preuss. Akad. Wiss. Berlin: 329". Nancy Simmons (pers. comm.) checked this and rejected this reference. Etymology: From the Greek adjective "μέγα" (méga), meaning "big" and the neuter Greek substantive "οὖς" (genitive "ώτός", "ûs", "ōtós"), meaning "ear" referring to the large ears of the species (see Lanza et al., 2015: 145). ? Hipposideros megalotis: (Name Combination, Current Combination) TAXONOMY: Placed in the subgenus Syndesmotis by Legendre (1982), Gaucher and Brosset (1990) and Horácek et al. (2000: 98) include it with a question mark). Horácek et al. (2000: 98) as part of the megalotis species group, followed by Simmons (2005: 374). Patterson et al. (2020: 121) placed it as only species in the megalotis group. COMMON NAMES: Chinese: 串耳蹄蝠. Czech: pavrápenec ušatý. English: Large-eared Leaf-nosed Bat, Ethiopian Large-eared Leaf-nosed Bat, Ethiopian Largeeared Roundleaf Bat, Big-eared leaf-nosed bat. French: Phyllorine à grandes oreilles d'Ethiopie, Phyllorine à grandes oreilles. German: GroßohrRundblattnase. Italian: Ipposìdero orecchiùto. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) because, although it is seldom recorded, it has a relatively wide distribution, is tolerant of a range of habitats, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ax; IUCN, 2009; Monadjem et al., 2017as). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017as). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008ax; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bu; IUCN, 2004). 2000: LR/nt (Hilton-Taylor, 2000). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species remain poorly known. Its is suspected to be threatened by habitat loss resulting from the conversion of land to agricultural use, and by disturbance of roosting sites (Mickleburgh et al., 2008ax; IUCN, 2009; Monadjem et al., 2017as). CONSERVATION ACTIONS: Mickleburgh et al. (2008ax) [in IUCN (2009)] and Monadjem et al. (2017as) report that there are no direct conservation measures in place for this 234 ISSN 1990-6471 species. It is not known if the species is present in any protected areas. Additional studies are needed into the distribution, abundance, general ecology and threats to this little-known species. GENERAL DISTRIBUTION: Hipposideros megalotis has been reported from Eritrea (including the type locality of Keren (Heuglin, 1861)), Ethiopia (including Sidamo in southern Ethiopia (Hayman and Hill, 1971)), Kenya (where it has been recorded from the localities of Kinangop and Nakuru (Hayman, 1954)), Somalia, Djibouti (Heuglin, 1861; Pearch et al., 2001) and from Jeddah in Saudi Arabia (Gaucher and Brosset, 1990). It has been reported at elevations of up to 2,000 m asl. Trend:- 2016: Unknown (Monadjem et al., 2017as). 2008: Unknown (Mickleburgh et al., 2008ax; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Eritrea, Ethiopia, Kenya. Native: Djibouti (Heuglin, 1861; Pearch et al., 2001); Eritrea (Heuglin, 1861); Ethiopia (Hayman and Hill, 1971); Kenya (Hayman, 1954); Saudi Arabia (Gaucher and Brosset, 1990). Presence uncertain: Somalia (Senna, 1905). POPULATION: Structure and Density:- Little is known about the population of this species. Very few animals have ever been recorded (Mickleburgh et al., 2008ax; IUCN, 2009; Monadjem et al., 2017as). Figure 64. Distribution of Hipposideros megalotis Hipposideros ruber (Noack, 1893) *1893. Phyllorhina rubra Noack, Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere, 7: 586, pl. 18, fig. 14, 15. Publication date: 23 December 1893. Type locality: Tanzania: Eastern province: Ngerengere River [ca 07 00 S 38 00 E] [Goto Description]. Holotype: ZMB 89571: ♂, skin only. Collected by: Dr. Emin Pasha; collection date: 20 May 1890. Formerly Noack collection number 5, see Turni and Kock (2008: 38). Holotype: ZMB no number: skin only. Collected by: Dr. Emin Pasha; collection date: 20 May 1890. Formerly Noack collection number 5; see Turni and Kock (2008: 38). - See ZMB 89571. - Comments: Noack (1893: 586) and Allen (1939a: 81) mentioned the type locality as "Lugerrunjere Fluss, Tanganyika Territory". Grubb et al. (1998: 80) mention the type locality as "Lugerrunjere Fluss = Ngerengere River, Tanzania - probably Ngerengere village, 32 miles east of Morogor according to Swynnerton, 1945, Proc. Zool. Soc. Lond., 115: 69". - Etymology: From the Latin "ruber" meaning red. 1906. Hipposiderus caffer centralis K. Andersen, Ann. Mag. nat. Hist., ser. 7, 17 (99): 275, 277. Publication date: 1 March 1906. Type locality: Uganda: Entebbe [Ntebbi] [00 04 N 32 28 E, 3 800 ft] [Goto Description]. Holotype: BMNH 1899.8.4.8: ad ♂, skin only. Collection date: 5 April 1893. Presented/Donated by: Sir Frederic John Jackson. 1906. Hipposiderus caffer guineensis K. Andersen, Ann. Mag. nat. Hist., ser. 7, 17 (99): 275, 278. Publication date: 1 March 1906. Type locality: Gabon: 70 mi (105 km) from Gaboon: Como River [ca. sea level] [Goto Description]. Holotype: BMNH 1897.12.1.11: ad ♀, skin only. Collected by: George Latimer Bates Esq. Collection date: 3 June 1897; original number: GLB 215. Note: caught under large rock, Fang name- Otan. Comments: Considered a valid subspecies by Grubb et al. (1998: 80). 1917. Hipposideros caffer niapu J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 431. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): NE Congo: Niapu [02 25 N 26 28 E] [Goto Description]. Holotype: AMNH 49414: ad ♂, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 27 January 1914; original number: 2322. Topotype: MCZ 17224: ♂, alcoholic (skull not removed). Collected by: Herbert Lang, James Paul African Chiroptera Report 2020 2016. 2016. 2016. 2018. 2018. 2019. 2020. 2020. 2020. 2020. 2020. ? ? ? ? ? ? ? 235 Chapin and The American Museum Congo Expedition; collection date: 27 February 1914. Presented/Donated by: ?: Collector Unknown. Hipposideros aff. ruber: Nkrumah, Vallo, Klose, Ripperger, Badu, Drosten, Kalko, Tschapka and Oppong, Acta Chiropt., 18 (1): 239. Publication date: June 2016. Comments: Nkrumah et al. (2016b: 3) indicate that this is the same form as "H. ruber lineage D from Vallo et al. (2009: 196), and which is evolutionarily distant to Hipposideros ruber s. str. from East Africa. (Name Combination) Hipposideros cf. ruber (lineage C1) Monadjem, Richards and Denys, Acta Chiropt., 18 (2): 365. Publication date: December 2016. - Comments: . Hipposideros cf. ruber (lineage E1) Monadjem, Richards and Denys, Acta Chiropt., 18 (2): 366. Publication date: December 2016. - Comments: . H[ipposideros] rubber: Malekani, Musaba, Gembu, Bugentho, Toengaho, Badjedjea, Ngabu, Mutombo, Laudisoit, Ewango, Van Cakenberghe, Verheyen, Asimonyo, Masudi, Bongo and Ngbolua, Nat. Conserv. Res., 3 (1): 67.. Publication date: January 2018. (Lapsus) Hip[posideros] caffer rufer: Bourgarel, Pfukenyi, Boué, Talignani, Chiweshe, Diop, Caron, Matope, Missé and Liégeois, Infec. Gen. Evol., 58: 255. Publication date: 10 January 2018. (Lapsus) Hipposideros caffer rubber: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) Hipposideros ruber 1: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicates that this is partially the same form as the Hipposideros ruber C1 lineage mentioned by Vallo et al. (2009) and the Hipposideros cf. ruber C1a and C1b lineages mentioned by Monadjem et al. (2013b). (Name Combination) Hipposideros ruber 2: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicates that this is partially the same form as the Hipposideros ruber C1 lineage mentioned by Vallo et al. (2009). (Name Combination) Hipposideros ruber 3: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicates that this is the same form as the Hipposideros ruber C2 lineage mentioned by Vallo et al. (2009) and the Hipposideros cf. ruber C2 lineage mentioned by Monadjem et al. (2013b). (Name Combination) Hipposideros ruber 4: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. (Name Combination) Hippposideros cf. ruber: Patterson, Webala, Lavery, Agwanda, Goodman, Kerbis Peterhans and Demos, ZooKeys, 929: 126. Publication date: 22 April 2020. - Comments: Patterson et al. (2020: 138) indicates that this is the same form as the Hipposideros ruber D lineage mentioned by Vallo et al. (2009) and the Hipposideros cf. ruber D1, D2, E1 and E2 lineages mentioned by Monadjem et al. (2013b). (Name Combination) Hipposideros caffer guineensis: Hipposideros caffer ruber: (Name Combination) Hipposideros ruber centralis: (Name Combination) Hipposideros ruber guineensis: (Name Combination) Hipposideros ruber niapu: (Name Combination) Hipposideros ruber ruber: (Name Combination) Hipposideros ruber: (Name Combination, Current Combination) TAXONOMY: Simmons (2005: 376) recognised it as part of the bicolor species group, and indicated that the subspecies limits are problematic, and it possible that this species complex includes more than one species. Vallo et al. (2007: 257; 2009: 193) indicate that two sympatric lineages exist in ruber based on analysis of complete sequences of the mitochondrial gene for cytochrome b, with possibly a third form in West Africa. Mitochondrial DNA analyses on specimens from the Niokolo Koba National Park in Senegal indicated that two distinct lineages of H. aff. ruber exists in that area (Vallo et al., 2011b: 84). Patterson et al. (2020: 121) place it in the ruber subgroup of the bicolor group. Decher et al. (2016: 265) indicate that they provisionally include all Simandou records 236 ISSN 1990-6471 (Guinea), previously named H. caffer or H. ruber in H. cf. ruber. Based on 11 cytochrome b microsatellite loci, Baldwin et al. (2014: 1) called the West African bats "Hipposideros aff. ruber" as these differed from East African H. ruber s. str., indicating it might represent a separate species. Nkrumah et al. (2016a: 240) also used the name "H. aff. ruber for their Ghanaian specimen (assigned to the D gentotype lineage), which is rather distant from H. ruber s.str. from East Africa. Nkrumah et al. (2017: 348) assigned their Ghanaian bats to H. cf. ruber as these include multiple genetic lineages. Examining cyt b sequences from various parts of Africa led Mande (2019: 5) to distinguish four clades: (a) restricted to the DRC, (b) spread from Liberia, through Cameroon, the DRC to Uganda, (c) occurring in Cameroon, Equatorial Guinea and São Tomé and Principé, and (d) restricted to Senegal. COMMON NAMES: Chinese: 诺氏蹄蝠. English: Noack's Leaf-nosed Bat, Noack's Roundleaf Bat, Red Leaf-nosed Bat. French: Phyllorine rouge, Phyllorine de Noack. German: Noacks Rundblattnase, KamerunBlattnase. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ay; IUCN, 2009; Monadjem et al., 2017at). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017at). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008ay; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bv; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is locally threatened by the loss and conversion of suitable habitat. It is also threatened in parts of its range by collection for subsistence use (Mickleburgh et al., 2008ay; IUCN, 2009; Monadjem et al., 2017at). CONSERVATION ACTIONS: Mickleburgh et al. (2008ay) [in IUCN (2009)] and Monadjem et al. (2017at) report that in view of its wide range it seems likely that the species is present in a number of protected areas. It has been recorded from the Manga Forest Reserve of Tanzania (Doggart et al., 1999b). No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Hipposideros ruber is a widespread species recorded throughout much of West, Central, and East Africa and part of southern Africa including Angola, southern Democratic Republic of the Congo, northern and eastern Zambia, southern Malawi and northwestern Mozambique. It is found between sea level and 2,300 m asl. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 625,018 km2. Native: Angola (Hayman, 1963; Crawford-Cabral, 1989; Monadjem et al., 2010d: 538); Benin; Burkina Faso (Kangoyé et al., 2015a 608); Burundi (Hayman et al., 1966); Cameroon; Central African Republic (Lunde et al., 2001: 537); Chad; Congo (Bates et al., 2013: 318 - 1st authenticated records); Congo (The Democratic Republic of the) (Allen, 1917; Römer, 1862; Hayman et al., 1966; Monadjem et al., 2010d: 538) ; Côte d'Ivoire; Equatorial Guinea; Ethiopia; Gabon; Gambia (Emms and Barnett, 2005: 50); Ghana (Decher and Fahr, 2007: 13 - 15); Guinea (Fahr and Ebigbo, 2003: 128; Denys et al., 2013: 283; Decher et al., 2016: 265 mention it as "Hipposideros cf. ruber"; Weber and Fahr, 2007); Guinea-Bissau; Kenya; Liberia (Monadjem and Fahr, 2007: 50); Malawi (Happold et al., 1988; Monadjem et al., 2010d: 538); Mali; Mozambique (Monadjem et al., 2010d: 538; Monadjem et al., 2010c: 381); Niger; Nigeria (Happold, 1987; Rwanda (Hayman et al., 1966); Saõ Tomé (Rainho et al., 2010a: 28); Senegal; Sierra Leone; Sudan; Tanzania , United Republic of (Cunneyworth, 1996b; Doggart et al., 1999b); Togo; Uganda (Kityo and Kerbis, 1996: 61); Zambia (Monadjem et al., 2010d: 538). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Lucati and López-Baucells (2016: Suppl.) report on five albino and nine piebald (all-white fur/skin patches, eyes always normally coloured) specimens from mines in Tanzania (Howell and Mainoya, 1974: 155; Howell, 1980). ECHOLOCATION: From Irangi (Kivu province, DRC), Heller (1992: 75) reported that H. ruber emitted calls with frequencies between 130 and 140 kHz, distinguishing them from H. caffer, which emitted calls at a higher frequency. African Chiroptera Report 2020 Fahr and Ebigbo (2003) reported for an individual of H cf. ruber from the Simandou Range in southeastern Guinea a CF-component of 146.7 kHz. At Sengwa in Zimbabwe, Jacobs (1996) and Fenton and Bell (1981) reported a highest frequency of 138 kHz. Monadjem et al. (2010c: 378) reported from Mozambique that the peak echolocation frequencies ranged between 132 136 kHz (ANABAT) and 130.5 kHz (Petterson D240x). Rainho et al. (2010a: 21) report the call parameters from Saõ Tomé Island. Monadjem et al. (2013b: 351) report on two lineages of H. cf. ruber from Mount Nimba: C1 with a Fmax of 147.7 ± 1.45 (145 - 150) Khz, and E1 with Fmax of 127.5 (127 - 128) kHz. Jones and Siemers (2010: 449 - 450) indicated that males emit pulses with a higher frequency than females, and that the CF-frequency is positively related to the body condition (mass/forearm length). Trentin and Rovero (2011: 56) describe the call from bats at Uzungwa Scarp Forest Reserve, Tanzania, as having a first modulated component starting at 143.9 kHz, which increases to a maximum frequency of 148.6 kHz, which continues for about 7 msecs, and then drops to 121.4 kHz in about 1 msec. The maximum intensity is at 144.8 kHz. For three Kenyan males and two females, Webala et al. (2019b: 15) reported Fmax between 132.6 and 134.5, and 140.4 and 145.3 kHz, whereas the duration ranged from 7.4 to 8.8 and 7.3 to 7.7 msec respectively. In Sierra Leone, Weber et al. (2019: 21) recorded calls from two types of H. ruber: C1 (n=3): 145.7 ± 4.5 (142.0 - 152.0) kHz and D 4 FF: 122.5 ± 2.4 (120.0 - 125.0) kHz and 2 MM: 127.0 kHz. Luo et al. (2019a: Supp.) reported the following data (Free flying bats): F peak: 131.96 kHz and duration: 8.06 msec. MOLECULAR BIOLOGY: DNA - Lim (2007), Vallo et al. (2007). Karyotype - Koubínová et al. (2010b) report on seven females from Senegal, 2n= 32, Fna= 60 and FN= 64, they comprised 30 biarmed autosomes, including four metacentric, eight submetacentric and three subtelocentric, eight submetacentric and three subtelocentric pairs, a secondary constriction situated near the centromere was observed in one pair of the medium-sized submetacentric autosomes (approximately the 9th largest pair). The X chromosome was identified 237 as a large metacentric element, the Y chromosome as a medium submetacentric. Denys et al. (2013: 284) report the following karyotype for a male H. cf. ruber from the Guinean Mount Nimba area: 2n = 32, FNa =60, including 15 pairs of meta- and submetacentric chromosomes. The X chromosome is middle-sized submetacentric and the Y chromosome is small subtelocentric. Protein / allozyme - Unknown. HABITAT: In Ghana, Nkrumah et al. (2016b: 5) reported H. aff. ruber to prefer foraging in seminatural habitats, followed by mixed farm areas, and, to a much lesser extent, tree farms and savanna grasslands. HABITS: Russo et al. (2010b: 701) report H. ruber to be diurnally active on the island of São Tomé, and indicate that adult males were much more active during the daytime than females, whereas during the night the opposite was true, possibly leading to a temporal niche partitioning. Russo et al. (2010a: 271) indicate that about one fifth of a colony may be out during daytime, also due to the absence of forest-dwelling avian predators. In Ghana, Nkrumah et al. (2016a: 239) found that during the night, males of H. aff. ruber returned more often to their cave roosts than females. Based on that data for 13 bats, they calculated the bat's homerange as 36 ± 35 (6 - 95) ha. The foraging range covered 50 % of this and the core area 2 %. Nkrumah et al. (2016b: 6) found that females had a night activity pattern that peaks between 6 and 8 p.m., with a smaller peak around midnight and another peak between 5 and 6 a.m. In between the flight duration dropped to almost zero. In males, the activity peaked around 8 p.m., midnight and between 4 and 5 a.m. However, their activity in the first part of the night did only decrease by a maximum of 10 minutes. Nkrumah et al. (2017: 350) found that activity patters of the bats differed from cave to cave. ROOST: Weber and Fahr (2006: 4) indicate that H. ruber largely or exclusively depends on the availability of caves as day roosts in Guinea. In Ghana, Nkrumah et al. (2016a: 239) found that caves were used as main roost, but that H. aff. ruber also used trees as individual night roosts. In the Mframabuom cave, the roost contained some 3,500 individuals (together with H. abae, H. jonesi and Nycteris cf. thebaica). 238 ISSN 1990-6471 DIET: Bell and Fenton (1984) [in Fenton, 1986a] found that H. ruber depends on flutter to find its prey and varies its hunting pattern by taking both flying prey and gleaning targets from surfaces such as foliage and water. They also found no restriction to a specific prey species or size class. Nkrumah et al. (2017: 353) reports that H. cf. ruber feeds primarlily on Lepidoptera in Ghana. POPULATION: Structure and Density:- This is one of the most common bats in Africa, with some colonies containing up to 500,000 individuals (Mickleburgh et al., 2008ay; IUCN, 2009; Monadjem et al., 2017at). Trend:- 2016: Unknown (Monadjem et al., 2017at). 2008: Unknown (Mickleburgh et al., 2008ay; IUCN, 2009). REPRODUCTION AND ONTOGENY: Trentin and Rovero (2011: 50) captured nine lactating and two pregnant females in December 2005 at Uzungwa Scarp Forest Reserve, Tanzania. In Kwamang (Ghana), the bats gave birth from late March to early May (Nkrumah et al., 2017: 348). In Sierra Leone, Weber et al. (2019: 23) reported two pregnant females (type "C1") on 1 and 7 April, and one pregnant (24 March) and one lactating (2 April) female (type "D"). PARASITES: Peralta-Rodríguez et al. (2012: 1006) report on the infection of H. ruber by Spirura hipposiderosi Khalil, 1975 (Nematode) in Tanzania. In his review of African chiggers (Acari: Trombiculidae), Stekolnikov (2018a: 49, 50, 72, 117, 119, 121, 123, 173, 179, 184) reported the presence of Whartonia lepidopteriscuta Vercammen-Grandjean, 1965, Whartonia oweni Vercammen-Grandjean and Brennan, 1957, Riedlinia willmanni Vercammen-Grandjean and Minter, 1964, Trombigastia ascoschoengastoides Vercammen-Grandjean and Fain, 1958, Trombigastia berghei Vercammen-Grandjean and Fain, 1958, Trombigastia laarmani VercammenGrandjean and Fain, 1958 and Trombigastia minor Vercammen-Grandjean and Fain, 1958, Trombigastia scapularia Vercammen-Grandjean and Fain, 1958, Trombigastia vinckei VercammenGrandjean and Fain, 1958, Afrotrombicula machadoi (Taufflieb, 1962), Microtrombicula tanzaniae Goff, 1982, Myotrombicula bidentipalpis Vercammen-Grandjean and Fain, 1958, and Oudemansidium howelli (Goff, 1983). Decher et al. (2016: 266) found the bat fly Penicillida allison Theodor, 1968 (Nycteribiidae) on a Guinean "H. cf. ruber". VIRUSES: Coronaviridae - Coronaviruses Alphacoronavirus: SARS-CoV (subgenus Duvinacovirus) - See also note under "viruses" for the genus as possiblitiy. Corman et al. (2015: 11859) tested 1,611 Ghanese H. cf. ruber specimens, of which 62 (3.8 %) tested positive for 229E-related coronavirsues. Anthony et al. (2017b: Suppl.) mention the betacoronavirus Predict_CoV_20. Willoughby et al. (2017: Suppl.) report Human coronavirus 229E. Nziza et al. (2019: 157) found a new coronavirus (PREDICT_CoV-43) in a rectal swap from a Rwandan bat. Maganga et al. (2020: 1) reported viruses belonging to the genus from Gabonese 13 out of 262 (4.96 %) H. cf. ruber bats, and found them to be significantly present in October and November. Flaviviridae In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against non-specific Flaviviruses. Hantaviridae (formerly included in Bunyaviridae) Hantavirus Witkowski et al. (2016: 113) report a new hantavirus from Gabon: Makokou virus (MAKV). Arai et al. (2019b: 2) and Arai and Yanagihara (2020: 7) tetatively assigned this virus to the genus Mobatvirus. Hepadnaviridiae Orthohepadnavirus Hepatitis B virus (HBV) Drexler et al. (2013c: 16152) report on four out of 51 examined H. cf. ruber specimens from Gabon that were positive for roundleaf bat HBV (RBHBV). Nie et al. (2017: 89) refer to this virus as horseshoe bat HBV (HBHBV). Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) reported that two of the 117 specimens they identified as belonging to H. cf. ruber from Ghana tested positive for Morbillivirus. Two of the 337 specimens from Gabon, they identified as H. cf. caffer/ruber also tested positive for Morbillivirus and one other specimen for Rubulavirus. In their overview table, Maganga et al. (2014a: 8) report the following viruses were already found on H. cf. ruber: Rift Valley Fever virus (RVF), African Chiroptera Report 2020 Rubulavirus, Coronavirus. Morbillivirus 239 unclassified, DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Burkina Faso, Burundi, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Malawi, Mali, Mozambique, Niger, Nigeria, Rwanda, São Tomé and Principé, Senegal, Sierra Leone, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia. Figure 65. Distribution of Hipposideros ruber Hipposideros tephrus (Cabrera, 1906) 1861. Phyllorrhina bicornis Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 7. Publication date: 1861. Type locality: Eritrea: Kérén region [ca. 15 45 N 38 20 E] [Goto Description]. Lectotype: ZMB 2795: ♂, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: undated. Formerly in SMNS. Lectotype designated by Turni and Kock (2008: 36). - Comments: Meester et al. (1986: 43) and Turni and Kock (2008:36) mentioned it as Phyllorhina bicornis. *1906. Hipposiderus tephrus Cabrera, Bol. r. Soc. espan. Hist. nat. , 6: 358. Publication date: June 1906. Type locality: Morocco: Mogador [=Essaouira] [31 31 N 09 46 W]. Holotype: MNCN 110: ad ♂. Collected by: Mr. Martínez de la Escalera; collection date: 29 August 1905. See Ibáñez and Fernández (1989: 11). Paratype: MNCN 111: ad ♀. See Ibáñez and Fernández (1989: 11). (Current Combination) 1939. Hipposideros braima Monard, Arq. Mus. Bocage, 10: 73, fig. 5. Publication date: March 1939. Type locality: Guinea-Bissau: Bagíngarà [12 35 N 14 28 W] [Goto Description]. Holotype: [Unknown] ad ♀. Original number: 602. See Monard (1939: 73). 1965. Hipposideros braimah: Rosevear, Bats of West Africa, 221, 226. - Comments: Lapsus for braima Monard, 1939 (see Kock, 1969a: 130). 2014. Hipposideros (caffer) tephrus: Moores and Brown, Go-South Bull., 11: 19. (Name Combination) ? Hipposideros caffer tephrus: ? Hipposideros caffer: ? Hipposideros tephrus: (Name Combination, Current Spelling) TAXONOMY: Hayman and Hill (1971) and Simmons (2005: 376) recognised it as a subspecies of caffer. Vallo et al. (2009: 200) suggest that this taxon is elevated to species rank based on the genetic distance, the small size and the wide geographical separation from South-African true Hipposideros caffer. Phyllorine cendrée, Rhinolophe de Cafrerie. German: Marokkanische Rundblattnase. Patterson et al. (2020: 121) place it in the ruber subgroup of the bicolor group. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001) in view of its wide distribution, presumed large population, and because it is unlikely to be declining at the rate required to qualify for listing in a threatened category (Monadjem and Shapiro, 2017a). COMMON NAMES: Czech: pavrápenec sahelský. Dutch: Cabrera's rondbladneus. English: Lesser leaf-nosed bat, Lesser (Sundevall’s) Leaf-nosed Bat. French: Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Shapiro, 2017b) 240 ISSN 1990-6471 Regional None known. MAJOR THREATS: Human disturbance to roosting sites (caves) could have a negative effect (Monadjem and Shapiro, 2017b). CONSERVATION ACTIONS: No specific conservation measures in place. May occur in several protected areas throughout its range (Monadjem and Shapiro, 2017b). GENERAL DISTRIBUTION: Morocco to Senegal, and Yemen (Vallo et al., 2009; Benda et al., 2011b: 29). H. tephrus is confined to the coastal areas of northern Morocco. Probably Mauritania (Padial and Ibáñez, 2005: 240); Ethiopia (Lavrenchenko et al., 2004b: 133); SW Saudi Arabia. Possibly Gambia (Emms and Barnett, 2005: 50), Guinea (Fahr and Ebigbo, 2003: 128, as "Hipposideros cf. caffer"), Benin (Capo-Chichi et al., 2004: 162), Liberia (Fahr, 2007a: 104). Native: Burkina Faso (Kangoyé et al., 2015a: 608); Ethiopia (Nyssen et al., 2020: 9); Mauritania (Allegrini et al., 2011: 2); Morocco (Disca et al., 2014: 209; Moores and Brown, 2014: 20); Yemen (Benda et al., 2011b: 29). ECHOLOCATION: Mayer et al. (2007) reported a CF-component of 133 to 137 kHz for Morocccan specimens. See also Jones et al. (1993). Karyotype - Koubínová et al. (2010b) report on three females and two males from Senegal, 2n= 32, Fna= 60 and FN= 64, they comprised of 30 biarmed autosomes, including four metacentric, eight submetacentric and three subtelocentric pairs. A secondary constriction situated near the centromere was observed in one pair of the medium-sized submetacentric autosomes (approximately the 9th largest pair). The X chromosome was identified as a large metacentric element, the Y chromosome as a medium submetacentric. Protein / allozyme - Unknown. POPULATION: Has a wide distribution and the population is presumed to be large, but more precise informations are not available (Monadjem and Shapiro, 2017b). Trend:- 2016: Unknown (Monadjem and Shapiro, 2017b). PARASITES: Okafor (1988: 11) reported the presence of the cestode Oochoristica agamae Baylis, 1919 (Eucestoda, Linstowiidae) from H. caffer tephrus specimens from Nsukka, Nigeria. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Burkina Faso, Côte d'Ivoire, Eritrea, Ethiopia, Ghana, Mali, Mauritania, Morocco, Niger, Nigeria, Senegal, Sierra Leone, South Sudan, Sudan, The Gambia, Uganda. Disca et al. (2014: 226) indicate that the type of call is FC-FM, with the following parameters: Fstart: 147.1 ± 1.1 kHz, Fend: 128.2 ± 5.4 kHz, Fpeak: 146.8 ± 1.3 kHz, bandwidth: 19 ± 4.7 kHz, duration: 6.6 ± 1.1 msec. They also mention (p. 213) that the maximum energy is in the FC part of the third harmonic. The start frequency is also at least 10 kHz above the value provided by Dietz et al. (2009). Moores and Brown (2014: 21) recorded the following values for social calls at Bou Jerif fortress, Morocco: Fpeak: 46.7 kHz, Fstart: 55 kHz, Fend: 29 kHz, duration: 8 msec. MOLECULAR BIOLOGY: DNA - Vallo et al. (2009). Figure 66. Distribution of Hipposideros tephrus Genus Macronycteris Gray, 1866 *1866. Macronycteris Gray, Proc. zool. Soc. Lond., 1866, I (vi): 82. Publication date: May 1866 [Goto Description]. - Comments: Type species: Rhinolophus gigas Wagner, 1845, by monotypy (see Decher and Fahr, 2005: 1; Jackson and Groves, 2015: 246). - Etymology: African Chiroptera Report 2020 241 From the Greek "μακρός", meaning large and "νυκτερις", meaning bat (see Palmer, 1904: 393). TAXONOMY: This taxon was longtime considered to be a synonym of Hipposideros, but molecular analyses of Cyt-b by Foley et al. (2017) indicated that its representatives are sufficiently different from that genus to be assigned to a separate genus. Currently (Simmons and Cirranello, 2020) recognized species of the genus Macronycteris: commersoni (E. Geoffroy Saint-Hilaire, 1813); cryptovalorona (Goodman, Schoeman, Rakotoarivelo and Willows-Murno, 2016); gigas (Wagner, 1845); thomensis (Bocage, 1891); vittatus (Peters, 1852). MOLECULAR BIOLOGY: Karyotype Foley et al. (2017: 11) indicate that the karyotype of the genus Hipposideros is very conservative, with almost all species having 2n = 32. Both M. commersoni and M. gigas, however, have 2n = 52. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". †Macronycteris besaoka (Samonds, 2007) *2007. Hipposideros besaoka Samonds, Acta Chiropt., 9 (1): 49, fig 5A - C, 6A - D. Type locality: Madagascar: Anjohibe cave [Goto Description]. Holotype: UA 9478:. Right maxialla: see Samonds (2007: 49). - Etymology: The word besaoka is derived from the Malagasy and means 'big chin' (see Samonds, 2007). - Phonetics: bay-SOH-ka. (Current Combination) 2019. M[acronycteris] besaoka: Rakotoarivelo, Goodman, Schoeman and Willows-Munro, PeerJ, 7 (e5866): 15. Publication date: 17 January 2019. (Current Combination) TAXONOMY: Goodman et al. (2016: 435) examined some besaoka skulls and found these to have a more robust mandibula and lower teeth than both other Madagascan "Hipposideros" [=Macronycteris"] species (commersoni and cryptovalorna), but further DNA study is required to see whether it is genetically distinct from either of them. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Samonds (2007: 49 - 53) found samples in deposits (TW-10) dated at 10,000 yrs old or younger within the Anjohibe cave, Madagascar. Timeframe: Pleistocene (Brown et al., 2019: Suppl.). HABITAT: Rakotoarivelo et al. (2019: 14) presume that M. besaoka was ecologically similar to M. commersoni. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Macronycteris commersoni (E. Geoffroy St.-Hilaire, 1813) *1813. Rhinolophus Commersonii E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 20: 263, pl. 5. Publication date: 1813. Type locality: Madagascar: Tuléar province: Fort Dauphin [25 02 S 47 00 E] [Goto Description]. Neotype: FMNH 175972: ad ♀, skull and alcoholic. Collected by: Steven M. Goodman; collection date: 9 December 2002; original number: SMG 13401. Presented/Donated by: ?: Collector Unknown. Type locality: Madagascar: Province de Fianarantsoa, Parc National de l’Isalo, along Sahanafa River, near foot of Bevato Mountain, 28 km south-east of Berenty-Betsileo, 22°19.0'S, 45°17.6'E, 550 m a.s.l. (see Goodman et al., 2016: 437). - Etymology: Geoffroy's description was based on drawings and notes found among Mr. Philibert Commerson's papers, in who's honour Geoffroy named this species (see Smithers, 1983: 133; Taylor, 2005). 2017. M[acronycteris] commersonii: Foley, Goodman, Whelan, Peuchmaille and Teeling, Acta Chiropt., 19 (1): 10. Publication date: June 2017. (Current Combination) 2017. Macronycteris commersonii: Sotero-Caio, Baker and Volleth, Genes, 8 (272): 9. Publication date: 13 October 2017. (Name Combination, Alternate Spelling) 2018. Macronycteris commersoni: Reher, Ehlers, Hajatiana and Dausmann, J. Comp. Physiol., B 188 (6): 1015. Publication date: 18 August 2018. (Name Combination) ? Hipposideros commersoni commersoni: (Name Combination) 242 ISSN 1990-6471 ? ? Hipposideros commersoni: (Name Combination) Hipposideros sp. cf. H. commersoni: TAXONOMY: Simmons (2005: 369, 372, 377) restricts "Hipposideros commersoni" to Madagascar, and assigns "H. gigas", "M. vittatus" (including marugensis) and "H. thomensis" as the mainland forms based on morphology and echolocation differences. A combined molecular and morphological study by Rakotoarivelo et al. (2015: 1) indicates that cryptic species exist in Madagascan M. commersoni, but also that a further fine-scale phylogeographic study is needed to fully resolve the systematics of this species (see M. cryptovalorona). Rakotoarivelo et al. (2019: 1) investigated this further and found several lineages in bats from the western part of Madagascar: the animals from northern Madagascar formed a single monophyletic clade (clade C), whereas the animals from the southwestern part of the island (clade B) showed more phylogeographical partitioning where different mtDNA haplotypes frequencies were detected between populations from different bioclimatic regions. Goodman et al. (2016: 439) point out that the orginal spelling by Geoffroy (1813) was "Rhinolophus Commersonii", but that most of the subsequent authors spelled it with only one "i". They refer to article 33.3.1 of the ICZN code to maintain the usage of the single "i" version. COMMON NAMES: Afrikaans: Commerson se bladneusvlermuis, Commerson-bladneusvlermuis, Commersonse Blaarneusvlermuis. Czech: pavrápenec Commersonův. English: Commerson's Leafnosed Bat, Giant Leaf-nosed Bat, Commerson's roundleaf bat, Commerson's Rhinolph. French: Chauve-souris de Commerson à nez feuillu, Phyllorine de Commerson. German: Madagaskar-Rundblattnase, RiesenRundblattnase, Marungu-Rundblattnase. Portuguese: Morcego de nariz enfolhado de commerson. Ukrainian: Листоніс Коммерсона [= Lystonis Kommersona]. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Sabatier and Legendre (1985: 23) reported the presence of M. commersoni (and Hipposideros sp.) in the Tsimanampetsotsa deposit. Samonds (2007: 45 - 46, 58 - 60) found samples in deposit (OLD SE locality) dated at 10,000 yrs old or younger and deposit (SS2) which was collected from the surface near the main cave entrance, the dating attemps revealed that this sample was contaminated, therefore no date is known, these two sites identified specimens belonging to M. sp. cf. commersoni. Samonds (2007: 54 - 55) also found M. commersoni within the deposit (NCC-1 locality) dating at 69,000 to 86,800 years old, from within the Anjohibe cave, Madagascar. Gunnell et al. (2014: 3) refer to Goodman and Junkers (2013), who report on fossil material from the Andrahomana cave in Madagascar. CONSERVATION STATUS: Global Justification This species is listed [as Hipposideros commersoni] as Near Threatened (NT ver 3.1 (2001)) in view of the significant threat from hunting in the west, which is likely to have resulted in a decline in the region of 20 - 25 % over the past 15 years. Almost qualifies as threatened under criterion A. However, as this species is generally widespread across Madagascar, occurs both in and outside a number of protected areas, and has an ability to tolerate some degree of habitat modification, it is unlikely to be declining fast enough to warrant listing in a threatened category (Andriafidison et al., 2008d; IUCN, 2009). Assessment History Global 2008: NT ver 3.1 (2001) (Andriafidison et al., 2008d; IUCN, 2009). 2004: Not assessed (IUCN, 2004). Regional None known. MAJOR THREATS: Andriafidison et al. (2008d) state that besides habitat loss, it is also threatened by hunting and is particularly vulnerable at roosting sites where bats are hunted as they emerge at dusk (Goodman et al., 2008d; Jenkins and Racey, 2008). It can be legally hunted on between the first of February and the first of May (Anonymus, 2007b: 6). In the extreme south-west of Madagascar, there were an estimated 140,000 individuals harvested for food annually between January and March (Goodman, 2006). This hunting is thought to occur throughout western Madagascar in areas where local people live in close proximity to roosting colonies (Jenkins and Racey, 2008). CONSERVATION ACTIONS: Andriafidison et al. (2008d) report that this species occurs in a number of protected areas as well as other forest areas that are actively managed for conservation (Goodman et al., 2005a; African Chiroptera Report 2020 Rakotoarivelo and Randrianandrianina, 2007; Ranivo and Goodman, 2007b). Due to the existence of several clades within M. commersoni, Rakotoarivelo et al. (2019: 16) stress the conservation importance of the localities where clade C was found to occur: Analamera, Ankarana, Montagne d'Ambre, and Marovaza. GENERAL DISTRIBUTION: Madagascar; adjacent small islands. GEOGRAPHIC VARIATION: Ranivo and Goodman (2007b: 48) indicate that there is a clinal variation in commersoni on Madagascar, where the individuals from the north are larger than those from the south. In the Isalo region, both sizes were found together, which might indicate that there are two cryptic species. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 56) reported that the baculum of commersoni has a bifurcated distal tip, a straight central shaft and a basal portion that is narrower than the distal part; length: 3.73 ± 0.540 (3.00 - 4.52) mm, width: 1.92 ± 0.330 (1.48 - 2.41) mm. In the bat's clades (sensu Rakotoarivelo et al., 2015), they found two morphotypes with some slight differences: Morphotype A/Clade B: distinct bicurcated distal tip, sometimes with an indentation; length: 3.77 ± 0.545 (3.00 - 4.52) mm, width: 1.94 ± 0.325 (1.50 - 2.41) mm; Morphotype B/Clade C: "This same structure is also found in certain animals allocated to Clade C, with the exception of two" (unfortunately, without further details); length: 3.05, 3.37 mm, width: 1.48, 1.69 mm. ECHOLOCATION: In Madagascar Russ et al. (2001) recorded a constant-frequency of 64 kHz. Kofoky et al. (2009: 379) reported the calls of 28 individuals from six different sites in Madagascar, which produced narrowband and long CF calls which terminate with brief FM elements with a high duty cycle (31.2 %) and maximum energy at about 66.6 kHz. The calls consisted of a maximum of three harmonics most energy in the fundamental or second harmonic. Ramasindrazana et al. (2015b: 85) reported a sexual dimorphism in bats from the western half of Madagascar. In males, the average forearm length was 93.1 mm and they emitted calls at 68.6 kHz. In females, the average forearm length was 83.9 mm and their calls were at 72.9 kHz. They also found that females from the far northern part of the island (Ankarana) emitted calls that were much lower than expected from size. 243 Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 61 kHz and duration: 12 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - 2N = 52 and FN = 60, containing one large metacentric, a medium-sized submetacentric, three pairs of medium-to-smallsized metacentrics. The others are acrocentric. The X chromosome is a large sub-metacentric one and the Y an acrocentric (see Richards et al., 2016d: 187, 191). Protein / allozyme - Unknown. HABITAT: Dammhahn and Goodman (2013: 108) indicate that the lower one-half of forest and perhaps partially open areas form the foraging habitat of this species. The home range of males is 31.8 ± 9.2 ha and of females 41.7 ± 12.9 ha (Razafimanahaka et al., 2016: 423). HABITS: Bouchard (2001b: 109) mentions that males scentmark their territories and mates by rubbing them with their glandular areas. ROOST: M. commersoni roosts singly in trees at some sites (Jenkins and Racey, 2008). Razafimanahaka et al. (2016: 427) found that these roosts are generally on braches of trees (Uapaca ferruginea Baill., Canarium boivinii Engl., Diospyros sp., Gaertnera sp. and Erythroxylum corymbosum Boivin ex Baill.) with a diameter at breast height of about 8.2 ± 0.7 cm and at 5.4 ± 0.2 m above the ground. Aucoumea klaineana Pierre [angouma, gaboon, or okoumé], an introduced tree in plantations, is hardly used by the bats. The mean distance between the roost and the foraging area was 313.9 ± 15.3 m (max: 1.6 km). Reher et al. (2018: 1024) indicated that the bats chose to roost in the hottest and most humid cave in the region, where the environment was more buffered than in the cooler caves, which were prefered by other bat species. MIGRATION: Reher et al. (2019: 121) found that M. commersoni was only present in two southwestern Madagascan caves during the wet season, whereas other species such as Triaenops menamena and Miniopterus mahafaliensis were present in both the dry and wet season. 244 ISSN 1990-6471 DIET: Razakarivony et al. (2005) examined the stomachs of 11 individuals and found that 90 % of the diet were Coleoptera. Rakotoarivelo et al. (2007) found M. commersoni mainly to consume Coleoptera, but also included large-bodied taxa such as Cicadidae, Lucanidae, and Passalidae. In the Tsimanampetsotsa NP, Ramasindrazana et al. (2012: 120-121) reported feacal pellets containing 76.2 ± 19.1 volume percent Coleoptera, 20.0 ± 20.0 Blattaria, 2.4 ± 1.0 Homoptera/Hemiptera and 1.4 ± 0.9 Hymenoptera. No Blattaria were found during the wet season, whereas the percentage Coleoptera rose to 93.0 (from 72.0 in the dry season). Rakotondramanana et al. (2015: 78) report the following prey items (mean volume percentages ± 1 SD, mean frequency percentages ± 1 SD): Coleoptera (46.6 ± 38.85, 77.7 ± 36.36); Hymenoptera (6.6 ± 11.49, 26.1 ± 40.70); Lepidoptera (8.2 ± 13.34, 26.1 ± 37.85); Isoptera (37.4 ± 40.53, 49.2 ± 48.41); Homoptera (0.9 ± 3.49, 6.7 ± 19.85); Blattoidea (0.2 ± 0.98, 0.2 ± 0.98); Arachnida (0.04 ± 0.20, 0.77 ± 3.92). PREDATORS: Goodman et al. (1991: 22) reported an almost intact skull in a pellet from the Madagascar Longeared Owl Asio madagascariensis. Additionally, Goodman and Griffiths (2006) found that M. commersoni comprised 0.6 % of the total biomass predated by the barn owl (Tyto alba). Mikula et al. (2016: Supplemental data) mention the Madagascan serpent eagle (Eutriorchis astur (Sharpe, 1875)) as avian diurnal predator. ACTIVITY AND BEHAVIOUR: Preliminary observations in Madagascar reveal that foraging behavior of M. commersoni involves short hunting flights from night perches (Russ et al., 2003). Reher et al. (2018: 1015) found that M. commersoni can enter in a torpor state to cope with the dry and unpredictable environmental conditions in the dry spiny forest of south-western Madagascar. On average, it took the bats 4.1 ± 1.4 minutes to enter torpor and 12.3 ± 3.3 minutes to arouse (p. 1020). REPRODUCTION AND ONTOGENY: Razafimanahaka et al. (2016: 429) recorded flying juveniles in early February and March, which suggests that births take probably place in January. They were also unable to catch females in nets in January, possibly because they were not foraging or because they had moved to materity roosts. PARASITES: BACTERIA: Gomard et al. (2016: 5) reported the presence of Leptospira bacteria in 13 out of 27 tested bats (48.1 %). VIRUSES: Nieto-Rabiela et al. (2019: Suppl.) reported the following viruses from "Hipposideros commersoni": African bat icavirus, Bat coronavirus, Bat flavivirus, Bat hepacivirus, Bat paramyxovirus, Bat pegivirus, Bat rhabdovirus, Leopards Hill Virus, Shimoni bat lyssavirus Rhabdoviridae The serum of one of six bats tested by Mélade et al. (2016a: 6) showed a reaction against Duvenhage lyssavirus, and one of four against European bat lyssavirus 1. UTILISATION: Goodman (2006) observed in February 2005, in south-western Madagascar, that local people were extensively hunting M. commersoni. This coincided with the onset of famine in the region. Goodman (2006) describes the hunting method used by local inhabitants. In PN d'Ankarafantsika, local people - while collecting forest tubers - occasionally encounter roosting M. commersoni which are taken back to the village and eaten (Jenkins and Racey, 2008). In the villages M. commersoni is either roasted over a fire or boiled. Goodman (2006) concluded that harvest level of M. commersoni (ca. 70,000 - 140,000 bats annually) surpasses the likely productivity of the species in the area. Goodman et al. (2002) suggested that hunting at the Mitoho Cave in PN de Tsimanampetsotsa may have been the reason M. commersoni abandoned the site. See also Goodman et al. (2008d) and Jenkins and Racey (2008). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar, Réunion. African Chiroptera Report 2020 245 Figure 67. Distribution of Macronycteris commersoni Macronycteris cryptovalorona (Goodman, Schoeman, Rakotoarivelo and Willows-Munro, 2016) *2016. Hipposideros cryptovalorona Goodman, Schoeman, Rakotoarivelo and Willows-Munro, Zool. J. Linn. Soc., 177 (2):428, 443. Publication date: 20 May 2016. Type locality: Madagascar: Fianarantsoa province: Parc National de l'Isalo, along Sahanafa River, near foot of Bevato: Berenty-Betsileo, 28 km SE [22 19.0 S 45 17.6 E, 550 m] [Goto Description]. Holotype: FMNH 175970: ad ♀, skull and alcoholic. Collected by: Steven M. Goodman; collection date: 9 December 2002; original number: SMG 13399. Presented/Donated by: ?: Collector Unknown. Type locality: Madagascar: Province de Fianarantsoa, Parc National de l’Isalo, along Sahanafa River, near foot of Bevato, 28 km south-east of Berenty-Betsileo, 22°19.0'S, 45°17.6'E, 550 m a.s.l. Paratype: FMNH 184173: ♀. Collected by: Steven M. Goodman; collection date: February 2005; original number: SMG 14582. Presented/Donated by: ?: Collector Unknown. - Etymology: The name is derived from the Greek "kryptos", meaning hidden or concealed, and from the Sakalava dialect of Malagasy "valorona", which is the local vernacular name of Hipposideros and refers to its distinct nasal structure, and can be translated as "eight nose", referring to the complicated and multiple layers of the nose structure (see Goodman et al. (2016: 444). 2017. M[acronycteris] cryptovalorona Foley, Goodman, Whelan, Puechmaille and Teeling, Acta Chiropt., 19 (1): 10. Publication date: June 2017. (Current Combination) 2019. Macronycteris cryptovalorona: Monadjem, Handbook Mammals of the World, vol 9 Bats: 232.. (Name Combination, Current Combination) TAXONOMY: Analysing the cyt b sequences of 148 Madagascan M. commersoni s.l. specimens, Goodman et al. (2016: 428) found three well-supported monophyletic clades: A, B, and C. Clades B and C form a monophyletic lineage, which they were able to refer to M. commersoni s.s. Clade A showed a 9 to 11 % sequence variation with M. commersoni s.s., enough to warrant a separate species status. This new species is known from two adult females only. COMMON NAMES: English: Madagascar cryptic leaf-nosed bat. French: La Phyllorhine cryptique de Madagascar. German: Kryptische Madagaskar-Rundblattnase. Figure 68. Distribution of Macronycteris cryptovalorona 246 ISSN 1990-6471 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Macronycteris gigas (Wagner, 1845) *1845. Rhinolophus gigas Wagner, Arch. Naturgesch. Berlin, 11 (1): 148. Publication date: 1845. Type locality: Angola: Benguela [12 34 S 13 24 E, 160 m]. 1906. H[ipposiderus]. gigas gambiensis K. Andersen, Ann. Mag. nat. Hist., ser. 7, 17 (97): 42. Publication date: 1 January 1906. Type locality: The Gambia: "Gambia" [Goto Description]. Holotype: BMNH 1842.9.27.36: ad ♀, alcoholic (skull not removed). Presented/Donated by: 13th Earl of Edward Smith Stanley Derby. 1917. Hipposideros gigas niangaræ J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 438, pl. 51, fig. 1. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Niangara [03 24 N 27 52 E] [Goto Description]. Holotype: AMNH 49103: ad ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 2 June 1913; original number: 2069. 1939. Hipposideros gigas viegasi Monard, Arq. Mus. Bocage, 10: 70. Publication date: March 1939. Type locality: Guinea-Bissau: Madina Boé [11 45 N 14 13 W] [Goto Description]. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 909. Syntype: [Unknown] ♂, skin and skull. Collected by: P. Rigaux; collection date: 9 March 1938; original number: 910. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 911. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 912. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 913. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 914. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 917. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 918. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 919. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 920. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 921. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 922. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 923. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 924. Syntype: [Unknown] ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 926. Syntype: NMBA 5266/9283: ♂. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 915. Syntype: NMBA 5267/9284: ♀. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 908. Syntype: RMCA 15305: ♂, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 925. Syntype: RMCA 15306: ♀, skin and skull. Collected by: Dr. Albert Monard; collection date: 9 March 1938; original number: 916. - Comments: Monard (1939: 70) mentions a series of 19 specimens (nrs 908 to 926 and 14 skulls). 2017. Macronycteris gigas: Foley, Goodman, Whelan, Puechmaille and Teeling, Acta Chiropt., 19 (1): 7. Publication date: June 2017. (Current Combination) ? Hipposideros commersoni gigas: (Name Combination) ? Hipposideros commersoni niangarae: ? Hipposideros commersoni: (Name Combination) ? Hipposideros gigas gigas: (Name Combination) ? Hipposideros gigas niangarae: ? Hipposideros gigas: (Name Combination, Current Combination) ? Hipposideros marungensis marungensis: ? Hipposideros marungensis: African Chiroptera Report 2020 247 TAXONOMY: McWilliam (1982: 9) found two forms of "Hipposideros commersoni" occurring sympatrically in Kenya, which he separated into "H. commersoni" and "H. gigas", with the latter species being significantly larger and heavier than the prior. Monadjem et al., 2010d: 538 as H. vittatus), Transvaal (South Africa (Monadjem et al., 2010d: 538 as H. vittatus), Malawi (Happold et al., 1988; Happold and Happold, 1997b: 818; Monadjem et al., 2010d: 538 as H. vittatus) and Mozambique (Monadjem et al., 2010d: 538, Monadjem et al., 2010c: 381 as H. vittatus). Simmons (2005: 369, 372, 377) restricts "Hipposideros commersoni" to Madagascar, and assigns H. gigas, H. vittatus (including marugensis) and H. thomensis as the mainland forms based on morphology and echolocation differences. Meester et al. (1986) state that Hayman and Hill (1971) note that marungensis may be antedated by vittata Peters, 1852, from Ibo Island, which is more often regarded as a synonym of the subspecies gigas Wagner, 1845 (Allen, 1939a, Ellerman et al., 1953, Hill, 1963a). Rainho and Ranco (2001: 54) report "H. commersoni" from Guinea-Bissau (see also Peters, 1870; Seabra, 1900c; Monard, 1939; Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992). The karyotypic studies for the African mainland forms (M. vittatus and M. gigas) indicate that they share the exact same numbers and forms (Porter et al., 2010); Rautenbach et al., 1993; Koubínová et al., 2010b). Koubínová et al. (2010b) suggest that African mainland forms may belong to the same species. COMMON NAMES: Afrikaans: Commerson se bladneusvlermuis, Commerson-bladneusvlermuis, Commersonse Blaarneusvlermuis. Chinese: 巨 蹄 蝠 . Czech: pavrápenec obrovský. English: Commerson's Leaf-nosed Bat, Giant Leaf-nosed Bat, Commerson's roundleaf bat, Commerson's Rhinolph. French: Chauve-souris de Commerson à nez feuillu, Phyllorine de Commerson. German: MadagaskarRundblattnase, Riesen-Rundblattnase, MarunguRundblattnase. Kinande (DRC): Mulima. Portuguese: Morcego de nariz enfolhado de commerson. GENERAL DISTRIBUTION: Gambia over Liberia (Monadjem et al., 2016y: 365 as Hipposideros gigas), Ghana (Decher and Fahr, 2007: 17 as Hipposideros gigas), Benin (Capo-Chichi et al., 2004: 162), Cameroon (Waghiiwimbom et al., 2019b: 6), Central African Republic (Lunde et al., 2001: 537), Republic of the Congo (Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 538 as H. vittatus; Bates et al., 2013: 335, as H. commersoni) to Ethiopia, south to Angola (Crawford-Cabral, 1989: 17, as H. c. gigas; Monadjem et al., 2010d: 538, as H. gigas and H. vittatus), Zambia (Ansell, 1967; Ansell, 1974; Monadjem et al., 2010d: 538 as H. vittatus), Zimbabwe (Monadjem et al., 2010d: 538 as H. vittatus), Namibia (Monadjem et al., 2010d: 538 as H. vittatus), Botswana (Smithers, 1971; DETAILED MORPHOLOGY: Baculum - Unknown. Brain - Kruger et al. (2010a) describe - based on three brains from Kenya, and using immunohistochemical methods - the nuclear organization of the cholinergic, catecholaminergic and serotonergic systems. Kruger et al. (2010b) also describe the distribution of Orexin-A immunoreactive cell bodies and terminal networks. ECHOLOCATION: O'Shea and Vaughan (1980) reported an ultrasonic frequency of 155 kHz for Masalani in Kenya. A highest frequency of 62 kHz was recorded at Sengwa in Zimbabwe by Jacobs (1996) and Fenton and Bell (1981), and at Pafuri in South Africa by Aldridge and Rautenbach (1987). Monadjem et al. (2010c: 381) reported from Mozambique [as M. vittatus] that the peak echolocation frequencies ranged between 64 - 66 kHz (ANABAT). Rainho et al. (2010a: 21) report [as Hipposideros thomensis] the calls of 10 individuals from Saõ Tomé. Monadjem et al. (2013b: 351) report the Fmax to be 52.0 ± 0.41 (51 - 53) kHz for four specimens from Mount Nimba. For two unsexed bats, two females and three males from the Shimoni cave in Kenya, Webala et al. (2019b: 15) reported an Fmax value ranging between 54.8 and 55.3, 53.4 and 54.3, and 53.4 and 54.1 kHz, respectively. The duration of the calls were respectively 12.3 to 14.6, 17.1 to 19.2, and 10.4 to 15.7 msec. Patterson et al. (2020: 137) mention a peak frequency of 53.4 - 54.8 kHz, which distinguishes this species from M. vittatta, which has a peak frequency of 64 - 70 kHz. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported for female individuals with a diplod number of 2n = 52, FN = 60?, and BA = 10?. Porter et al. (2010) 248 ISSN 1990-6471 found the same diploid number described by Rautenbach et al. (1993) for a single male Gabonese specimen reported as Hipposideros gigas, with six pairs of biarmed and 20 pairs of acrocentric chromosomes (including the sex chromosomes). Koubínová et al. (2010b) reported for two females from Senegal [as "Hipposideros gigas"] a 2n= 52, and FNa= 60, supposing the X chromosomes to be (sub)metacentric. The autosomes would then consist of one pair of large metacentrics, two pairs of medium-sized sub-metacentrics, one pair of small submetacentrics, one pair of small metacentrics, and 20 pairs of acrocentrics, the two smallest acrocentric pairs were of a dot-like appearance. Protein / allozyme - Unknown. HABITAT: Monadjem et al. (2016y: 365) recorded this species widely in forested and disturbed habitats in the Mount Nimba area, at altitudes between 460 and 1,060 m. In Angola, it occurs in lowland forest and moist savanna (Beja et al., 2019: 388). DIET: Andreas (2010: 266) indicates that there is some proof from Senegal that M. gigas might be carnivorous, as was already suggested by Eger and Mitchell (2003) for M. commersoni. REPRODUCTION AND ONTOGENY: Kurta and Kunz (1987: 82) report that, at birth, the young has about 25.2 % of the mother's body mass (28.0 versus 111.0 g). Kurta and Kunz (1987: 227) indicate that in Gabon, all adult females (of M. commersoni) were pregnant between July and October (with implantation occurring in mid June), and generally gave birth in October. The young is being fed with milk until the beginning of May. The males were in full spermiogenesis by mid April. PARASITES: DIPTERA: Nycteribiidae: Nycteribia schmidlii scotti Falcoz, 1923, Eucampsipoda africana Theodor, 1955, and Penicilidia fulvida Bigot, 1885 (Obame-Nkoghe et al., 2016: 5, Gabon) Streblidae: Brachytarsina allaudi (Falcoz, 1923) (Obame-Nkoghe et al., 2016: 5, Gabon) Szentiványi et al. (2019: Suppl.) indicated that a similar bat fly from a Gabonese "Hipposideros gigas" was carrying an Arsenophonus sp. bacteria. Raymondia huberi Frauenfeld, 1856 group (Obame-Nkoghe et al., 2016: 5, Gabon). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in "Hipposideros commersonii" [as they considered "H. vittatus" to be a separate species, the bats probably represent M. gigas]: Astroviruses. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 to 1999, for antibodies for SARS-CoV in sera of 16 individuals from the Oriental Province, DRC. None tested positive (0/16). Tong et al. (2009) found coronavirus RNA in this bat, collected in 2006 in Kenya. In Nigeria, Quan et al. (2010) screened the gastrointestinal tissue for the presence of coronaviruses by PCRs. Maganga et al. (2020: 1) reported viruses belonging to the genera Alphacoronavirus and Betacoronavirus from bats in Gabon. These were significantly present in July and October. Filoviridae - Filoviruses Marburgvirus: Towner et al. (2007) reported on one individual [as Hipposideros gigas] from Gabon, which they tested for Marburg virus RNA by conventional and real-time RT-PCR and for anti-Marburg virus IgG antibodies by ELISA. The results were negative. Ebolavirus: Fiorillo et al. (2018: 7) included "Hipposideros gigas" in their Ebola distribution model, indicating that there is evidence that suggests their relation with filoviruses infection cases, although they provide no details on this. Flaviviridae Pegivirus (BPgV) - 1 out of 5 Nigerian specimens examined by Quan et al. (2013: Table S5) was infected by clade K type Pegivirus. Maganga et al. (2014a: 5) tested 227 bats from Gabon of which only one tested positive for a Flavivurus. Nairoviridae Orthonairovirus Ishii et al. (2014: 2) isolated a new Nairovirus (tentatively called Leopards Hill Virus - LPHV) from a H. gigas specimen collected in the Leopards Hill cave in Lusaka, Zambia. In a test on 129 specimens from Gabon, Müller et al. (2016: 3) found 32 of them positive for Crimean Congo hemorrhagic fever virus (CCHFV). Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that two of the 20 specimens they examined from Ghana tested positive for Morbillivirus, and one for Rubulavirus. African Chiroptera Report 2020 In their overview table, Maganga et al. (2014a: 8) report that the following viruses were already found on "Hipposideros gigas": Rubulavirus, Morbillivirus unclassified, Flavivirus, Shimoni bat virus, SARS-like CoV. 249 (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Niger, Nigeria, São Tomé and Principé, Senegal, Tanzania, The Gambia, Togo. Picornaviridae Mischivirus - This virus was reported by Zeghbib et al. (2019: Suppl.) from a Congolese bat. Rhabdoviridae Lyssavirus - Rabies related viruses: In Nigeria, Quan et al. (2010) did not detect any lyssavirus-specific antigens from the brains by use of direct fluroescent anitbody testing. Shimoni: Kuzmin et al. (2010a) detected a new lyssavirus, Shimoni bat lyssavirus from the brain of a dead "H. commersoni" bat found in the coastal region of Kenya. Genetic distances demonstrated that this may be a new lyssavirus species. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Burkina Faso, Cameroon, Central African Republic, Congo, Congo Figure 69. Distribution of Macronycteris gigas Macronycteris vittatus (Peters, 1852) *1852. Phyllorrhina vittata Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 32, pl. 6; pl. 13, fig 7 - 13. Publication date: 1852. Type locality: Mozambique: Cap Delgado group [=Querimba islands]: Ibo Island [ca. 12 20 S] [Goto Description]. Syntype: ZMB 363: ♀, alcoholic, skull and skeleton. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008). Syntype: ZMB 37150: ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. Legs and wings missing, see Turni and Kock (2008: 40). - Etymology: From the feminine Latin adjective vittàta meaning "striped", referring to the occurrence of dorsal light bands in this species (Lanza et al., 2015 149). 1879. Phyllorhina commersoni: Dobson, Proc. zool. Soc. Lond., 1878, IV: 879. Publication date: April 1879. 1887. Phyllorhina commersonii var. marungensis Noack, Zool. Jb. Syst., 2 (2): 272, pl. 10, fig. 31 - 33. Publication date: 7 May 1887. Type locality: Congo (Democratic Republic of the): W Shore Lake Tanganyika, Marungu: Qua Mpala [=Mpala / Pala] [06 43 S 29 31 E] [Goto Description]. Syntype: ZMB 6312: ♂, skin and skull. Collected by: Richard Böhm; collection date: 12 July 1883. See Turni and Kock (2008). Syntype: ZMB 86257: skin and skull. Collected by: Richard Böhm; collection date: 12 July 1883. Old number A4269; see Turni and Kock (2008). Syntype: ZMB 86258: skull only. Collected by: Richard Böhm; collection date: 12 July 1883. Old number A4268; see Turni and Kock (2008). Syntype: ZMB 86259: skull only. Collected by: Richard Böhm; collection date: 12 July 1883; original number: A4270. See Turni and Kock (2008). - Comments: Actually, in Noack (1887: 272), the name is mentioned as "Phyllor hinacommersonii Peters, var. marungensis N.". Noack (1887: 272) also mentions five specimens, including one female, collected in July-August. Ansell and Dowsett (1988: 34) refer to Hayman and Hill (1971: 28) who indicate that marungensis may be antedated by vittata Peters, 1852. 1891. Phyllorhina commersoni var. thomensis Bocage, J. Sci. mat. phys. nat., ser. 2, 2 (6): 88. Publication date: September 1891. Type locality: São Tomé and Principé: São Tomé Island [Goto Description]. - Comments: Bocage (1891: 88) mentions one male and one female specimen. 250 ISSN 1990-6471 1904. 1993. 2017. 2017. 2018. 2020. ? ? ? ? ? Hipposideros Commersoni mostellum Thomas, Ann. Mag. nat. Hist., ser. 7, 13 (77): 385. Publication date: 1 May 1904. Type locality: Kenya: Tana River [ca. 02 33 S 40 31 E, 700 ft] [Goto Description]. Holotype: BMNH 1889.3.8.3: ad ♂, skin and skull. Presented/Donated by: H.C.V. Hunter Esq. Hipposideros commersoni matungensis: Koneval, Honors Projects - Illinois Wesleyan University: 1. (Lapsus) M[acronycteris] thomensis: Foley, Goodman, Whelan, Puechmaille and Teeling, Acta Chiropt., 19 (1): 10. Publication date: June 2017. (Current Combination) M[acronycteris] vittatus: Foley, Goodman, Whelan, Puechmaille and Teeling, Acta Chiropt., 19 (1): 10. Publication date: June 2017. (Current Combination) Macronycteris vittata: Musila, Monadjem, Webala, Patterson, Hutterer, De Jong, Butynski, Mwangi, Chen and Jiang, Zool. Res. (China), 40 (1): 21 (for 2019). Publication date: 17 October 2018. (Current Combination) Macronycteris vitattus: Weier, Keith, Neef, Parker and Taylor, Diversity, 12 (188): 5. Publication date: 11 May 2020. (Lapsus) Hipposideros commersoni marungensis: Hipposideros commersoni: (Name Combination) Hipposideros thomensis: (Name Combination) Hipposideros vittatus: (Name Combination, Current Combination) Macronycteris vittatus: (Alternate Spelling) TAXONOMY: Simmons (2005: 369, 372, 377) restricts "Hippposideros commersoni" to Madagascar, and assigns H. gigas, H. vittatus (including marugensis) and H. thomensis as the mainland forms based on morphology and echolocation differences. Meester et al. (1986) state that Hayman and Hill (1971) noted that marungensis may be antedated by vittata Peters, 1852, from Ibo Island, which is more often regarded as a synonym of the subspecies gigas Wagner, 1845 (Allen, 1939a, Ellerman et al., 1953, Hill, 1963a). The karyotypic studies for the African mainland forms (M. vittatus and M. gigas) indicate that they share the exact same numbers and forms (Porter et al., 2010); Rautenbach et al., 1993; Koubínová et al., 2010b). Koubínová et al. (2010b) suggests that African mainland forms may belong to the same species. Foley et al. (2017: 3) indicate that the morphological characters used to distinguish M. gigas from M. vittatus are not straightforward and that these are generally used in combination with habitat and distribution data, which is complicated by the fact that their distributions partially overlap. They also indicate that the differences between the forms gambensis [sic], marungensis, niangarae and viegasi have been based on non-molecular characters, and as such the assignment of these synonyms is not yet definitive. COMMON NAMES: Chinese: 大 白 纹 蹄 蝠 . Czech: pavrápenec pruhovaný. English: Striped Leaf-nosed Bat. German: Streifen-Rundblattnase. Italian: Ipposìdero vittàto. CONSERVATION STATUS: Assessment History Global 2008: NT ver 3.1 (2001) (Mickleburgh et al., 2008z; Mickleburgh et al., 2008z). Bioko: Conenna et al. (2017: Suppl.) mentioned the status of "Hipposideros thomensis" as Least Concern (LC). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016v). 2004: considered a vagrant not assessed, single locality (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). GENERAL DISTRIBUTION: Gambia over Benin (Capo-Chichi et al., 2004: 162), Ghana (Decher and Fahr, 2007: 17 as H. gigas), Burkina Faso (Kangoyé et al., 2015a: 608); Central African Republic (Lunde et al., 2001: 537), Republic of the Congo (Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 538) to Ethiopia, south to Angola (CrawfordCabral, 1989: 17, as H. c. gigas; Monadjem et al., 2010d: 538, as H. gigas and H. vittatus; Taylor et al., 2018b: 62), Zambia (Ansell, 1967; Ansell, 1974; Monadjem et al., 2010d: 538), Zimbabwe (Monadjem et al., 2010d: 538), Namibia (Monadjem et al., 2010d: 538), Botswana (Smithers, 1971; Monadjem et al., 2010d: 538), Transvaal (South Africa (Monadjem et al., 2010d: 538)), Malawi (Happold et al., 1988; Happold and Happold, 1997b: 818; Monadjem et al., 2010d: 538) and Mozambique (Monadjem et al., 2010d: 538, Monadjem et al., 2010c: 381); Madagascar; African Chiroptera Report 2020 251 adjacent small islands; Saõ Tomé (Rainho et al., 2010a: 26 as H. thomensis). duration was 14.3 to 17.9 msec for the males and 10.6 to 17.8 for the females. Reported from the Tanzanian island of Pemba by O'Brien (2011: 287). In Sierra Leone, Weber et al. (2019: 21) recorded calls from a male and a female with a CF value of 62.5 (62.0 - 63.0) kHz. Rainho and Ranco (2001: 54) report "H. commersoni" from Guineu-Bissau (see also Peters, 1870; Seabra, 1900c; Monard, 1939; Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,043,916 km2. DETAILED MORPHOLOGY: Baculum - Unknown. Brain - Using immunohistochemical methods, Kruger et al. (2010a) describe the nuclear organization of the cholinergic, catecholaminergic and serotonergic systems based on three brains from Kenya. Kruger et al. (2010b) also describe the distribution of Orexin-A immunoreactive cell bodies and terminal networks. Koneval (1993) described several significant differences in the hyoid morphology of "Hipposideros commersoni matungensis" (from Kenya) and other bats of the family Rhinolophidae (including Hipposideridae): a modified stylohyal, fused to the auditory bulla; loss of the stylohyoid; addition of a new muscle, possibly from the stylopharyngeus; a modified mylohyoid profundus; and insertion of the ceratohyoid onto only the stylohyal. She found this morphology to be closest related to this of the east Asian H. armiger. ECHOLOCATION: O'Shea and Vaughan (1980) reported an ultrasonic frequency of 155 kHz for Masalani in Kenya. A highest frequency of 62 kHz was recorded at Sengwa in Zimbabwe by Jacobs (1996) and Fenton and Bell (1981), and at Pafuri in South Africa by Aldridge and Rautenbach (1987). From Mozambique, Monadjem et al. (2010c: 381) reported that the peak echolocation frequencies ranged between 64 - 66 kHz (ANABAT). Rainho et al. (2010a: 21) reports [as H. thomensis] the calls of 10 individuials from Saõ Tomé. Musila et al. (2018c: 44) [referring to Monadjem et al. (2010d: 162]) mention a high-duty, constant frequency call with a peak frequency of 61 kHz. Two males and 7 females from three localities in Kenya had Fmax values between respectively 67.9 and 70.1, and 64.3 and 69.7 kHz, whereas the Weier et al. (2020: Suppl.) reported on four calls from the Okavango River Basin with the following characteristics: Fmax: 60.23 ± 6.07 kHz, Fmin: 58.34 ± 6.08 kHz, Fknee: 59.52 ± 6.12 kHz, Fchar: 59.17 ± 6.21 kHz, slope: 3.49 ± 9.39 Sc, duration: 3.84 ± 1.36 msec. Patterson et al. (2020: 137) mention a peak frequency of 64 - 70 kHz, which distinguishes this species from M. gigas, which has a peak frequency of 53.4 - 54.8 kHz. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported for female individuals with a diplod number of 2n = 52, FN = 60?, and BA = 10?. Porter et al. (2010) found the same diploid described by Rautenbach et al. (1993) for a single male Gabonese specimen reported as "Hipposideros gigas", with six pairs of biarmed and 20 pairs of acrocentric chromosomes. While, Koubínová et al. (2010b) reported for two females from Senegal [as H. gigas] a 2n= 52, and FNa= 60, including a pair of large metacentrics, three pairs of medium-sized sub-metacentrics, one pair of small submetacentrics, one pair of small metacentrics, and 20 pairs of acrocentrics, the two smallest acrocentric pairs were of a dot-like appearance. Protein / allozyme - Unknown. ROOST: In Angola, M. vittata roosts in very large colonies in cave, mainly in the southern part of the country (Beja et al., 2019: 388). PREDATORS: Messana et al. (1985: 337) shows a photo, taken in a cave in Somalia, on which a dead body of "Hipposideros commersoni" is being fed on by Phaeophilacris sp. (Insecta: Orthoptera: Grillidae) and Tura mesopotamica Taiti & Ferrara, 1985 (Isopoda: Porcellionidae). REPRODUCTION AND ONTOGENY: A pregnant female was recorded on 26 March by Weber et al. (2019: 23) in Sierra Leone. PARASITES: HAEMOSPORIDA: Blood samples from Zambian M. vittata examined by Qiu et al. (2019: 236) revealed the presence of 252 ISSN 1990-6471 two Trypanosoma sp., very similar to T. conorhini and T. dionisii clades. DIPTERA: Streblidae: Ascodipteron variisetosum Maa, 1965 from Robertspoort, Liberia and Ghana, where the cysts were found under the upper arm (Haeselbarth et al., 1966: 107, hosts identified as H. commersoni). Raymondia huberi huberi Frauenfeld, 1855 (Shapiro et al., 2016: 254). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in "Hipposideros vittatus": Paramyxovirus. Coronaviridae - Coronaviruses Alphacoronavirus: SARS-CoV (subgenus Duvinacovirus) - Müller et al. (2007b) tested between 1986 to 1999, for antibodies for SARS-CoV in sera of 16 individuals from the Oriental Province, DRC. None tested positive (0/16). Tong et al. (2009) found coronavirus RNA (Kenya-bat-coronavirus [BtKY07]) from fecal material in 2006 in Kenya, and was similar to Rousettus sp. coronaviruses from China. In Nigeria, Quan et al. (2010) screened the gastrointestinal tissue for the presence of coronaviruses by PCRs. One out of 123 (0.8 %) Kenyan bats tested by Tao et al. (2017: Suppl.) was positive for CoV. Filoviridae - Filoviruses Marburgvirus - Towner et al. (2007) as H. gigas, tested 1 individual from Gabon for Marburg virus RNA by conventional and real-time RT-PCR and for anti-Marburg virus IgG antibodies by ELISA; the results were negative. Lyssavirus - Rabies related viruses In Nigeria, Quan et al. (2010) did not detect any lyssavirus-specific antigens from the brains by use of direct fluroescent antibody testing. Shimoni - Kuzmin et al. (2010a) detected a new lyssavirus, Shimoni bat lyssavirus from the brain of a dead "H. commersoni" bat found in the coastal region of Kenya. Genetic distances demonstrated that this may be a new lyssavirus species. Fikirini rhabdovirus (FKRV) Kading et al. (2013) report on this new rhabdovirus from a M. vittata specimen collected at Three Caves, Shimoni cave in Southern Kenya. Horton et al. (2014: Table S1) tested 77 Kenyan "H. commersoni" specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). ANTHROPOPHILOUS: Dandurand et al. (2018: 284) studied the influence of condensation-corrosion processes caused by urine and droppings on the walls of Drotsky's cave (Botswana) and their consequences on archaeological remains and cave art. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Botswana, Burundi, Cameroon, Central African Republic, Chad, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Ghana, Guinea-Bissau, Kenya, Madagascar, Malawi, Mozambique, Namibia, Niger, Rwanda, São Tomé and Principé, Senegal, Sierra Leone, Somalia, South Africa, Sudan, Tanzania, The Gambia, Togo, Zambia, Zimbabwe. Flaviviridae Hepacivirus (BHV) - Quan et al. (2013: Table S5) report 2 with clade A infected Kenyan specimens on a total of 75 tested (2.7 %), and 4 specimens (5.3 %) that were infected with clade D virus. Pegivirus (BPgV) - Quan et al. (2013: Table S5) also indicated that 2 of the 75 specimens were infected with a clade K type Pegivirus. Polyomaviridae Polyomavirus - Conrardy et al. (2014: 259) examined nine "Hipposideros commersoni" specimens from the Shimoni cave (Kenya), and found one of them testing positive for this virus. Figure 70. Distribution of Macronycteris vittatus Rhabdoviridae †Genus Palaeophyllophora Revilliod, 1917 1917. Palaeophyllophora Revilliod, Abh. Grossherz.-hess. Geol. Landesanst., 7: 1??. African Chiroptera Report 2020 253 †Palaeophyllophora tunisiensis (Ravel, 2016) *2016. ?Palaeophyllophora tunisiensis Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 368, figs 6, 7. Publication date: 30 September 2016. Type locality: Tunisia: Kassérine province: Dejbel Chambi National Park: Chambi [35 14 03 N 08 45 29 E, 630 m] [Goto Description]. - Etymology: The species name refers to the country from which uit was described: Tunisia. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. Family MEGADERMATIDAE H. Allen, 1864 *1864. Megadermatidae H. Allen, Monograph of North American Bats, xxiii, 1. - Comments: Type genus: Megaderma Geoffroy, 1810. See Handley (1980: 10) for the correct formation of the family name. (Current Combination) 1866. Megadermata Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 1865: 256. - Comments: In Part (see Taylor, 1934). Originally included the genera Rhinopoma É. Geoffroy, 1818; Megaderma É. Geoffroy, 1810; Nycteris É. Geoffroy & G. Cuvier, 1795; and Nyctophilus Leach, 1821 (see Jackson and Groves, 2015: 241). 1866. Megadermina Gray, Proc. zool. Soc. Lond., 1866, I: 83. Publication date: May 1866. 1872. Megadermidae Gill, Arr. Families of Mammalia, p. 17. - Comments: Originally included the subfamilies Vampyrinae Peters, 1865 [ = Family Phyllostomidae Gray, 1825], Glossophaginae Bonaparte, 1845 and Stenoderminae Gill, 1872 (see Jackson and Groves, 2015: 241). 1893. Megadermatini Winge, Samling af Afhandlinger. E Museo Lundii, 2 (1): 24. - Comments: Proposed as tribe (?) and originally included the genera Nycteris É. Geoffroy and G. Cuvier, 1795 and Megaderma É. Geoffroy, 1810 (see Jackson and Groves, 2015: 241). TAXONOMY: Based on morphological data, the Family Megadermatidae was placed in the superfamily Rhinolophoidea with the families Nycteridae, Rhinolophidae and Hipposideridae (Simmons, 1998; Simmons and Geisler, 1998), but more recent molecular studies have contradicted many groupings based on morphological data. Hand (1985, 1996) and Griffiths et al. (1992) have provided alternative phylogenies for the group. COMMON NAMES: Czech: lyronosovití, kožnatcovití, megadermovití. Dutch: Onechte vampieren. English: False Vampire Bats, Large-winged Bats. Finnish: Valevampyyrit. French: Mégadermatidés. German: Grossblattnasen, Großblattnasen, Falsche Vampire. Italian: Megadermàtidi, Fàlsi vampìri. Norwegian: Storøreflaggermus, falske vampyrer. Russian: Лжевампиры. Vietnamese: Họ dơi ma. See Handley (1980) [in Corbet and Hill (1992: 89)] for correct formation of family name. ETYMOLOGY OF COMMON NAME: The common name of the family, false vampire bats, stems from the old, erroneous belief that they fed on blood, as do the true vampire bats of the New World (Happold, 2013ay: 401). There are no blood-sucking bats in Africa. No subfamilies recognized (Simmons, 2005: 379). Currently (Simmons and Cirranello, 2020) recognized genera of the family Megadermatidae: Cardioderma Peters, 1873; Eudiscoderma Soisook, Prajakjitr, Karapan, Francis and Bates, 2015; Lavia Gray, 1838; Lyroderma Peters, 1872; Macroderma Miller, 1906; Megaderma E. Geoffroy Saint-Hilaire, 1810. Additionally, there is one extict genus: †Saharaderma Gunnell, Simons and Seiffert, 2008 PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The Megadermatidae appear in the late Eocene or early Oligocene of Europe (Butler, 1978). Butler and Hopwood (1957) reported an indetermined species of Megadermatidae from the early Miocene at Rusinga. The locality was corrected by Butler (1984: 185) to Kaswanga, where it was found in the Hiwegi Formation. 254 ISSN 1990-6471 Russell and Sigé (1970) suggest that Afropterus gigas Lavocat, 1961, is a megadermatid. DENTAL FORMULA: 0113/ 2123 = 26. Except in one non-African species, which has two upper premolars. Shi and Rabosky (2015: 1532) indicate that the Megadermatidae split of from the Rhinopomatidae some 34 million years ago. They also reported (p. 1537) the family-level stem age to be 53.4 Mya and the crown age 27.2 Mya. MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) indicate that the 2n value for the family varies between 38 and 54. Genus Cardioderma Peters, 1873 *1873. Cardioderma Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 488. Publication date: 23 June 1873. - Comments: Type species: Megaderma cor Peters, 1872. - Etymology: From the Greek "καρδία", meaning heart and "δέρμα", meaning skin, referring to the "cordiform" base of the central longitudinal crest of the nose-leaf (see Palmer, 1904: 160; Csada, 1996: 3). (Current Combination) TAXONOMY: See Simmons (2005). Currently (Simmons and Cirranello, 2020) recognized species of the genus Cardioderma: cor (Peters, 1872). Additionally, there is the extinct African †leakeyi Gunnell, Butler, Greenwood and Simmons, 2015. COMMON NAMES: Czech: srdcoví lyronosi. English: Heart-nosed Bats, African False Vampire Bats. French: Cardiodermes. German: HerznasenFledermäuse. Italian: Cardiodèrma. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Cardioderma sp. has been found in the early Pleistocene deposits in Olduvai (Butler and Greenwood, 1965; Butler, 1978). †Cardioderma leakeyi Gunnell, Butler, Greenwood and Simmons, 2015 *2015. Cardioderma leakeyi Gunnell, Butler, Greenwood and Simmons, Am. Mus. Novit., 3846: 1, 7, figs 4B-D, 5C. Publication date: 16 December 2015. Type locality: Tanzania: Arusha province: Olduvai Gorge: Bed I, FLK NI, Layer 2. [Goto Description]. - Etymology: Named in honor of Louis S.B. Leakey who was instrumental in initiating and leading the search for vertebrate fossils, especially fossil humans, in East Africa (Gunnell et al., 2015a: 7). SIMILAR SPECIES: The teeth of C. leakeyi are on average 18 - 20 % larger than those of C. cor (Gunnell et al., 2015a: 8). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pleistocene (Brown et al., 2019: Suppl.). Cardioderma cor (Peters, 1872) *1872. Megaderma cor Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 194. Type locality: Ethiopia: "Ethiopia". Holotype: ZMB 4181: ad ♂, skull and alcoholic. Collection date: undated. Vendor Gerrard; see Turni and Kock (2008: 28). - Etymology: From the neuter Latin substantive cor meaning "heart", used as an apposition, referring to the heart-shaped noseleaf (see Lanza et al., 2015: 167). ? Cardioderma cor: (Current Combination) African Chiroptera Report 2020 TAXONOMY: See Csada (1996, Mammalian Species, 519). COMMON NAMES: Chinese: 非洲假吸血蝠. Czech: lyronos srdcový, megaderma africká. Dutch: Hartneusvleermuis. English: Heart-nosed Bat, African Heart-nosed Bat, African False Vampire Bat, Heart-nosed Bigeared Bat. French: Cardioderme à nez en cœur, Pseudo-vampire à nez en cœur. German: Herznasen-Fledermaus. Italian: Cardiodèrma nàso a cuòre, Fàlso vampìro dàl nàso a cuòre. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008b; IUCN, 2004; Monadjem et al., 2017i). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017i). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008b; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004a; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Mickleburgh et al. (2008b) [in IUCN (2009)] and Monadjem et al. (2017i) report that there appear to be no major threats to this species, however, further studies are needed into the impact of disturbance on roosting sites. CONSERVATION ACTIONS: The species has been recorded from protected areas in Kenya (e.g. Tsavo West National Park (Vaughan, 1976; Aggundey and Schlitter, 1984) and is likely to be present in protected areas in some other East African countries. Additional studies are needed into the impact of disturbance on this species (Mickleburgh et al., 2008b; IUCN; Monadjem et al., 2017i). GENERAL DISTRIBUTION: Cardioderma cor is distributed from north-east Sudan (near the Red Sea) south to central Tanzania. The distribution extends from the border area of Uganda and Sudan in the west, to east Somalia (near the tip of the Horn of Africa) in the east. 255 Native: Djibouti (Pearch et al., 2001: 397); Eritrea; Ethiopia (Glass, 1965: 179); Kenya; Somalia; Sudan; Tanzania; Uganda, Zanzibar. DETAILED MORPHOLOGY: Baculum - Unknown Brain - Based on three brains from Kenya, Kruger et al. (2010a) describe the nuclear organization of the cholinergic, catecholaminergic and serotonergic systems, using immunohistochemical methods. Kruger et al., 2010b) describe the distribution of Orexin-A immunoreactive cell bodies and terminal networks based on three brains from Kenya. ECHOLOCATION: See O'Shea and Vaughan (1980) and Taylor et al. (2005). Davies et al. (2013b: Table S8) report a peak frequency of 56.7 kHz and a range between 39.6 and 90.9 Khz. Kanuch et al. (2015: 55) found that the bats emitted relative broadband FM signals with a very sort duration (2 ms). The average peak frequency was 50.4 kHz (49.2 - 51.7 kHz), and the mean inter-signal interval was 186 ms (130 - 286 ms). The signals also had a second (around 65 kHz) and third (around 88 kHz) component, which are probably not true harmonics. Smarsh and Smotherman (2016: 429) report up to four harmonics, of which the second and third are emphasized and the first suppressed. They also present the following data (p. 435): Fpeak: 49.13 ± 1.39 kHz, Fmax: 62.19 ± 2.29 kHz, Fmin: 40.14 ± 0.73 kHz, bandwidth: 22.04 ± 2.72 kHz, duration: 1.34 ± 0.06 ms, and interpulse interval: 47.53 ± 22.19 ms. Furthermore, males had significantly lower Fmin values than females. Luo et al. (2019a: Supp.) reported the following data (): Fpeak: 49.13 kHz and duration: 1.34 msec. HABITAT: Typically this species has been recorded from lowland savanna, shrubland, and the coastal strip, and in some instances, it may be observed in river valleys (Varty and Hill, 1988; Csada, 1996). Highest altitudinal record is 940 m (Csada, 1996). Stanley et al. (2007c: 58) mention that C. cor is a common resident of baobab trees (Adansonia digitata L.). During their study of the bat diversity at the Arabuko-Sokoke Forest and adjacent farmlands, Musila et al. (2019b: 8) captured 621 C. cor specimens in the farmlands and only two in the forest areas. 256 ISSN 1990-6471 HABITS: McWilliam (1987a: 243) reported that C. cor males sing to mark their feeding territories. Contrary to echolocation calls, which are produced nasally, these songs are produced orally (Smarsh and Smotherman, 2017: 2), and are seasonally. Smarsh and Smotherman (2017: 11) also found that C. cor reacted to play-back sounds by producing faster, lower-frequency songs. Smarsh and Smotherman (2016: 429) [referring to Bogdanowicz and Owen (1990)] mention that Cardioderma cor is a surface gleaner, which sits in perches and scans the environment, rotating its body and pinnae until it detects a prey to pick off the surface in the dense scrub habitat. Vaughan (1976) [in Fenton, 1986a], however, indicated that C. cor alternates between prey on the ground and flying prey, either from a perch about 1 m from the ground or from flight. ROOST: It roosts alone, or in small numbers, in caves, hollow trees (e.g. baobab) and abandoned buildings (Kingdon, 1974; Vaughan, 1976; Ryan and Tuttle, 1987; Csada, 1996; Pearch et al., 2001; Smarsh and Smotherman, 2016: 429). When roosting in trees, open sites are preferred, since this species doesn't seem to be very agile in roosting situations (Kunz, 1996: 50). In the Mago National Park (Ethiopia), Kanuch et al. (2015: 55) found that dozens of male and female bats were roosting in two underground cesspits of the toilet, which they entered through waste holes. DIET: The diet of C. cor generally consists of larger (> 25 mm) terrestrial arthropods, although small vertebrates (including frogs and bats) are eaten too (Csada, 1996: 2, 3). POPULATION: Structure and Density:- Although there is little overall information on the abundance of this species, up to 81 bats have been found roosting in a hollow baobab tree (Vaughan, 1976). Trend:- 2016: Unknown (Monadjem et al., 2017i). 2008: Unknown (Mickleburgh et al., 2008b; IUCN, 2009). 2004: Unknown (Mickleburgh et al., 2004a; IUCN, 2004). ACTIVITY AND BEHAVIOUR: Smotherman et al. (2016: 537) refer to Vaughan (1976) and McWilliam (1987a) in describing the song structure for this species as consisting of different syllable types in a basic motif. REPRODUCTION AND ONTOGENY: Tuttle and Stevenson (1982: 121) mention that both males and females need about 16 months to attain sexual maturity. Matthews (1941) and Krutzsch (2000: 95) indicate that the central duct of the ampullary gland seems to act as a temporary storage site for spermatozoa. They also mention that C. cor is the only polyoestrous species in the Megadermatidae, all others are seasonally monoestrous. Vaughan (1976) and Kunz and Hood (2000: 418) suggest that paternal recognition of young would occur in monogamous species, such as C. cor, where males participate in the care of young. The males also defend the foraging territories in which young bats learn to feed (Kunz and Hood, 2000: 440). MATING: Vaughan (1976), McWilliam (1987a), and McCracken and Wilkinson (2000: 351) report this species to be monogamous. PARASITES: PROTOZOA Kassahun et al. (2015: 168) tested one C. cor specimen from Awash-Methara (Ethiopia) and found it to be infected by Leishmania tropica Wright, 1903. ACARI Trombiculidae: Grandjeana kanuchi Kalúz and Ševcík, 2015 collected and described from C. cor in Ethiopia (Kalúz and Ševcík, 2015: 381; Stekolnikov, 2018a: 136). DIPTERA Streblidae: Raymondia huberi huberi Frauenfeld, 1855 (Shapiro et al., 2016: 254). Raymondia planiceps Jobling, 1930 (Haeselbarth et al., 1966: 102; Kanuch et al., 2015: 55; Shapiro et al., 2016: 255); Raymondia seminuda Jobling, 1954 in Ethiopia (Jobling (1954) in (Haeselbarth et al., 1966: 104). Nycteribiidae: Basilia daganiae Theodor and Moscona, 1954 from Mombasa, Kenya (Haeselbarth et al., 1966: 110). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Cardioderma cor: Adenoviruses and Astroviruses. Coronaviridae - Coronaviruses SARS-CoV - Tong et al. (2009) found coronavirus RNA in fecal samples collected of one bat collected in Kenya in 2006. African Chiroptera Report 2020 257 Tao et al. (2017: 3) indicate that one (25 %) of the Kenyan C. cor they examined tested positive for CoV. Paramyxoviridae Conrardy et al. (2014: 259) tested 14 individuals from Kenya with the use of an RT-PCR assay. Of the 10 individuals tested from Panga Yambo cave, one individual tested positive for paramyxovirus RNA that grouped phylogenetically to an Otomops martiensseni-related paramyxovirus. Polyomaviridae Cardioderma PyV KY336 - Tao et al., 2012 detected polyomavirus DNA in fecal/oral swabs from Kenya in 2006. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Egypt, Eritrea, Ethiopia, Kenya, Somalia, South Sudan, Sudan, Tanzania, Uganda. Figure 71. Distribution of Cardioderma cor Genus Lavia Gray, 1838 *1838. Lavia Gray, Mag. Zool. Bot., Edinburgh, 2 (12): 490. Publication date: 1 February 1838. - Comments: Type species: Megaderma frons E. Geoffroy Saint-Hilaire, 1810. Etymology: From "lavo" meaning fine, elegant, splendid (see Vonhof and Kalcounis, 1999: 4). Lanza et al. (2015: 172), however, indicate that lavo is a Latin verb meaning "wash, wash away, wash off, wet, moisten, bathe, soak", while its passive Latin perfect participle lauta (corresponding to a feminine verbal adjectiv) actually means - besides "washed", "clean", and "sumptuous" - "splendid" and "magnificent", referring to the beauty of the animal. (Current Combination) 1846. Livia: Agassiz, Nomenclator. Zool., Mamm., addenda 6. (Lapsus) TAXONOMY: See Simmons (2005). Currently (Simmons and Cirranello, 2020) recognized species of the genus Lavia: frons (E. Geoffroy Saint-Hilaire, 1810). COMMON NAMES: Czech: žlutokřídlí lyronosi. English: Yellowwinged Bats. French: Mégadermes à ailes orangées. German: Gelbflügel-Fledermäuse. Italian: Fàlsi vampìri dàlle àli giàlle. Lavia frons (E. Geoffroy St.-Hilaire, 1810) 1800. Vespertilio megalotis Bechstein. Type locality: South Africa: Great Namaqualand: Orange River, 50 mi [=75 km] N. - Comments: Nomen oblitum. See Grubb (2004: 91 92) for a discussion. *1810. Megaderma frons E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 15: 192. Type locality: Senegal: "Senegal". Holotype: MNHN 928:. Number 175 in Rode (1941: 236). Skull number A6844. Paratype: MNHN 929: skin only. Skull removed and lost. Number 175a in Rode (1941: 236). Paratype: MNHN 929.C: skin only. Number 175b in Rode (1941: 2436). Paratype: MNHN 929.D: skin only. Number 175c in Rode (1941: 236). Etymology: From the Latin "frons" meaning forehead (see Vonhof and Kalcounis, 1999: 4). 1905. Lavia rex Miller, Proc. Biol. Soc. Wash., 18: 227. Publication date: 9 December 1905. Type locality: Kenya: Taveta [03 23 S 37 40 E] [Goto Description]. Holotype: USNM 18993/38197: ad ♂, skin and skull. Collected by: Dr. William Louis Abbott; collection date: 1889. Skin catalogued 24 June 1890; skull catalogued 18 February 1904: see Miller (1905: 227), Lyon and Osgood (1909: 260), Poole and Schantz (1942: 118). 258 ISSN 1990-6471 1907. ? ? ? Lavia frons affinis K. Andersen and Wroughton, Ann. Mag. nat. Hist., ser. 7, 19 (110): 138, 140. Publication date: 1 February 1907. Type locality: Sudan: White Nile: Kaka [10 41 N 32 13 E] [Goto Description]. Holotype: BMNH 1901.8.8.3: ad ♂, skin only. Collected by: R. McDonald Hawker Esq. - Comments: Not mentioned by Vonhof and Kalcounis (1999: 1). Lavia frons frons: (Name Combination) Lavia frons rex: (Name Combination) Lavia frons: (Name Combination, Current Combination) TAXONOMY: See Vonhof and Kalcounis (1999, Mammalian Species, 614). Grubb (2004: 91) calls Megaderma frons a "nomen protectum". COMMON NAMES: Azande (DRC): Ndima. Chinese: 黄 翼 蝠 . Czech: lyronos žlutokřídlý, weloblánec lupenatý, megaderma žlutokřídlá. English: Yellow-winged Bat, African Yellow-winged Bat. Finnish: Keltapoimulepakko. French: Feuille de Daubenton, Megaderme à ailes orangées, Lavie à ailes jaunes, Le Megaderme feuille, Megaderme à ailes orangées. German: Gelbflügelige Großblattnase, Gelbflügel-Fledermaus. Italian: Fàlso vampìre dàlle ali giàlle. Kiluba (DRC): Kasusu. Kinande (DRC): Kakoro kombe. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008n; IUCN, 2009; Monadjem et al., 2017j). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017j). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008n; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004y; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: While Vonhof and Kalcounis (1999) state that the effects of human activity on population dynamics of this bat are unknown, there do not appear to be any major threats to the species (Mickleburgh et al., 2008n; IUCN, 2009; Monadjem et al., 2017j). CONSERVATION ACTIONS: Mickleburgh et al. (2008n) [in IUCN (2009)] and Monadjem et al. (2017j) report that it is present in a number of protected areas (e.g. Garamba National Park, Democratic Republic of the Congo). Other than studies into the possible effects of human activity on population dynamics of this bat, and additional research into the range of this species, no conservation measures are currently needed for this widespread species. GENERAL DISTRIBUTION: Lavia frons is broadly distributed south of the Sahara from Gambia and Senegal, through much of West and Central Africa, to southern and central Sudan, Eritrea, Ethiopia and Somalia in the east; ranging southwards as far as northern Zambia and northern Malawi. It is typically a lowland species found below 2,000 m asl. Native: Benin (Capo-Chichi et al., 2004: 162); Burkina Faso (Kangoyé et al., 2015a: 610); Burundi; Cameroon; Central African Republic; Chad; Congo (The Democratic Republic of the) (Schouteden, 1944; Hayman et al., 1966; Monadjem et al., 2010d: 539); Côte d'Ivoire; Eritrea; Ethiopia (Yalden et al., 1996; Lavrenchenko et al., 2004b: 132); Gabon; Gambia (Grubb et al., 1998); Ghana; Guinea; GuineaBissau (Seabra, 1900a; Monard, 1939; Rainho and Ranco, 2001: 46); Kenya (Aggundey and Schlitter, 1984); Malawi (Happold et al., 1988; Ansell and Dowsett, 1988; Vonhof and Kalcounis, 1999; Monadjem et al., 2010d: 539); Mali; Niger; Nigeria; Rwanda (Baeten et al., 1984; Senegal; Sierra Leone; Somalia (Funaioli and Simonetta, 1966); Sudan; Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 61); Zambia (Ansell, 1967; Ansell, 1978; Monadjem et al., 2010d: 539). Presence uncertain: Namibia, Zanzibar. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data: Fa: 59.5 mm, mass: 23 g, wing loading: 7.8 N/m 2, aspect ratio: 5.2. DENTAL FORMULA: Dorst (1953a: 83 - 84) indicates that he was unable to find any trace of a milk dentition in a foetus of about 35 mm in size (Fa: 25 mm), but also that the permanent teeth were already perfectly formed, but were not yet grown above the gum. African Chiroptera Report 2020 SEXUAL DIMORPHISM: Krutzsch (2000: 115) mentions the presence of dorsal skin glands in males. Kangoyé et al. (2015a: 610) found no differences in body and cranial measurements between the two sexes. ECHOLOCATION: See Taylor et al. (2005). Davies et al. (2013b: Table S8) report a peak frequency of 18.5 kHz and a range between 16.9 and 82.9 kHz. Smarsh and Smotherman (2016: 429) report up to four harmonics of which the second and third are emphasized and the first suppressed, with males having a significantly lower Fmin. They also provide the following data: Fpeak: 42.21 ± 2.35 kHz, Fmax: 48.50 ± 2.08 kHz, F min: 35.08 ± 1.25 kHz, bandwidth: 13.42 ± 3.04 kHz, duration: 3.25 ± 2.98 ms, and interpulse interval: 53.99 ± 32.24 ms. Luo et al. (2019a: Supp.) reported the following data (): Fpeak: 42.21 kHz and duration: 3.25 msec. HABITAT: R.B. Woosnam [in Thomas, 1910b: 488] mentions it to be plentiful among the acacia trees on the plains around the south end of Ruwenzori at 3,400 ft (1,000 m), where it was found hanging fully exposed to the sun. Conenna et al. (2019: 1) radio-tracked 22 bats during April - May 2017 (rainy season, n = 13) and January - February 2018 (dry season, n = 9) in a desert in northern Kenya. The bat's home ranges averaged 5.46 ± 11.04 (0.45 - 42.32) ha and they travelled a minimum distance of 99.69 ± 123.42 m/hour. These ranges were larger during the dry season than during the wet season. HABITS: Vaughan and Vaughan (1987) [in McCracken and Wilkinson (2000: 353)] found that the single young remains closely associated with both parents at roost sites and on the foraging area for up to 50 days after it becomes volant. Smarsh and Smotherman (2016: 430) consider L. frons to be an aerial-hawker, capturing insects in vegetation gaps. ROOST: L. frons roosts in trees, but prefers relatively open sites as it seems to be less agile in roosting situations (Kunz, 1996: 50). McCracken and Wilkinson (2000: 351) indicate that male/female pairs roost in the same territory, and that pair- 259 specific roost sites and foraging areas remain stable for at least several months. Smarsh and Smotherman (2016: 429) indicate that L. frons roost in male-female pairs in Acacia trees within a foraging territory. PREDATORS: Schätti (1984: 339) refers to Loveridge (in Allen, 1940), who found a specimen in the stomach of a Black Mamba (Dendroaspis polylepis). Mikula et al. (2016: Supplemental data) mention the Grey kestrel (Falco ardosiaceus Vieillot, 1823), Common kestrel (Falco tinnunculus Linnaeus, 1758) and Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as avian diurnal predators. POPULATION: Structure and Density:- Although there is little information on the abundance of this species, it is probably uncommon (Vonhof and Kalcounis, 1999). Trend:- 2016: Stable (Monadjem et al., 2017j). 2008: Stable (Mickleburgh et al., 2008n; IUCN, 2009). ACTIVITY AND BEHAVIOUR: Smotherman et al. (2016: 537) refer to Wickler and Uhrig (1969) and Vaughan and Vaughan (1986) in describing the song structure for this species as consisting of simple syllable types in a basic motif. REPRODUCTION AND ONTOGENY: Vaughan and Vaughan (1987: 217) report that females give birth to one young, which hangs on to its mother for about a week, and then starts flying. The young is weaned about 20 days after its first flight, but remains at least 50 days in close contact with its parents, during which time it learns to become a skilled opportunistic forager. In Malawi, births were reported at the end of the dry season (October) (Happold and Happold, 1990b: 566). MATING: Duron et al. (2014: 351) reports that L. frons is a monogamous species. PARASITES: SIPHONAPTERA Pulicidae: Echidnophaga aethiops Jordan and Rothschild, 1906 locality not mentioned (Haeselbarth et al., 1966: 135). Ischnopsyllidae: Chiropteropsylla brockmani Rothschild, 1915 widespread (Haeselbarth et al., 1966: 192). 260 ISSN 1990-6471 ANTHROPOPHILOUS: Jeffreys (1944: 73) [referring to Schweinfurt (1878)] mentions the following "Among the Bongo of the Anglo-Egyptian Sudan much the same superstition lurks - 'Spirits, devils and witches have their general appellation of 'Bitaboh', wood-goblins being specially called 'Ronga'. Comprehended under the same term are all the bats (especially the Megaderma frons, which flutters about from tree to tree in broad daylight).'" DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Congo (Democratic Republic of the), Côte d'Ivoire, Egypt, Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Malawi, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia. Figure 72. Distribution of Lavia frons Genus Megaderma E. Geoffroy St.-Hilaire, 1810 *1810. Megaderma E. Geoffroy St.-Hillaire, Ann. Mus. Hist. nat. Paris, 15: 197. - Comments: Type species: Vespertilio spasma Linnaeus, 1758. - Etymology: From the Greek "μέγας", meaning great or large and "δέρμα", meaning skin, referring to the large wings and interfemoral membrane (see Palmer, 1904: 403). (Current Combination) 1847. Eucheira Hodgson. - Comments: not Eucheira Westwood, 1838, an insect. 1866. Spasma Gray. 1872. Lyroderma Peters. 1961. Afropterus Lavocat, Notes Mém. Serv. Mines Carte geol. Maroc, 155: ?. - Comments: Mid Miocene-Pliocene in Morocco. TAXONOMY: Includes Lyroderma, but see Hand (1985). Two subgenera (Megaderma and Lyroderma) are recognized by Corbet and Hill (1992) and Simmons (2005: 380). Currently (Simmons and Cirranello, 2020) recognized species of the genus Megaderma: spasma (Linnaeus, 1758) – Sri Lanka and India through south east Asia (including Vietnam) to Lesser Sundas, the Philippines and Molucca Isls, various adjacent islands (Simmons, 2005: 380). Additionally there are the extinct †gigas Lavocat, 1961; †jaegeri Sigé, 1976. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: gigas (Lavocat, 1961) fossil bat from the Pliocene. †Megaderma gaillardi (Trouessart, 1898) *1898. Miomegaderma gaillardi Trouessart, Catalogus Mammalium tam viventium quam fossilium. Berlin., 84. Type locality: France: La Grive St. Alban. 1961. Afropterus gigas Lavocat, Notes Mém. Serv. Mines Carte geol. Maroc, xxx.. Type locality: Morocco: Beni Mellal. 2013. Megaderma gaillardia: Stoetzel, Palaeog., Palaeoclim., Palaeoecol., 392: 363. (Lapsus) ? Megaderma gaillardi: PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Middle Miocene (Astaracanian / Langhian, Serravallian - Brown et al., 2019: Suppl.). GENERAL DISTRIBUTION: Europe and Morocco Hugueney et al. (2015: 472). African Chiroptera Report 2020 261 †Megaderma gigas (Lavocat, 1961) *1961. Afropterus gigas Lavocat, Notes Mém. Serv. Mines Carte geol. Maroc, 155: ?. Type locality: Morocco: Beni Mellal. - Comments: Mio-Pliocene. ? Megaderma gigas: (Name Combination, Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pliocene. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ghana. †Megaderma jaegeri Sigé, 1976 *1976. Megaderma jaegeri Sigé, Géol. Médit., 3 (2): xxx. (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Middle Miocene (Langhian, Serravallian - Brown et al., 2019: Suppl.). Type locality: Morocco: Béni Mellal. GENERAL DISTRIBUTION: Morocco (Hugueney et al. (2015: 472). †Genus Saharaderma Gunnell, Simons and Seiffert, 2008 *2008. Saharaderma Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 3, 7. - Comments: Type species - Saharaderma pseudovampyrus. - Etymology: For the 'Sahara Desert' in combination with derma, Greek for skin, and the common ending for several genera of megadermatids (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). Currently recognized species of the genus Saharaderma: †pseudovampyrus Gunnel, Simons and Seiffert, 2008. GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). SIMILAR SPECIES: See Gunnell et al. (2008). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). †Saharaderma pseudovampyrus Gunnell, Simons and Seiffert, 2008 *2008. Saharaderma pseudovampyrus Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 3, 7, fig. 7. Type locality: Egypt: Fayum Depression: Quarry L-41: Lower Sequence, Jebel Qatrani Formation. Holotype: CGM 83672:. Right dentary p4-m3: see Gunnell et al. (2008: 7). - Comments: latest Eocene, Priabonian. - Etymology: 'Pseudo- and Vampyrus' in reference to the common name of megadermatids (False Vampire Bats), from Slavic, 'wampir', blood-sucking ghost (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). SIMILAR SPECIES: See Gunnell et al. (2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). Timeframe: Latest Eocene (Priabonian - Brown et al., 2019: Suppl.), Lower Sequence, Jebel Qatrani formation. GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). 262 ISSN 1990-6471 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). Family RHINOLOPHIDAE Gray, 1825 *1825. Rhinolophidae Gray, Zool. Journ., 2 (6): 242. - Comments: Type genus: Rhinolophus Lacépède, 1799. Corbet and Hill (1992: 90) mention "Bell, 1836: 599" as author. Jackson and Groves (2015: 242) mention "T. Bell, 1836: 599" as author of this spelling, but attributed the family name to Gray, 1825. (Current Combination) 1825. Rhinolophina Gray, Ann. Philos., 10 (5): 338. - Comments: Proposed as tribe and originally containing the genera Megaderma É. Geoffroy, 1810; Rhinolophus Lacépède, 1799; Nycteris É. Geoffroy & G. Cuvier, 1795; Mormoops Leach, 1821 and Nyctophilus Leach, 1821. Jackson and Groves (2015: 241, 242) placed this name in the synonymy of both Rhinolophoidea and Rhinolophidae. 1827. Rhinolophina Lesson, Mannuel de mammalogie, p. 81. - Comments: In Part (see Taylor, 1934). 1854. Rhinolophides Gervais, Histoire naturelle des mammifères, p. 200. - Comments: In Part (see Taylor, 1934). 1855. Histiorhina Van der Hoeven, Handboek Dierkunde, 2nd ed., 2: 1033. - Comments: Proposed as tribe (?), which was not based on a genus name (see Jackson and Groves, 2015: 242). 1865. Rhinolophi Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, p. 256. - Comments: In Part (see Taylor, 1934). Originally included the genera Rhinolophus Lacépède, 1799; Phyllorrhina Bonaparte, 1837 [1832–1841] [ = Hipposideros J. Gray, 1831b]; and Coelops Blyth, 1848 (see Jackson and Groves, 2015: 242). 1883. Phyllorhinidae de Rochebrune, Act. Soc. Linn. Bordeaux, 37 (4) 7: 94. - Comments: Originally included the genus Phyllorhina Leach, 1816 [= Rhinolophus Lacépède, 1799]. Howev,er the name is unavailable (see Jackson and Groves, 2015: 242). 1893. Rhinolophini Winge, Samling af Afhandlinger. E Museo Lundii, 2 (1): 24. - Comments: Proposed as tribe (?) and originally included the genera Phyllorhina [sic] Bonaparte, 1837 [1832–1841] [= Hipposideros J. Gray, 1831]; Anthops Thomas, 1888: Rhinonicteris J. Gray, 1847; Triaenops Dobson, 1871; Coelops Blyth, 1848; and Rhinolophus Lacépède, 1799 (ee Jackson and Groves, 2015: 242). TAXONOMY: Koopman and Jones, 1970; Swanepoel et al. (1980: 157), Harrison and Bates (1991: 51), Corbet and Hill (1992: 104 - with comment), Pavlinov et al. (1995), Peterson et al. (1995: 66), Hutson et al. (2001: 16) and Simmons (2005: 350) included the Hipposideridae. Hill (1982b) listed Hipposideridae as a distinct family, without comment and Hill and Smith (1984) retained this separation and discussed the question. English: Horseshoe bats. Finnish: Hevosenkenkäyököt, Kaviokuonoyököt. French: Rhinolophidés, Fers-à-cheval, Rhinolophes. German: Hufeisennasen, HufeisennasenFledermäuse. Italian: Rinolòfidi. Norwegian: Hesteskoneser, Ekte hesteskonese. Polish: podkowcowate. Romanian: Lilieci cu pliuri nazale. Russian: Подковоносые. Ukrainian: Підковикові [= Pidkovykovi]. Vietnamese: Họ dơi lá. Foley et al. (2014: 319) state that the Rhinolophidae form a fully supported monophyletic group, but that the intrafamilial relationships are less well resolved than for the Hipposideridae. ETYMOLOGY OF COMMON NAME: They are called horseshoe bats because the anterior nose-leaf is shaped like a horseshoe. Currently (Simmons and Cirranello, 2020) recognized genera of the family Rhinolophidae: Rhinolophus Lacépède, 1799. COMMON NAMES: Czech: vrápencovití, vlastní vrápenci. Dutch: Hoefijzerneusvleermuizen, Hoefijzerneuzen. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The Rhinolophidae, date from the late Eocene or early Oligocene of Europe and have been recorded in Africa only from the Mio-Pliocene of Morocco (Lavocat, 1961) and the late Pliocene of Makapansgat (De Graaf, 1960). The family-level stem and crown ages were calculated as 49.9 and 49.8 Mya by Shi and Rabosky (2015: 1537). African Chiroptera Report 2020 263 BIOGEOGRAPHY: During the Miocene at about 17 Ma, the Rhinolophus diversified into distinct clades (Foley et al., 2014: 320). MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) indicated that the karyotype for all the members of this family has a 2n value varying between 28 and 62. FUNCTIONAL MORPHOLOGY: Using micro computed tomography (micro-CT) scans, Curtis A. A. and Simmons N. B. (2016: 310) examined the turbinal morphology in the Rhinolophidae and found strangely formed maxilloturbinal bones within the respiratory airway. These looked distinct in form from the scrolled, branched, or plate shaped turbinals reported in other mammals, including other nasophonating bats. VIRUSES: Coronaviridae SARS-related coronaviruses (subgenus Sarbecovirus; see Markotter et al., 2020: 5) have only been detected in the Rhinolophus genus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Tanzania. Genus Rhinolophus Lacépède, 1799 *1799. Rhinolophus Lacépède, Tabl. Div. Subd. Orders Genres Mammifères, 15. - Comments: Type species (by monotypy): Vespertilio ferrum-equinum Schreber, 1774. Conserved by Opinion 91 (1926) and Direction 24 (1955) (see Corbet and Hill, 1992: 91). - Etymology: From the Greek "ρις" or "ρινός", meaning nose and "λόφος", meaning crest, referring to the complex nose-leaf, consisting of three parts (Palmer, 1904: 606; Taylor, 2005). Lanza et al. (2015: 76) translate "λόφος" as "comb", which leads to "with a comb on the nose or, more properly, on the snout". (Current Combination) 1816. Phyllorhina Leach, Systematic Catalogue of the species of indigenious Mammals in the British Museum, 5. - Comments: Type species: Vespertilio minutus Montagu, 1808. By subsequent designation (see Jackson and Groves, 2015: 243). Not Phyllorrhina Bonaparte, 1831 (Mammalia, Chiroptera, Rhinolophidae), a Nomen nudum according to Palmer (1904: 535), nor Phyllorrhina Bonaparte, 1837 [1832–1841] (Mammalia, Chiroptera, Hipposideridae) (see Jackson and Groves, 2015: 243). 1836. Rhinocrepis Gervais, Dictionnaire Pittoresque d'Histoire naturelle, 4 (2): pt. 318, p. 617. Comments: Type species: Vespertilio ferrum-equinum Schreber, 1774. By "implication" according to Allen (1939a: 72) or by subsequent designation (Palmer, 1904: 606). 1847. Aquias Gray, Proc. zool. Soc. Lond., 1847, XV (clxix): 15. Publication date: 13 April 1847. - Comments: Type species: Rhinolophus luctus Temminck, 1838 / Rhinolophus trifoliatus Temminck, 1834. Restricted to R. luctus by Ellerman et al. (1953): see Meester et al. (1986: 36). 1866. Phyllotis Gray, Proc. zool. Soc. Lond., 1866, I (vi): 81. Publication date: May 1866. Comments: Type species (by monotypy): Rhinolophus philippinensis Waterhouse, 1843. Preoccupied by Phyllotis Waterhouse 1837 (Mammalia, Rodentia, Cricetidae) (see Jackson and Groves, 2015: 243). 1867. Cœloephyllus Peters, Proc. zool. Soc. Lond., 1866, III: 427. Publication date: April 1867. - Comments: Type species: Rhinolophus cœlophyllus Peters, 1867. 1867. Coelophyllus Peters, Proc. zool. Soc. Lond., 1866, III: 427. Publication date: April 1867. - Comments: Type species (by monotypy): Rhinolophus coelophyllus Peters, 1867 (see Jackson and Groves, 2015: 243). 1901. Euryalus Matschie, Sber. Ges. naturf. Freunde Berlin, 225. - Comments: Type species (by monotypy): Rhinolophus mehelyi Matschie, 1901. Described as a subgenus of Rhinolophus (see Meester et al., 1986: 36; Jackson and Groves, 2015: 243). - Etymology: From the specific name euryale ("Ευρυάλη"), being one of the Gorgons (see Palmer, 1904: 279). 1941. Rhinophyllotis Troughton, Furred Animals of Australia, 1st edn: 342. - Comments: Type species (by monotypy): Rhinolophus megaphyllus Gray, 1834. Redescription for Rhinophyllotus Iredale and Troughton, 1934, "nomen nudum" (see Meester et al., 1986: 36). 1951. Rhinomegalophus Bourret, Bull. Mus. natn. Hist. nat., Paris, sér. 2, 23 (6): 607. Comments: Type species (by monotypy): Rhinomegalophus paradoxolophus Bourret, 1951 (see Jackson and Groves, 2015: 243). 264 ISSN 1990-6471 1973. 1993. 2003. 2015. 2015. 2018. ? Rhinomegaphyllus: Thonglongya, Mammalia, 37: 587. - Comments: Lapsus for Rhinomegalophus Bourret, 1951 (see Eger and Fenton, 2003: 1). (Lapsus) Rhinilophus: Amr and Qumsiyeh, Ent. News, 104 (1): 44. (Lapsus) Indorhinolophus Guillén-Servent, Francis and Ricklefs, in: Csorba, Ujhelyi and Thomas, Horseshoe Bats of the World. Rhrynolophus: Thies and Lewis, Occ. Pap. Mus. Texas Tech Univ., 330: 7. Publication date: 10 February 2015. (Lapsus) Rhynolophus: Thies and Lewis, Occ. Pap. Mus. Texas Tech Univ., 330: 7. Publication date: 10 February 2015. (Lapsus) Rhinolphus: Leopardi, Holmes, Gastaldelli, Tassoni, Priori, Scaravelli, Zamperin and De Benedictis, Infec. Gen. Evol., 58: 283. Publication date: 30 January. (Lapsus) Rhinolophus sp.: TAXONOMY: The radiation (about 15 million years ago) of Rhinolophidae into five main clades apparently occurred rapidly and the distinct clades could be regarded as different genera (Servent et al., 2003: xvi). Following tradition Servent et al. (2003: xvi) split the six subgenera (Aquias, Phyllorhina, Rhinolophus, Indorhinolophus, Coelophyllus, Rhinophyllotis) into groups corresponding to smaller lineages in the phylogeny. All African and Palaearctic species are contained within a single clade (Servent et al., 2003: xvii). This confirms the views proposed by Bogdanowicz (1992) and Bogdanowicz and Owen (1992), who suspected that the relationship was monophyletic. Qumsiyeh et al. (1988) studied protein electrophoresis of a limited number of African taxa, which grouped together in their phylogeny. All African and Palaeartic Rhinolophid species karyotyped show a diploid number of 58, which differs from all other Rhinolophus with the exception of some populations of R. hipposideros (Zima et al., 1992b). Servent et al. (2003: xvii) note that the geographical distribution of taxa and the phylogeny are strongly congruent. Species groups recognized by Servent et al. (2003: xvi), for African taxa: Rhinolophus: R. landeri - landeri, alcyone, ?guineensis. R. euryale - blasii, euryale, mehelyi. R. capensis - ?capensis, ?swinnyi, ?denti, simulator. R. adami - ?adami, ?maendeleo. R. ferrumequinum ferrumequinum, clivosus, ?silvestris, ?deckenii. R. maclaudi - ?maclaudi, ruwenzorii. R. fumigatus - fumigatus, eloquens, hildebrandti, darlingi. Stoffberg et al. (2010a: 6 - 7) refer to GuillénServent et al. (2003), who, based on the complete cytochrome b gene, propose that Rhinolophus should be divided into six subgenera: Aquias (trifoliatus group), Coelophyllus (affinis, coelophyllus, euryotis, and pearsonii groups), Indorhinolophus (rouxi group), Phyllorhina (hipposideros group), Rhinolophus (adami, capensis, euryale, ferrumequinum, fumigatus, landeri, and maclaudi groups), and Rhinophyllotis (acuminatus, borneensis, macrotis, malayanus, megaphyllus, and pusillus groups), and confirm most of these. See Csorba et al. (2003: 1 - 3) for key to species. See Taylor et al. (2019a: 316) for a discussion on the recent increase in Afro-Madagascan species. Nesi et al. (2015: 1) indicate that the taxonomy and true number of species of African Rhinolophus remains unresolved because of the highly convergent morphology observed across taxa, and that the current species delimitation is based on slight variations in morphological measurements and echolocation call frequency. Due to the extensive phenotypic variation, these characters are not always very clearly separable and as such the true diversity of the the genus is most probably underestimated, based on these methods alone. Using mitochondrial cytochrome-b and four nuclear introns, Demos et al. (2019a) found indications that twelve undescribed cryptic species might be present in Africa, and that two to five of the currently accepted species are invalid. They distinguish the following putative species: alcyone, blasii 1, blasii 2, capensis, clivosus 1, clivosus 2, clivosus 3, clivosus 4, clivosus 5, clivosus 6, cohenae, damarensis, darlingi, deckenii, denti, cf. denti, cf. denti/simulator, euryale, ferrumequinum 1, fumigatus/eloquens 1, fumigatus/eloquens 2, fumigatus/eloquens 3, fumigatus/eloquens 4, fumigatus/eloquens 5, fumigatus/eloquens 6, fumigatus/eloquens 7, fumigatus/eloquens 8, gorongosae, hildebrandtii 1, hildebrandtii 2, horaceki, kahuzi, landeri, cf. landeri, lobatus, mabuensis, mehelyi, mossambicus, rhodesiae, ruwenzorii, simulator 1, simulator 2, smithersi, swinnyi, willardi. Currently (Simmons and Cirranello, 2020) species of the genus Rhinolophus: acuminatus Peters, 1871 – Thailand, Laos, Cambodia, Peninsular African Chiroptera Report 2020 Malaysia and Sabah, Borneo, Sumatra (including Nias and Engano Isls), Java, Karkatau, Lombok, and Bali (Indonesia), Palawan, Balabac, Busuanga (Philippines) (Csorba et al., 2003: 98; Simmons, 2005: 350); adami Aellen and Brosset, 1968; affinis Horsfield, 1823 – India and Nepal to south China and Vietnam, through Malaysia to Borneo and Lesser Sunda Isls, Andaman Isls (India) (Csorba et al., 2003: 68; Simmons, 2005: 351); alcyone Temminck, 1853; arcuatus Peters, 1871 – Sumatra to Philippines, New Guinea and South Molucca Isls (Csorba et al., 2003: 20; Simmons, 2005: 351); beddomei K. Andersen, 1905 – southern India, Sri Lanka (Csorba et al., 2003: 123; Simmons, 2005: 351); belligerator Patrick, McCulloch, and Ruedas, 2013 – Indonesia (Sulawesi); blasii Peters, 1867; bocharicus Kastchenko and Akimov, 1917 – Kyrgyzstan, western Tajikistan, northeastern Iran, Uzbekistan, Turkmenistan, Afghanistan (Csorba et al., 2003: 35; Simmons, 2005: 352); borneensis Peters, 1861 – Borneo, Labuan and Banguey Isls (Malaysia), Java, Karimata Isls and South Natuna Isls (Indonesia), Cambodia, Laos and Vietnam (Csorba et al., 2003: 70; Simmons, 2005: 352); canuti Thomas and Wroughton, 1909 – Java, Bali, Timor (Indonesia) (Csorba et al., 2003: 22; Simmons, 2005: 352); capensis Lichtenstein, 1823; celebensis K. Andersen, 1905 – Java, Bali, Timor, Sulawesi, Sangihe, Kangean and Talaud Isls (Indonesia) (Csorba et al., 2003: 72; Simmons, 2005: 352); chiewkweeae Yoshiyuki and Lim, 2005 – Malaysia; clivosus Cretzschmar, 1828; coelophyllus Peters, 1867 – western Malaysia, Thailand, Burma, Laos (Csorba et al., 2003: 24; Simmons, 2005: 353); cognatus K. Andersen, 1906 – Andaman Isls (India) (Csorba et al., 2003: 99; Simmons, 2005: 353); cohenae Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012; convexus Csorba, 1997 – Peninsular Malaysia, Laos (Csorba et al., 2003: 100; Simmons, 2005: 353); cornutus Temminck, 1834 – Japan (Csorba et al., 2003: 102; Simmons, 2005: 353); creaghi Thomas, 1896 – Borneo, Madura Isl, Kalimantan (Indonesia), Sabah, Sarawak (Malaysia) (Csorba et al., 2003: 25; Simmons, 2005: 354); damarensis Roberts, 1946; darlingi K. Andersen, 1905; deckenii Peters, 1868; denti Thomas, 1904; eloquens K. Andersen, 1905; episcopus G.M. Allen, 1923 – China; euryale Blasius, 1853; euryotis Temminck, 1835 – Aru Isls, Buru, Bacan, Amboina, Seram and Tanimbar Isls, Kai Isls, Halmahera and Sulawesi (Indonesia), New Guinea, Bismarch Arch, adjacent small islands (Csorba et al., 2003: 27; Simmons, 2005: 355); ferrumequinum (Schreber, 1774); formosae Sanborn, 1939 – Taiwan (Csorba et al., 2003: 124; Simmons, 2005: 355); francisi Soisook, Struebig, Bates and Miguez, 2015 – Known from 6 specimens only: two 265 in Sabah, three in Indonesian Borneo, and one in Thailand; fumigatus Rüppell, 1842; gorongosae Taylor, MacDonald, Goodman, Kearney, Cotterill, Stoffberg, Monadjem, Schoeman, Guyton, Naskrecki and Richards, 2018; guineensis Eisentraut, 1960; hildebrandtii Peters, 1878; hilli Aellen, 1973; hillorum Koopman, 1989; hipposideros (Borkhausen, 1797); hirsutus K. Andersen, 1905 – Philippines; horaceki Benda and Vallo, 2012; imaizumii Hill and Yoshiyuki, 1980 – Iriomote Isl and Yaeyama Isl (Japan: Ryukyu Isls) (Csorba et al., 2003: 104; Simmons, 2005: 357); inops K. Andersen, 1905 – Philippines except Palawan region (Csorba et al., 2003: 29; Simmons, 2005: 357); kahuzi Fahr and Kerbis Peterhans, 2013; keyensis Peters, 1871 – Many islands in Indonesia (Simmons, 2005: 357); landeri Martin, 1838; lanosus K. Andersen, 1905 – China; lepidus Blyth, 1844 – Afghanistan, Pakistan, northern India, Nepal, Burma, Thailand, Szechwan and Yunnan (China), Peninsular Malaysia, Sumatra (Indonesia) (Csorba et al., 2003: 106; Simmons, 2005: 358); lobatus Peters, 1852; luctoides Volleth, Loidl, Mayer, Yong, Müller and Heller, 2015 – Peninsular Malaysia; luctus Temmink, 1834 – India, Nepal, Burma, Sri Lanka, south China, Vietnam, Cambodia, Laos, Thailand, Peninsular Malaysia, Borneo, Sumatra, Java and Bali (Indonesia) (Csorba et al., 2003: 126; Simmons, 2005: 358); mabuensis Taylor, Stoffberg, Monadjem, Scoeman, Bayliss and Cotterill, 2012; maclaudi Pousargues, 1898; macrotis Blyth, 1844 – Pakistan, northern India, Nepal to south China, Burma, Thailand, Laos, Vietnam and Peninsular Malaysia, Sumatra (Indonesia), Philippines (Csorba et al., 2003: 87; Simmons, 2005: 358); madurensis K. Andersen, 1918 – Madura and Kangean Isls (Indonesia) (Simmons, 2005: 358); maendeleo Kock, Csorba and Howell, 2000; malayanus Bonhote, 1903 – Thailand, Burma, Cambodia, Laos, Vietnam, Peninsular Malaysia (Csorba et al., 2003: 74; Simmons, 2005: 359); marshalli Thonglongya, 1973 – Thailand, Burma, Vietnam, Laos, Peninsular Malaysia (Csorba et al., 2003: 89; Simmons, 2005: 359); mcintyrei Hill and Schlitter, 1982 – New Guinea; megaphyllus Gray, 1834 – eastern New Guinea; Misima Isl (Louisiade Arch.), Goodenough Isl (D’Entrecasteaux Isls) and Bismarck Arch. (Papua New Guinea), Moluccas, Lesser Sundas, eastern Queensland, eastern New South Wales and eastern Victoria (Australia) (Csorba et al., 2003: 76; Simmons, 2005: 359); mehelyi Matschie, 1901; microglobosus Csorba and Jenkins, 1998 – Cambodja, Laos, Thailand (N of Kra isthmus), N + C Vietnam, China (Yunnan); mitratus Blyth, 1844 – India (Csorba et al., 2003: 131; Simmons, 2005: 360); monoceros K. Andersen, 1905 – Taiwan (Csorba et al., 2003: 108; Simmons, 2005: 360); montanus Goodwin, 266 ISSN 1990-6471 1979 – Timor (Indonesia) (Csorba et al., 2003: 91; Simmons, 2005: 360); morio Gray, 1842 – Sumatra (part); Peninsular Malaysia; mossambicus Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012; nereis K. Andersen, 1905 – Anamba and North Natuna Isls (Indonesia) (Csorba et al., 2003: 78; Simmons, 2005: 360); osgoodi Sanborn, 1939 – Yunnan (China) (Csorba et al., 2003: 109; Simmons, 2005: 360); pearsoni Horsfield, 1851 – northern India, Nepal, Bhutan, Burma, Tibet, Szechwan, Anhwei and Fukien (China) to Vietnam, Laos, Thailand, Peninsular Malaysia (Csorba et al., 2003: 83; Simmons, 2005: 360); perditus K. Andersen, 1918 – Ryukyu Islands, Ishigaki Island (Japan); philippinensis G.R. Waterhouse, 1843 – Phillipines, Kai Isls, Sabah, Sarawak and Sulawesi (Indonesia), Borneo, New Guinea, northeastern Queensland (Australia) (Csorba et al., 2003: 94; Simmons, 2005: 361); proconsulis Hill, 1959 – Borneo; pusillus Temminck, 1834 – India, Nepal, Thailand, Burma, Laos, south China, Peninsular Malaysia, Mentawai Isls, Java and Lesser Sunda Isls (Indonesia), small adjacent islands (Csorba et al., 2003: 112; Simmons, 2005: 361); refulgens K. Andersen, 1905 – Malaysia; rex G. M. Allen, 1923 – southwestern China (Csorba et al., 2003: 96; Simmons, 2005: 361); rhodesiae Roberts, 1946; robinsoni K. Andersen, 1918 – western Malaysia, Thailand, adjacent small islands (Simmons, 2005: 361); rouxii Temminck, 1835 – Sri Lanka, peninsular India to southern Burma and Vietnam (Csorba et al., 2003: 117; Simmons, 2005: 362); rufus Eydoux and Gervais, 1836 – Philippines (Csorba et al., 2003: 30; Simmons, 2005: 362); ruwenzorii J. Eric Hill, 1942; sakejiensis Cotterill, 2002; schnitzleri Wu and Thong, 2011 – only known from the type locality in China; sedulus K. Andersen, 1905 – Peninsular Malaysia, Sarawak and Sabah (Malaysia), Borneo (Indonesia) (Csorba et al., 2003: 128; Simmons, 2005: 362); septentrionalis Sanborn, 1939 – Yunnan province (China); shameli Tate, 1943 – Burma, Thailand, Laos, Cambodia, Peninsular Malaysia (Csorba et al., 2003: 31; Simmons, 2005: 363); shortridgei K. Andersen, 1918 – northern India, Burma (Csorba et al., 2003: 114; Simmons, 2005: 363); siamensis Gyldenstolpe, 1917 – Thailand, Laos, Vietnam (Simmons, 2005: 363); silvestris Aellen, 1959; simulator K. Andersen, 1904; sinicus K. Andersen, 1905 – southern China, Nepal, northern India, Vietnam (Csorba et al., 2003: 119; Simmons, 2005: 363); smithersi Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012; stheno K. Andersen, 1905 – Vietnam, Thailand, Laos, Peninsular Malaysia, Sumatra and Java (Indonesia) (Csorba et al., 2003: 80; Simmons, 2005: 363); subbadius Blyth, 1844 – northeastern India, Nepal, Vietnam, Burma (Csorba et al., 2003: 115; Simmons, 2005: 364); subrufus K. Andersen, 1905 – Philippines except Palawan region (Csorba et al., 2003: 33; Simmons, 2005: 364); swinnyi Gough, 1908; thailandensis Wu, Harada and Motokawa, 2009 – N Thailand (Chiang Mai); thomasi K. Andersen, 1905 – Burma, Vietnam, Thailand, Laos (Csorba et al., 2003: 121; Simmons, 2005: 364); trifoliatus Temminck, 1834 – northeastern India, southwestern Thailand and Burma, Peninsular Malaysia, Sarawak and Sabah (Malaysia), Singapore, Borneo, Sumatra, Riau Archipelago, Banguey Isl, Java, Banka Isl and Nias Isl (Indonesia) (Csorba et al., 2003: 130; Simmons, 2005: 364); virgo K. Andersen, 1905 – Philippines (Csorba et al., 2003: 81; Simmons, 2005: 364); willardi Kerbis Peterhans and Fahr, 2013 – Democratic Republic of the Congo (Albertine Rift Valley); xinanzhongguoensis Zhou, Guillén-Servent, Lim, Eger, Wang and Jiang, 2009 Yunnan Province (China); yunanensis Dobson, 1872 – Yunnan (China), Burma, Thailand, northeastern India (Csorba et al., 2003: 85; Simmons, 2005: 365); ziama Fahr, Vierhaus, Hutterer and Kock, 2002. Additionally, in Africa there is also the extinct †maghrebensis Gunnell, Eiting, and Geraads, 2011. COMMON NAMES: Czech: vrápenci, wrápenecové, vrápníkové, vrápeníkové, podkováčkové, vrápenec. Dutch: Hoefijzerneuzen. English: Horseshoe bats. French: Rhinolophes, Fers-à-cheval. German: Hufeisennasen, Hufeisennasen-Fledermäuse. Italian: Fèrri di cavàllo, Rinòlofi. Kiluba (DRC): Kasusu. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The approximate fossil date for the crown Rhinolophus is placed at about 37.2 MYA by Shi and Rabosky (2015: 1532). Three or four Rhinolophus species were reported by Avery (2007: 619) from Pleistocene deposits at Wonderwerk Cave, South Africa. Geraads et al. (2010: 279) reported the presence of Rhinolophus in Late Cenozoic (ca. 2.5 MYA) deposits in Ahl al Oughlam, Morocco. BIOGEOGRAPHY: See Servent et al. (2003: xii - xxiv) for discussion on phylogeny and biogeography of the horseshoe bats. During the Miocene at about 17 Mya, Rhinolophus diversified into three distinct clades, two of these clades showed distinct geographical origins (Foley et al., 2014: 320). In Eastern/South Eastern Asia represented by R. trifoliatus and R. luctus and East Asia represented by R. pearsoni, R. sinicus, R. creaghi, R. shameli and R. pusillus. While the African Chiroptera Report 2020 clade that contains R. hipposideros, R. ferrumequinum and R. euryale is less clear indicating a widespread geographical origin including Europe, East Asia, Middle East and Africa. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Schmieder et al. (2015) studied the wing morphology of five species of European Rhinolophid bats, which also occur in northern Africa: R. blasii, R. euryale, R. ferrumequinum, R. hipposideros, and R. mehelyi, and they provide a table of geometric differences (p. 8). DENTAL FORMULA: The dental formula is variable, usually 32. 1123/ 2133 = GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: Santana and Lofgren (2013: 2521) investigated the evolution of skull morphology and modularity in the recent radiation of rhinolophid bats and predict that nasal echolocation has resulted in the evolution of a third cranial module (the 'nasal dome'), in addition to the two cranial modules conserved across mammals. FUNCTIONAL MORPHOLOGY: Jacobs et al. (2007a) found that in Africa the peak echolocation frequency was inversely correlated with size in the Rhinolophidae, while for rhinolophids from around the world, peak frequency was inversely correlated with forearm length. Similarly, there was an inverse correlation between mass and peak frequency, between forearm and peak frequency and between wing loading and peak frequency in the African Rhinolophidae (Jacobs et al., 2007a). ECHOLOCATION: Jacobs et al. (2014: 2829) found that echolocation frequency was positively correlated with bite force, which suggests that its evolution is influenced by a trade-off between the masticatory and sensory functions of the skull. In support of this, variation in skull shape was explained by both echolocation frequency (80 %) and bite force (20 %). They also found that selection has acted on the nasal capsules, which have a frequency-specific impedance matching function during vocalization. In the Mapungubwe National Park (RSA), Parker and Bernard (2018: 56, 57) recorded a Rhinolophus species with a Fchar = 100.72 ± 0.52 kHz, Fmax: 100.93 ± 0.77 kHz, Fmin: 91.34 ± 6.08 kHz, Fknee: 100.72 ± 0.52 kHz, duration: 34.89 ± 14.42 msec, with 8.32 ± 3.39 calls/sec. This type of call was also recorded by Taylor et al. (2013b: 267 14) from the Soutpansberg area, and probably is produced by an undescribed species. PREDATORS: Mikula et al. (2016: Supplemental data) mention the following diurnal avian predators: African fish eagle (Haliaeetus vocifer (Daudin, 1800)), Bat hawk (Macheiramphus alcinus Bonaparte, 1850) [on either landeri or fumigatus or hildebrandti]. PARASITES: BACTERIA Bartonellae Kamani et al. (2014: 628) tested 12 Nigerian "Rhinolophus" sp. and found at least 8 of them testing positive for Bartonella DNA. Of the six Nigerian bats, examined by Di Cataldo et al. (2020: 2), 2 were found to be infected by hemoplasma bacteria. HAEMOSPORIDA Schaer et al. (2015: 381) found Nycteria sp. parasites in Rhinolophus sp. 1 from South Sudan. The same bat species was also infected by Nycteria cf. gabonensis. Landau et al. (2012: 142) and Perkins and Schaer (2016: Suppl.) also reported Nycteria krampitzi Rosin, Landau and Hugot, 1978 from the Republic of Congo and Gabon. Wünschmann et al. (2010: 178) refer to Boulard (1975), who reported the first case of renal coccidiosis in a bat (on H. caffer and Rhinolophus sp.) from Sierra Leone and the Congo and caused by Klossiella killicki. Garnham (1973: 236) refers to Anciaux de Faveaux for a Polychromophilus ? murinus from "Africa". HEMIPTERA Cimicidae: Haeselbarth et al. (1966: 10) records that only three males of Loxaspis setipes Ferris and Usinger, 1957 are known from unknown Rhinolophus, from Lukolela, Congo. DIPTERA Streblidae: Brachytarsina africana (Walker, 1849) is a typical ectoparasite of the genus Rhinolophus in sub Saharan Africa (Haeselbarth et al., 1966: 100). Ascodipteron brevior Maa, 1965 from Sudan (Haeselbarth et al., 1966: 106). Ascodipteron lophotes Monticelli, 1898 from Yemen (Haeselbarth et al., 1966: 106). Raymondia alulata Speiser, 1908 (Shapiro et al., 2016: 253). Raymondia aspera Maa, 1968 (Shapiro et al., 2016: 253). Raymondia boquieni Vermeil, 1965 from Congo and the DRC (Shapiro et al., 2016: 254). Raymondia huberi huberi Frauenfeld, 1855 (Shapiro et al., 2016: 254). 268 ISSN 1990-6471 Raymondia intermedia Jobling, 1936 (Shapiro et al., 2016: 255). Raymondia seminuda Jobling, 1954 (Shapiro et al., 2016: 255). Raymondia setiloba (Jobling, 1954) (Shapiro et al., 2016: 256). Raymondia waterstoni Jobling, 1931 (Shapiro et al., 2016: 256). Raymondia sp. B (Shapiro et al., 2016: 257). Nycteribiidae: Nycteribia integra Theodor and Moscona, 1954 common in Egypt (Haeselbarth et al., 1966: 109). Nycteribia ovalis Theodor, 1957 from Tanzania (Haeselbarth et al., 1966: 110). Nycteribia tecta Theodor, 1957 (Haeselbarth et al., 1966: 110). Dipseliopoda biannulate (Oldroyd, 1953) from the Sudan (Haeselbarth et al., 1966: 116). Kamani et al. (2014: 628) reported the presence of Cyclopodia greeffi Karsch, 1884 on bats from Nigeria. SIPHONAPTERA Ischnopsyllidae: Members of the genus Rhinolophopsylla Oudemans, 1909 are known to be parasites of Miniopterus and Rhinolophus (Haeselbarth et al., 1966: 191). ACARI Myobiidae: Fain (1994: 1280) indicates that only two endemic genera are found on Rhinolophus species: Neomyobia with 13 species and three subspecies, and Rhinomyobia with one species. Another genus/species - Calcarmyobia rhinolophi - was described from Rhinolophus lobatus, but this probably should be from Miniopterus natalensis arenarius (Uchikawa, 1985a: 15). VIRUSES: Adenoviridae These viruses were found by Waruhiu et al. (2017) in bats from Kenya. Coronaviridae Tao et al. (2017: 3) found 28.9 % (13 out of 45) of the Kenyan Rhinolophus sp. Specimens they tested to be positive for CoV. Tao and Tong (2019: 1) report the complete genomic sequence of a strain of Betacoronavirus (BtKY72/Rhinolophus sp./Kenya/2007 - BtKY72 subgenus Sarbecovirus). Paramyxoviridae 1 out of nine tested Rhinolophus sp. from Cameroon was found to test positive for Paramyxovirus sequences by Mortlock et al. (2015: 1841). Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 16 Kenyan Rhinolophus sp. specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Angola, Benin, Burundi, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Egypt, Equatorial Guinea, Eswatini, Ethiopia, Kenya, Malawi, Morocco, Mozambique, Namibia, Senegal, Somalia, South Africa, South Sudan, Tanzania, Togo, Tunisia, Uganda, Zambia, Zimbabwe. †Rhinolophus maghrebensis Gunnell, Eiting and Geraads, 2011 *2011. Rhinolophus maghrebensis Gunnell, Eiting and Geraads, N. Jb. Geol. Paläont. Abh., 260 (1): 56, figs 3A, 7A. Publication date: January 2011. Type locality: Morocco: Ahl al Oughlam [ca. 33 35 N 07 30 W] [Goto Description]. Holotype: INSAP AaO 4655: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Right maxilla with P4-M3. Paratype: INSAP AaO 4673: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Left dentary p4-m3. - Etymology: Name taken from Maghreb ("countries of the setting sun"), the Arabic name for north-western African countries (Morocco, Algeria, Tunisia) (see Gunnell et al., 2011: 56). (Current Combination) 2013. Rhinolophus maghrebiensis: Stoetzel, Palaeog., Palaeoclim., Palaeoecol., 392: 373. Publication date: 29 September 2013. (Lapsus) TAXONOMY: Gunnell et al. (2011: 56) describe it as a large species of the Rhinolophus ferrumequinum phenotypic group larger than extant African Rhinolophus species R. alticolus, R. blasii, R. euryale, R. denti, R. landeri, R. mehelyi, R. hipposideros, and R. simulator, smaller than R. hildebrandtii. African Chiroptera Report 2020 269 †Rhinolophus mellali Lavocat, 1961 *1961. Rhinolophus ferrumequinum mellali Lavocat, Notes Mém. Serv. Mines Carte geol. Maroc, 155: xxx. Type locality: Morocco: Beni Mellal. - Comments: The holotype consists of a left mandibular fragment with I, C, P2-M1 (Marandat, 1994: 60). ? Rhinolophus mellali: (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Early late Miocene. GENERAL DISTRIBUTION: Northern Africa (Gunnell et al., 2011: 70). Rhinolophus adami Aellen and Brosset, 1968 *1968. Rhinolophus adami Aellen and Brosset, Rev. suisse Zool., 75 (14): 443. Type locality: Congo: Kouilou [ca. 04 00 N 12 00 E] [Goto Description]. Holotype: MNHN ZM-MO1968-408: ad ♀, alcoholic (skull not removed). Collected by: Jean-Paul Adam; collection date: 9 January 1967; original number: 12/8G. 3 paratypes see Aellen and Brosset (1968: 443). Paratype: MHNG 1129.084: skull only. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. - Etymology: In honour of J.-P. Adam, the collector of the type specimen. (Current Combination) TAXONOMY: Kock et al. (2000) place it in the adami species group, followed by Csorba et al. (2003: xvi, 4 - 5) and Simmons (2005: 351). Species group reviewed by Kock et al. (2000). See Csorba et al. (2003: 5) for taxonomic background. CONSERVATION ACTIONS: Jacobs et al. (2008x) [in IUCN (2009)] report that it is not known if the species is present in any protected areas. Further studies are needed into the distribution, abundance, natural history, and threats to this species. COMMON NAMES: Czech: vrápenec Adamův. English: Adam's Horseshoe Bat, Congo Horseshoe Bat. French: Rhinolophe du Congo. German: Adams Hufeisennase. GENERAL DISTRIBUTION: Rhinolophus adami is known only from the type series collected in Kimanika cave, Kouilou, in the Republic of Congo. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Jacobs et al., 2008x; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Jacobs et al., 2008x; IUCN, 2009). 2004: DD ver 3.1 (2001) (Jacobs et al., 2004ab; IUCN, 2004). 1996: DD (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are currently unknown. It is possible that it is threatened by cave disturbance, however, this needs to be confirmed (Jacobs et al., 2008x; IUCN, 2009). Native: Congo (Aellen and Brosset, 1968; Csorba et al., 2003: 5; Simmons, 2005:351). DETAILED MORPHOLOGY: Baculum - See Csorba et al. (2003: 4). POPULATION: Structure and Density:- It is known only from ten specimens held in the Paris and Geneva museums (Aellen and Brosset, 1968), and may be naturally rare, however, this requires confirmation (Jacobs et al., 2008x; IUCN, 2009). Trend:- 2008: Unknown (Jacobs et al., 2008x; IUCN, 2009). PARASITES: Adam and Landau (1973a: 5) report on the presence of a protozoan parasite of the genus Polychromophilus (Haemoproteidae) in R. adami from the Republic of Congo. About 50 % of the specimens they examined were found to be infected. Gametocytes were found year-round. The bats were also infested (in low numbers) by the nycteribiid flies Penicillidia fulvida Bigot, 1885 270 ISSN 1990-6471 and Nycteribia schmidlii scotti Falcoz, 1923, of which the prior was a major carrier of Polychromophilus sporozoites. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo. Figure 73. Distribution of Rhinolophus adami Rhinolophus alcyone Temminck, 1853 *1853. Rhinolophus alcyone Temminck, Esquisses zoologiques sur la Côte de Guiné. 1e partie, les Mammifères., 80. Type locality: Ghana: Boutry River [=Butre river] [ca. 04 49 N 01 55 W] [Goto Description]. Holotype: RMNH MAM.35892:. Allen (1939a: 73) notes: 'Type in Leiden Mus., but description has not "one single word of any value for identifying the species or determining its affinities" - K. Andersen'. - Comments: Allen (1939a: 73), Kock (1969a: 121), Grubb et al. (1998: 77) mention 1852 as year of publication. - Etymology: alcyone: "[=Alkuone] mythological name for the kingfisher" and this is sometimes written as "Halcyon". Alkuone herself is the "daughter of Aiolos, who threw herself in the sea, where her husband Keüks (or Ceyx) was drowned. Thetis changed her in a kingfisher and her husband in a seagull". (Current Combination) TAXONOMY: Forms part of the landeri species group (Csorba et al., 2003: 55 - 57; Simmons, 2005: 351). See Csorba et al. (2003: 56) for remarks on taxonomy. COMMON NAMES: Castilian (Spain): Murciélago de Herradura. Czech: vrápenec Alcyone. English: Halcyon Horseshoe Bat. French: Rhinolophe Alcyon, Rhinolophe d'Halcyon. German: Temmincks Hufeisennase, Rothen Kamnase, Halcyon Rundblattnase. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008y; IUCN, 2009; Monadjem et al., 2017cu). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cu). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008y; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004y; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: This widespread species is threatened in parts of its range by habitat loss, largely resulting from logging and conversion of land to agricultural use. In some areas the species is hunted for food, although it is unclear whether this represents a major threat (Jacobs et al., 2008y; IUCN, 2009; Monadjem et al., 2017cu). CONSERVATION ACTIONS: Jacobs et al. (2008y) [in IUCN (2009)] and Monadjem et al. (2017cu) reported that it is not known if the species is present in any protected areas. Further studies are needed into the taxonomy and distribution of this species. African Chiroptera Report 2020 GENERAL DISTRIBUTION: Rhinolophus alcyone ranges through much of West and Central Africa. It has been recorded from Senegal in the west through to Togo, and then from Nigeria to southern Sudan and western Uganda, with patchy records from the Congo basin (although it likely occurs throughout the Congo). It ranges as far south as central Democratic Republic of the Congo. 271 fourth one was lactating. One birth was reported in September. PARASITES: HAEMOSPORIDA Schaer et al. (2013a: 17416; 2015: 381) report the presence of hemosporidian parasites of the genus Nycteria in one investigated bat from Côte d'Ivoire. In the DRC, Karadjian et al. (2016: 9835) found R. alcyone specimens infected with Nycteria gabonensis Rosin, Landay & Hugot 1978. The record of this species from Gabon needs to be reexamined as it may represent a misidentified specimen of Rhinolophus sylvestris (Dowsett et al., 1991). DIPTERA Streblidae: Raymondia allisoni Theodor, 1968 from Ghana (Shapiro et al., 2016: 253). Amori et al. (2016: 219) pointed out that a number of localities, originally situated in Togo, are currently in Ghana. Nycteribiidae: Nycteribia inopinata Theodor, 1957 from Ebolowa, Cameroon (Haeselbarth et al., 1966: 109). Native: Bioko, Burkina Faso (Kangoyé et al., 2012: 6025; 2015a: 610); Cameroon; Central African Republic (Lunde et al., 2001: 537); Congo (Dowsett et al., 1991: 259); Congo (The Democratic Republic of the) (Monadjem et al., 2010d: 555); Côte d'Ivoire; Equatorial Guinea; Gabon (?); Ghana; Guinea (Fahr and Ebigbo, 2003: 128); Guinea-Bissau (Rainho and Ranco, 2001: 47); Kenya; Liberia (Monadjem and Fahr, 2007: 50); Nigeria; Senegal; Sierra Leone; Sudan; Togo (?); Uganda. Presence uncertain: Gabon. VIRUSES: Hepadnaviridiae Hepatitis B virus (HBV) Drexler et al. (2013c: 16152) report on one R. alcyone on 16 examined specimens from Gabon that was positive for horseshoe bat HBV (HBHBV). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Young (1975: 76) indicates that two distinct colour phases can be found: rufous (orange) and grey. ECHOLOCATION: In Guinea, Fahr and Ebigbo (2003) reported the frequency of the CF-component at 67.4 kHz. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003) and Lim (2007). Karyotype - Unknown. Protein / allozyme - Unknown. POPULATION: Structure and Density:- This species is easily found in colonies of up to 20 animals (Jacobs et al., 2008y; IUCN, 2009; Monadjem et al., 2017cu). Trend:- 2016: Unknown (Monadjem et al., 2017cu). 2008: Unknown (Jacobs et al., 2008y; IUCN, 2009). REPRODUCTION AND ONTOGENY: Csorba et al. (2003: 57) refer to Kingdon (1984), who indicated that three females from western Uganda were pregnant in mid-June, whereas a Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that none of the 15 specimens they examined from Gabon tested positive for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. Rhabdoviridae Lyssavirus - Rabies related viruses 2006 in Nigeria, Dzikwi et al. (2010: 269) tested 43 brains by direct fluorescent antibody (DFA) and mouse inoculation test (MIT) which tested negative for lyssavirus antigens. Lagos bat virus (LBV) - In 2006, 2 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), none tested positive for neutralizing anitibodies (Dzikwi et al., 2010: 269). Mokola virus (MOKV) - In 2006, 2 individuals from Nigeria were serum tested by a modified rapid fluorescent focus inhibition test (RFFIT), none tested positive for neutralizing anitibodies (Dzikwi et al., 2010: 269). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Nigeria, Senegal, Sierra Leone, Uganda. 272 ISSN 1990-6471 Figure 74. Distribution of Rhinolophus alcyone Rhinolophus blasii Peters, 1867 1857. Rhinolophus clivosus: Blasius, Säugethiere Deutschlands, 33, figs 10, 11. Type locality: Italy: "Italy". - Comments: Preoccupied by clivosus Cretzschmar, 1828 (see Corbet and Hill, 1992: 100 and Pavlinov et al., 1995: 81). *1867. Rhinolophus blasii Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 17 (for 1866). Publication date: 1867. Type locality: Italy: "Italy". Holotype: ZMB 557: ♂, alcoholic (skull not removed). Collected by: Johan Heinrich Blasius. See Turni and Kock (2008). - Comments: Type locality originally SE Europe, but restricted to Italy by Ellerman et al. (1953: 59), and Trieste (see Turni and Kock (2008). - Etymology: In honour of the German naturalist Johann Heinrich Blasius (1809 - 1870) (see Kozhurina, 2002: 14). (Current Combination) 1904. Rhinolophus empusa K. Andersen, Ann. Mag. nat. Hist., ser. 7, 14 (83): 378. Publication date: 1 Novmber 1904. Type locality: Malawi: Zomba [15 23 S 35 19 E,, 800 - 900 m] [Goto Description]. Holotype: BMNH 1893.7.9.33: ad ♀. Collected by: Alexander ("Alex") Whyte (F.Z.S); collection date: January 1893. - Comments: Considered a valid subspecies by Ansell and Dowsett (1988: 33), Taylor (1998: 37). 1905. Rhinolophus andreinii Senna, Arch. Zool. ital., 2: 256, pl. 16, f. 1; pl. 18, f. 7 - 16. Publication date: 30 September 1905. Type locality: Eritrea: Adi Ugri [14 53 N 38 49 E, 1 900 m]. 1910. Rhinolophus blasiusi Trouessart, Fauna des Mammifères d'Europe, 9. 1910. Rhinolophus brockmani Thomas, Ann. Mag. nat. Hist., ser. 8, 5 (26): 192. Publication date: 1 February 1910. Type locality: Somalia: Upper Sheikh [4 500 ft] [Goto Description]. Holotype: BMNH 1909.12.17.4: ad ♀. Collected by: Dr. Ralph Evelyn Drake-Brockman; collection date: 13 November 1909; original number: 237. Thomas (1910a: 193) mentions two specimens. - Etymology: In honour of Dr. R.E. DrakeBrowkman, the collector of the type specimen. 2019. Rhinolophus basii: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) ? Rhinolophus blasii andreinii: (Name Combination) ? Rhinolophus blasii empusa: (Name Combination) African Chiroptera Report 2020 TAXONOMY: Figure 75. Rhinolophus blasii. Includes brockmani; see Koopman (1975: 383). Replacement for clivosus Blasius, 1857, which was preoccupied (see Ansell and Dowsett, 1988: 33). Part of the landeri species group (Csorba et al., 2003: 57 - 59; Simmons (2005: 351 - 352). See Csorba et al. (2003: 58 - 59) for remarks on taxonomy. Herkt et al. (2017: Appendix S9) indicate that three subspecies are recognized in continental Africa which are relatively isolated geographically: blasii in Northwest Africa, andreinii in Ethiopia and empusa in southern Africa. No genetic tests have been performed yet to check the validity of these taxa. The analyses carried out by Herkt et al. (2017), however, indicate substantial ecological niche differences as well as major geographic gaps in suitable habitat areas between them. COMMON NAMES: Afrikaans: Spitssaalneusvlermuis. Albanian: Lakuriq nate hundëpatkua i Blasius-it. Arabian: Khaffash. Armenian: Բլազիի կամ, միջերկրածովային պայտաքիթ. Azerbaijani: Blazius nalburunu. Basque: Blasius ferrasaguzar. Belarusian: Падкаванос Блазіўса. Bosnian: Blazijev potkovasti ššmiš. Breton: Frigribell Blasius. Bulgarian: Средиземноморски подковонос. Castilian (Spain): Murciélago de herradura de Blasius, Rinolofo de Blasius, Murciélago dalmata de herradura. Catalan (Spain): Ratpenat de ferradura de Blasius. Croatian: Blazijev potkovnjak. Czech: Vrápenec Blasiův, vrapenec pahrbeční. Danish: Blasius' hestekonæse. Dutch: Blasius' hoefijzerneus, Blasiushoefijzerneus. English: Peters's Horseshoe Bat, Blasius's Horseshoe Bat, Peak-saddle Horseshoe Bat. Estonian: Hele-sagarnina. Finnish: Vaaleaherkko. French: Rhinolophe de Blasius, Rhinolophe à selle pointue. Frisian: Blasius' hoefizernoas. Galician (Spain): Morcego de ferradura de Blasius. Georgian: ბლაზის ცხვირნალა. German: Blasius-Hufeisennase. 273 Greek: Μεσορινόλοφος, Ρινόλοφος του Blasius. Hebrew: ‫מצוי פרסף‬, parsaf matzuy, Parsaf Matsui. Hungarian: Csúcsosnyergű patkósdenevér. Irish Gaelic: Crú-ialtóg Blasius. Italian: Rinolofo di Blàsius, Ferro di cavallo di Blàsius. Latvian: Blāzija pakavdegunis. Lithuanian: Blazijaus pasagnosis. Luxembourgish: BlasiusHuffeisennues. Macedonian: Бласиев потковичар [= Blasiev Potkovichar]. Maltese: Rinolofu ta' Blasius. Montenegrin: Sredozemni potkovičar. Norwegian: Balkanhesteskonese. Polish: Podkowiec Blasiusa. Portuguese: Morcego-de-ferradura de Blasius, Morcego ferradura de nariz de sela. Rhaeto-Romance: Rinolof da Blasius. Romanian: Liliacul cu potcoavă a lui Blasius, Liliacul-lui-Blasius. Russian: Подковонос средиземноморский. Serbian: Јужни потковичар [= Južni potkovičar]. Scottish Gaelic: Crudh-ialtag Blasii. Slovak: Podkovár Blasiov. Slovenian: Blasijev podkovnjak. Swedish: Balkanhästskonäsa. Turkish: Blasius Nalburunlu Yarasa. Ukrainian: Підковик (Підковоніс) Блазіуса. Welsh: Ystlum pedol Blasius. CONSERVATION STATUS: Global Justification Assessed as Least Concern (LC ver 3.1 (2001)). Widespread, although patchily distributed. There are some large colonies and the global population is likely to considerably exceed 10,000. Although the population is declining in some areas (e.g. western Balkans), it is stable in others (Jacobs et al., 2008ag; IUCN, 2009; Taylor, 2016f). Assessment History Global 2016: LC ver 3.1 (2001) (Taylor, 2016f). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008ag; IUCN, 2009). 2004: NT ver 3.1 (2001) (Jacobs et al., 2004x; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional South Africa:- 2016: NT D1 ver 3.1 (2001) (Jacobs et al., 2016b). 2004: VU D2 ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: Threats to the species include loss of Mediterranean woodlands, disturbance and loss of underground habitats, and destruction of roost sites (Kryštufek, 1999). In a number of range states the species is disturbed by tourist visits to caves and by use of the caves as shelters for livestock (Jacobs et al., 2008ag; IUCN, 2009; Taylor, 2016f). 274 ISSN 1990-6471 The most important risk factors related to climatic change were identified by Sherwin et al. (2012: 174) as being their roosting in caves/trees, the aerial feeding style and its long-distance dispersal. Bilgin et al. (2012: 434) predict that R. blasii will have a major range contraction as a result of climatic change. recent records in Romania and northern Bulgaria despite intensive work by Christian Dietz (Z. Nagy pers. comm., 2006). Past records from this area are disputed: no specimen has been found in museums in Romania, and the presence of R. euryale in the same area might have caused confusion (Z. Nagy pers. comm., 2006). CONSERVATION ACTIONS: Taylor (2016f) supports Jacobs et al. (2008ag) [in IUCN (2009)] who reported that Rhinolophus blasii is protected by national legislation in some range states. There are international legal obligations for the protection of this species through the Bonn Convention (Eurobats) and Bern Convention in areas to which these apply. It is included in Annex II (and Annex IV) of the EU Habitats and Species Directive, and hence requires specific conservation measures in some range states, including the designation of Special Areas for Conservation. It occurs in some protected areas. Taxonomic research is needed to clarify the status of the African populations. Monitoring and protection of caves is also required. It occurs from sea level to 2,215 m in Yemen. GENERAL DISTRIBUTION: Rhinolophus blasii has a large range in the Palaearctic and the Afrotropics, throughout which it is widely but patchily distributed. Its range extends marginally into the Indomalayan region. In Africa, it occurs from northeastern South Africa and the Democratic Republic of Congo, through south Malawi, to East African, Ethiopia and Somalia, and in North Africa. Follow Taylor (2000) for southern African distribution. In North African it is only present in Morocco and Algeria (it may occur in Tunisia but there are no confirmed records yet, and likewise for Egypt). Altitude range is from sea level to 1,400 m. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 602,154 km 2. In South Africa, its distribution is strongly associated with the amount of precipitation in the warmest quarter of the year (Babiker Salata, 2012: 49). In Asia, it has a patchy distribution extending from Turkey in the west to Pakistan in the east, and from the Caucasus in the north to Yemen in the south (Simmons, 2005). It was confirmed in Georgia in 2006 (Z. Nagy pers. obs.). In Europe, it is extinct in northeastern Italy and has not been recorded in Slovenia during the last 50 years (Kryštufek, 2001). Also recorded from western Anatolia and from the Levant (Syria, Lebanon, Jordan, Palestine and Israel). It is now restricted to the Balkan peninsula and to some Mediterranean islands including Crete and Cyprus. There are no Native: Afghanistan; Albania; Algeria; Armenia; Austria; Azerbaijan; Bosnia and Herzegovina; Botswana; Bulgaria; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 555); Croatia; Cyprus; Eritrea; Ethiopia; Greece (Kriti); Iran, Islamic Republic of; Israel; Italy; Jordan; Libyan Arab Jamahiriya; Malawi (Ansell, 1978; Happold et al., 1988; Happold and Happold, 1997b: 818; Monadjem et al., 2010d: 555); Montenegro; Morocco (Benda et al., 2010a: 157; El Ibrahimi and Rguibi Idrissi, 2015: 359); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 555; Monadjem et al., 2010c: 377); Oman; Pakistan; Palestinian Territory, occupied; Serbia; Somalia; South Africa (Monadjem et al., 2010d: 555); Swaziland (Monadjem, 2005: 5; Monadjem et al., 2010d: 555); Syrian Arab Republic; Tanzania; Tunisia (Aellen and Strinati, 1970; Cockrum, 1996; Dalhoumi et al., 2011; 2014: 53); Turkey; Turkmenistan; Yemen; Yugoslavia; Zambia (Ansell, 1969; Ansell, 1978; Monadjem et al., 2010d: 555); Zimbabwe (Monadjem et al., 2010d: 555). Possibly extinct: Slovenia. DETAILED MORPHOLOGY: Baculum - See Csorba et al. (2003: 4). In their study on take-off performance, Gardiner et al. (2014: 1059) determined a wing beat frequency of 12.96 ± 0.27 Hz. FUNCTIONAL MORPHOLOGY: In a study of flight-initiating jumps, Gardiner and Nudds (2011: 2187) found that the ground gleaning R. blasii did not outperform the fast-flying hawker (Miniopterus schreibersii) nor the trawling (Myotis capaccinii), although it spends a major part of its time foraging on the ground. ECHOLOCATION: Jacobs et al. (2007a) reported for individuals from South Africa a peak frequency of 86.6 (± 0.7) kHz (detector type not reported), while Monadjem et al. (2007a), using an Anabat detector, reported the same peak frequency albeit a different standard deviation (± 0.23) for an individual from Swaziland. Also from Swaziland, Monadjem et al. (2017c: 179) reported Fmin: 79.4 ± 1.86 (77.9 - 81.5) kHz, African Chiroptera Report 2020 Fknee: 86.3 ± 0.24 (86.1 - 86.6) kHz, Fc: 86.2 ± 0.21 (86.0 - 86.4) kHz, duration: 20.8 ± 0.72 (19.9 21.3) msec. Monadjem et al. (2010c: 377) reported for Mozambique that peak echolocation frequencies ranged between 93.2 - 95.4 kHz (ANABAT, Petterson D240x). Jones and Siemers (2010: 449 - 450) indicated that females emit pulses with a higher frequency than males, and that juveniles emit lower frequencies than adults. Papadatou et al. (2008b: 132) report the following data for 37 Greek specimens: Fstart: 90.3 ± 4.85 kHz, Fend: 78.1 ± 4.48 kHz, Fpeak: 94.0 ± 0.92 kHz, bandwidth: 4.0 ± 1.65 kHz, duration 44.1 ± 12.14 msec and interpulse interval: 71.8 ± 23.10 msec. The following data for 10 Iranian calls were reported by Benda et al. (2012a: 182): Fstart: 94.0 ± 0.1 (94.2 - 94.4) kHz, Fend: 92.0 ± 0.1 (91.8 - 92.1) kHz, Fpeak: 93.1 ± 0.1 (93.0 - 93.3) kHz, duration: 48.6 ± 9.4 (40.1 - 68.3) msec and interpulse interval: 109.3 ± 5.6 (103.1 - 117.1) msec. Walters et al. (2012: suppl.) report the following figures for 26 calls (4 sequences) from bats from Algeria and Greece: duration: 44.90 ± 8.52 msec, Fmax: 95.51 ± 1.34 kHz, Fmin: 80.58 ± 3.52 kHz, bandwidth: 14.93 ± 3.82 kHz, Fpeak: 95.41 ± 1.33 kHz. Luo et al. (2019a: Supp.) reported the following data: Fpeak: 94 kHz, Fstart: 80.3 kHz, Fend: 78.1 kHz, and duration: 44.1 msec. Disca et al. (2014: 226) indicate that in Morocco the type of call is FM-FC-FM, with Fpeak: 96.7 ± 0.3 kHz and duration: 20.7 ± 4.9 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Ðulic (1967) reported 2n = 58, FN = 60, BA = 4, a submetacentric X and an acrocentric Y for specimens from Croatia. Qumsiyeh et al. (1986) and Rautenbach (1986) also reported 2n = 58, FN = 60, BA = 4, but differed in the X chromosome being reported in the respective publications as metacentric and submetacentric, and the Y chromosome as acrocentric and metacentric. Arslan and Zima (2014: 9) report that the autosomal complement consists of two medium-size bi-armed pairs and 26 acrocentric pairs, with one of the acrocentrics having a secondary constriction. Protein / allozyme - Unknown. 275 HABITS: Siemers and Ivanova (2004: 470) report that R. blasii uses aerial and foliage-gleaning in flight, as well as ground-gleaning (including landing) to detect and capture their prey. DIET: In northern Algeria, Ahmim and Moali (2013: 46) found that R. blasii droppings contained 96.87 % Insecta and 3.13 % Chilopoda. The most frequent insect preys were Diptera (37.50 %), specifically Chironomidae/Ceratopogonidae (9.38 %), Culicidae, Anisopodidae and Sphaeroceridae (6.25 %). Trichoptera also were frequently present (15.63 %), followed by Lepidoptera and Hemiptera (both 12.50 %). Gardiner et al. (2014: 1058) assessed its foraging strategy as "low altitude slow aerial hawking", sometimes with gleaning. POPULATION: Structure and Density Africa: Not very common (Jacobs et al., 2008ag; Taylor, 2016f). Asia: This species has a widespread distribution and the populations in Pakistan and Afghanistan seem to be stable and doing well (Molur et al., 2002). Europe: A rare or infrequent species, probably the rarest horseshoe bat in Europe (Kryštufek, 1999). Summer colonies of ca. 20 30 are typical, although up to 400 females may be found in a single colony. In winter, it congregates in mixed-species clusters with other Rhinolophus species (up to 2,000 animals in Serbia). There are large colonies in Serbia, Bulgaria and Greece. It is suspected to be declining because of loss of Mediterranean woodlands and cave disturbance, and is considered vulnerable in many range states (e.g., the western Balkans); however, the populations in the eastern Balkans are stable (Mediterranean Workshop, 2007) (Jacobs et al., 2008ag; IUCN, 2009; Taylor, 2016f). Trend 2016: Decreasing (Taylor, 2016f). 2008: Decreasing (Jacobs et al., 2008ag; IUCN, 2009). R. blasii populations within the Asia Minor and Levant region are predicted to have a declining population trend, under climate change scenarios (Bilgin et al., 2012: 433). REPRODUCTION AND ONTOGENY: In Malawi, single young are born from midNovember to early January (beginning of the wet season) and these are lactated for about a month (Happold and Happold, 1990b: 566). The females are also not in close reproductive synchrony. 276 ISSN 1990-6471 PARASITES: Clément et al. (2019: 5) reported on 10 European or southern African R. blasii specimens of which one was infected by Trypanosoma livingstonei_A. Ševcík et al. (2012: 35) report on the presence of the bat flies Phthiridium biarticulatum Hermann, 1804 (Nycteribiidae) and Brachytarsina flavipennis Macquart, 1851 (Streblidae) on bats from Cyprus. The same parasites were found by Benda et al. (2010b: 243) in Jordan. B. flavipennis was also reported from Iran by Benda et al. (2012a: 255). Benda et al. (2010b) also refer to previous records of Penicillidia conspicua Speiser, 1901 and P. dufourii (Westwood, 1835) from Palestine, Phthiridium integrum (Theodor and Moscona, 1954) and Penicillidia fulvida (Bigot, 1885) from Yemen (all Nycteribiidae); a streblid Ascodipteron africanum rhinolophi Jobling, 1958 from Yemen; and an ischnopsillid Rhinolophopsylla unipectinata (Taschenberg, 1880) from Yemen. Nycteridocoptes eyndhoveni Fain, 1959 (Acari: Sarcoptidae) was reported from "R. blasii empusa" from Tanganika Moba, DCR by Fain (1959a: 341). Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Raymondia alulata Speiser, 1908 (Shapiro et al., 2016: 253). Raymondia hardyi Fiedler, 1954 (Shapiro et al., 2016: 254). Raymondia waterstoni Jobling, 1931 (Haeselbarth et al., 1966: 104). Ascodipteron lophotes Monticelli, 1898 (Haeselbarth et al., 1966: 106, but Maa (1965) suggests that these records need verification). VIRUSES: Coronaviridae - Coronaviruses Luis et al. (2013: suppl.) mention the presence of BatCoV BB98-15 and a SARS coronavirus-related virus. Alphacoronavirus Kohl and Kurth (2014: 3112) reported this virus from German bats. Betacoronavirus Reported from bats from Bulgaria and Germany by Kohl and Kurth (2014: 3112). Picornaviridae Nieto-Rabiela et al. (2019: Suppl.) mentioned the presence of Picornavirus. Polyomaviridae Carr et al. (2017) reported two new virsuses from R. blasii from Zambia: Rhinolophus blasii polyomavirus 1 and Rhinolophus blasii polyomavirus 2. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Congo (Democratic Republic of the), Eritrea, Eswatini, Ethiopia, Italy, Liberia, Malawi, Morocco, Mozambique, Somalia, South Africa, Tanzania, Tunisia, Zambia, Zimbabwe. Nycteribiidae: Nycteribia integra Theodor and Moscona, 1954 (Haeselbarth et al., 1966: 109). Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114). Bendjeddou et al. (2017: 15) reported the following ectoparasites from Algerian bats: Nycteribiidae: Nycteribia (Nycteribia) latreillii (Leach, 1817), Penicillidia (Penicillidia) dufourii Westwood, 1835 and Phthiridium biarticulatum Hermann, 1804; Streblidae: Brachytarsina (Brachytarsina) flavipennis Macquart, 1851 and Arachnida: Eyndhovenia euryalis (G. Canestrini, 1885). Figure 76. Distribution of Rhinolophus blasii Rhinolophus capensis Lichtenstein, 1823 *1823. Rhinolophus capensis Lichtenstein, Verzeichniss der Doubletten des Zool. Mus. K. Univ. Berlin, 4. Type locality: South Africa: Cape province: Cape of Good Hope [ca. 33 56 S 18 28 E]. Lectotype: ZMB 377: juv, skin and skull. Collected by: Krebs; collection date: 1822. Skull damaged. Lectotype designated by Turni and Kock (2008: 32). - Comments: Turni and Kock (2008: 32) also indicate that originally there were two additional specimens, which might have been sold to an unknown collection. Furthermore, Turni and Kock (2008: 32) mention that ZMB 376 and 378, which were longtime considered syntypes of African Chiroptera Report 2020 1860. 277 capensis, were re-identified by Karl Koopman as belonging to Rhinolophus clivosus Cretzschmar, 1828. - Etymology: Referring to Cape of Good Hope or the Cape province, from where the type specimen was obtained. (Current Combination) Rhinolophus auritus Sundevall, Kongl. Svenska Vet.-Akad. Hand., ser. 2, 2: no 10, p. 13, footnote. Type locality: South Africa: Southern Cape province: Cape of Good Hope, near Knysna: Belvedere [ca. 33 53 S 22 59 E]. Holotype: NRM ♂. Collection date: 3 April 1854. Victorin; Grill 1859; Mam. Ex. No. 1907: see Andersen (1904c: 456). TAXONOMY: Part of the capensis species group (Csorba et al., 2003: 7 - 8; Simmons, 2005: 352). See Csorba et al. (2003: 8) for remarks on taxonomy. Reviewed by Stoffberg (2008). COMMON NAMES: Afrikaans: Kaapse saalneusvlermuis, Kaapse Vlermuis. Czech: vrápenec jihoafrický. English: Cape Horseshoe Bat, Southern Africa horseshoe bat. French: Rhinolophe d'Afrique du Sud, Rhinolophe du Cap. German: KapHufeisennase, Capischen Kamnase, Russbraunen Kamnase. Portuguese: Morcego ferradura de cabo. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: De Graaf (1960) identified material as R cf. capensis from the Late Pliocene, at Makapansgat, South Africa. Avery and Avery (2011: 16) reported Holocene remains from Blinkklipkop, Powerhouse Cave, Wonderwerk (Northern Cape province, South Africa), as well as Pleistocene remains from the latter locality. CONSERVATION STATUS: Global Justification An endemic species to South Africa. While declining in parts of its range, the species is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, its known large population (there are many records of this species occurring in high numbers in coastal caves), and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008z; IUCN, 2009; Jacobs and Monadjem, 2017). Assessment History Global 2016: LC ver 3.1 (2001) (Jacobs and Monadjem, 2017). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008z; IUCN, 2009). 2004: NT ver 3.1 (2001) (Jacobs et al., 2004w; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016c). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: The species is threatened in parts of its range by disturbance of cave roosts (often by recreational and tourism activities), and the conversion of suitable foraging habitat to agricultural use (Jacobs et al., 2008z; IUCN, 2009; Jacobs and Monadjem, 2017). CONSERVATION ACTIONS: Jacobs and Monadjem (2017) the species is recorded from more than 10 protected areas including: West Coast National Park; De Hoop Nature Reserve; Garden Route National Park; Langeberg Nature Reserve; Addo Elephant National Park; Great Fish Nature Reserve; Kologha Forest Reserve and Kubusi Indigenous State Forest. While no urgent conservation interventions are necessary, the species would benefit from further protected area establishment once key roost sites have been identified; and artificial wetlands in agricultural landscapes should be managed for biodiversity by conserving patches of native vegetation around the waterbodies (Sirami et al., 2013). Further studies are needed into the distribution of this bat it may occur in southern Namibia (Sirami et al., 2013). GENERAL DISTRIBUTION: Rhinolophus capensis is restricted to the coastal belt of the Northern Cape, the Western Cape and the Eastern Cape of South Africa, as far east along the coast as the vicinity of East London (Csorba et al., 2003: 8; Stoffberg, 2008: 2). Odendaal et al. (2014: 4) indicate that the geographic range of R. capensis is determined by two environmental gradients that influence the regional rainfall patterns. First, there is the increasing aridity towards the north and secondly a seasonality shift from west to east, whereby the rains are dominant during the winter in the west, then becoming aseasonal and finally being dominant during the summer in the extreme east. Simmons (2005: 352) follows Ansell and Dowsett (1988: 33) and Koopman (1993a: 164), who list South Africa, Zimbabwe and Mozambique, but does indicate doubt about its occurrence outside of South Africa. Koopman (1993a: 164) states that the records from Zambia and Malawi are erroneous. Griffin (1999: 32), however, suspects its occurrence in Namibia. In the RSA, the 278 ISSN 1990-6471 species' distribution is best predicted by biome: desert, Albany thicket, fynbos and grassland (Babiker Salata, 2012: 50, 118). and produced calls with higher frequencies, and therefore suggest that there might be a potential social role for the peak frequency. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 277,925 km2 (or 4 % of the total surface of the area). When flying with conspecifics, Fawcett et al. (2013: 49) found that the terminal FM sweep became longer and wider in bandwidth than when flying alone. Fawcett et al. (2015: 693), furthermore, found that R. capensis produced longer frequency-modulated downward sweeps at the end of their call when they flew together with bats from the same species or with Miniopterus natalensis than when they flew alone. Additionally, when they were flying with R. clivosus (a larger high-duty cycle bat), its calls were shorter than when flying alone or with other R. capensis individuals. Native: South Africa (Eastern Cape Province, Northern Cape Province, Western Cape Province) (Ellerman et al., 1953; Maree and Grant, 1997; Csorba et al., 2003: 8; Monadjem et al., 2010d: 556). Presence uncertain: Namibia. FUNCTIONAL MORPHOLOGY: Jacobs et al. (2007a), at De Hoop, South Africa, found R. capensis to have a lower mean wing loading and aspect ratio than R. clivosus, but no differences in their mean shape indices. SEXUAL DIMORPHISM: At De Hoop, South Africa, Jacobs et al. (2007a) found females were heavier than males, but there were no differences in forearm length, or wingspan. ECHOLOCATION: Jacobs et al. (2007a) reported a peak frequency of 83.9 (+ 1.2) kHz for individuals from South Africa. Odendaal and Jacobs (2009: 62) found some sort of local dialects in the echolocation calls for South African R. capensis: 84.60 ± 0.82 kHz at De Hoop, 85.84 ± 0.73 kHz at Table Farm, and 80.66 ± 0.50 kHz at Steenkampskraal, which is probably the result of interactions with other Rhinolophus species: R. swinnyi at Steenkampskraal, and R. darlingi at Table Farm, as well as with R. clivosus. Bastian and Jacobs (2013: 18) and Odendaal et al. (2013: 112) indicate that the peak frequency ranges from 75.7 kHz at the western end of the species' coastal distribution in South Africa to 85 86.5 kHz at the eastern end. This increase seems to be linked with the mean annual rainfall (Odendaal et al., 2013). Bastian and Jacobs (2013: 18) also found that the 85 kHz specimens were able to discriminate between 85 kHz calls from conspecifics and calls from R. damarensis, which also peaks at 85 kHz. The 75 kHz specimens were unable to make that discrimination. Odendaal and Jacobs (2010b: 240) reported that there was no relation between body size and peak frequency across populations. However, nasal chamber length was found to be the best predictor on peak frequency. They also found that in each population, the females were larger than the males Bastian and Jacobs (2015) investigated the ability for R. capensis to discriminate their own calls from those of allopatric (R. damarensis) and sympatric (R. clivosus) congeners. The bats had no problems with the calls from R. clivosus, but the calls from R. damarensis could only be discriminated by one of the test groups. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach (1986) reported 2n = 58, FN = 60, BA = 4, and the X and Y chromosomes as submetacentric. Erasmus and Rautenbach (1984: 94) mention 52 acrocentric and four biarmed autosomal chromosomes, resulting in 60 autosomal arms. Protein / allozyme - see Maree and Grant (1997). HABITAT: Sirami et al. (2013: 34) recorded in the Western Cape Province, South Africa that R. capensis activity was not significantly, nor positively influenced by size of wetland, while habitats 100 m surrounding wetlands were also significantly and positively influenced by the water body. R. capensis preferentially flew near trees, orchards surrounding wetlands and over these wetlands (Sirami et al., 2013: 35). HABITS: Jacobs et al. (2007a) at De Hoop, South Africa found that the average esitmated flying height above ground 1.00 ± 0.77 m, n= 11. At the same locality, Thomas and Jacobs (2013: 129) found that Lepidoptera eating bats (e.g. R. capensis and Nycteris thebaica) emerged earlier than bats feeding on Diptera (e.g. Tadarida aegyptiaca). DIET: Jacobs et al. (2007a) at De Hoop, South Africa, found that the diet mainly comprised Lepidoptera African Chiroptera Report 2020 and Coleoptera with lesser amounts of Neuroptera, Hemiptera and Diptera. The mean size of measurable prey taken by R. capensis was smaller than that taken by R. clivosus, although the range of prey sizes taken by the two species overlapped (Jacobs et al., 2007a). PREDATORS: Avery et al. (2005: 1054) found this species in pellets from Tyto alba in South Africa. POPULATION: Structure and Density:- This species is common throughout its range (Bernard, 2013a), and is relatively well represented in museums (Monadjem et al., 2010d). Skinner and Chimimba (2005) state that 'they are abundant in the Western Cape and the Eastern Cape, where there are many records from coastal caves. It can be found in colonies consisting of thousands of individuals (Herselman and Norton, 1985; Taylor, 2000; Skinner and Chimimba, 2005). Example, there are an estimated 19,000 individuals in De Hoop Guano Cave (McDonald et al., 1990b). Trend:- 2016: Stable (Jacobs and Monadjem, 2017). 2008: Decreasing (Jacobs et al., 2008z; IUCN, 2009). months 2000). 279 (see also Bernard, 1984; Krutzsch, Bernard (1985: 134) indicates that females do not become reproductively mature in their first year, and that males would only be able to mate in their second year. PARASITES: Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Nycteribiidae: Nycteribia schmidlii Schiner, 1853 (Haeselbarth et al., 1966: 108). SIPHONAPTERA Ischnopsyllidae: Rhinolophopsylla ashworthi (Waterson, 1913) known only from a few specimens from the Cape Province (King William’s Town, Willowmore and Oudtshoorn), South Africa (Haeselbarth et al., 1966: 191). Rhinolophopsylla capensis Jordan and Rothschild, 1921 (Haeselbarth et al., 1966: 191). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Namibia, South Africa. REPRODUCTION AND ONTOGENY: Bojarski and Bernard (1992) [in Anthony (2000: 16)] report that immunocytochemical indices of gonadotropic function rise significantly during the spermatogenic period, coincident with high plasma testosterone levels. Bernard (1985: 129) and Martin and Bernard (2000: 36) mention that spermatogenesis starts in spring (October in South Africa) and ends after early winter (May) although sperm remain in the caudae epididymides through winter (May to September). Copulation and ovulation occur in August to September, at the end of winter hibernation. Parturition occurs in November and December after a gestation period of three to four Figure 77. Distribution of Rhinolophus capensis Rhinolophus clivosus Cretzschmar, 1828 *1828. Rhinolophus clivosus Cretzschmar, in: Rüppell, Atlas Reise Nördlichen Afrika, Zoologie Säugethiere, 1: 47, pl. 18. Type locality: Saudi Arabia: Red Sea Coast: Al Muwaylih [=Mohila] [ca. 27 49 N 35 30 E] [Goto Description]. Paratype: SMF 12311: skin only. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Lectotype: SMF 4371: mounted skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell; collection date: 1826. Old catalog: II.F.4.a; see Mertens (1925: 20). Comments: Allen (1939a: 74) mentions "1826" (as mentioned on the title page) as year of publication of Cretzschmar's work, other authors mention it as "1827", "1828", "18301831". - Etymology: From the masculine Latin adjective clivósus, meaning "characterized by slopes, steep", referring to a seemingly unreliable distinctive character of the noseleaf 280 ISSN 1990-6471 1829. 1861. 1904. 1904. 1904. 1904. 1905. 1916. 1932. 1934. "Rhinolophus apparato olfactorio externo clivis gradatim elatis non dissimili" (Cretzschmar 1828) (see Lanza et al. (2015: 80). (Current Combination) Rhinolophus Geoffroyii A. Smith, Zool. Journ., 4 (16): 433. Publication date: May 1829. Type locality: South Africa: "South Africa" [Goto Description]. - Comments: Roberts (1951: 61) nominated Cape Town as type locality; see Meester et al. (1986: 38). Allen (1939a: 75) mentioned geoffroyii as a separate species. Meester et al. (1986: 38) follow Ellerman et al. (1953: 57) in considering geoffroyii as unidentifiable. Rhinolophus acrotis Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 10. Publication date: 1861. Type locality: Eritrea: Kérén [ca. 15 45 N 38 20 E] [Goto Description]. Holotype: SMNS 986a: ad ♂, alcoholic (skull not removed). Collected by: Martin Theodor von Heuglin; collection date: 1862. Paratype: SMNS 986b: juv ♂, alcoholic (skull not removed). Collected by: Martin Theodor von Heuglin; collection date: 1862. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 293) mention this specimen as a possible paratype, although they do indicate that Heuglin based his description on one single specimen (probably SMNH 986a). They also indicate that this specimen was marked as "Cotype" by Krauss. - Etymology: From the two neuter Greek substantives "ἄκρον" (akron), meaning "extremity" and "οὖς", meaning "ear" (see Lanza et al., 2015: 80). Rhinolophus Andersoni Thomas, Ann. Mag. nat. Hist., ser. 7, 14 (80): 156. Publication date: 1 August 1904. Type locality: Egypt: Eastern desert of Egypt [ca. 22 N 35 E] [Goto Description]. Holotype: BMNH 1904.11.4.2: imm ♂. Collected by: Arthur M. MacKilligan; collection date: 3 August 1903; original number: 26. - Comments: Kock (1969a: 113) mentions: "Eastern Desert, im Gebiet des Wadi Alagi, 22 N 35 E, Sudan". Rhinolophus augur K. Andersen, Ann. Mag. nat. Hist., ser. 7, 14 (83): 380. Publication date: 1 November 1904. Type locality: South Africa: Northern Cape province: [formerly Bechuanaland]: Kuruman [26 28 S 22 27 E, 4 000 ft] [Goto Description]. Holotype: BMNH 1904.10.1.1: ad ♂, skin and skull. Collected by: Mr. R.B. Woosnam; collection date: 19 April 1904; original number: 26. Rhinolophus augur zambesiensis K. Andersen, Ann. Mag. nat. Hist., ser. 7, 14 (83): 383. Publication date: 1 November 1904. Type locality: Malawi: Fort Hill [=Chitipa] [09 42 S 33 16 E, 1 300 m] [Goto Description]. Holotype: BMNH 1897.10.1.18: ad ♂, skin and skull. Collected by: Alexander ("Alex") Whyte (F.Z.S); collection date: July 1896; original number: 136. Presented/Donated by: Sir Harry Hamilton Johnston. - Comments: Considered a synonym of zuluensis by Ansell and Dowsett (1988: 32). Rhinolophus augur zuluensis K. Andersen, Ann. Mag. nat. Hist., ser. 7, 14 (83): 383. Publication date: 1 November 1904. Type locality: South Africa: KwaZulu-Natal: 20 mi [=32 km] NW Eshowe: Jususie valley [=Insuzi valley] [28 53 S 31 03 E, 1 000 ft] [Goto Description]. Holotype: BMNH 1904.5.1.8: ad ♀, skin and skull. Collected by: Captain Claude Henry Baxter Grant; collection date: 17 November 1903; original number: 602. Presented/Donated by: C.D. Rudd Esq. - Comments: Considered a valid subspecies by Ansell and Dowsett (1988: 32), Taylor (1998: 35). Rhinolophus acrotis brachygnatus K. Andersen, Ann. Mag. nat. Hist., ser. 7, 15 (85): 73. Publication date: 1 January 1905. Type locality: Egypt: Giza province: Giza [30 01 N 31 13 E] [Goto Description]. Holotype: BMNH 1892.9.9.7: ad ♂, alcoholic (skull not removed). Collected by: Dr. John Anderson; collection date: 16 December 1891. See Qumsiyeh (1985: 35). Paratype: [Unknown] imm ♀, alcoholic (skull not removed). See Qumsiyeh (1985: 35). Rhinolophus keniensis Hollister, Smiths. Misc. Coll., 66 (1) (2406): 2. Publication date: 10 February 1916. Type locality: Kenya: W side Mount Kenya [ca. 00 10 S 37 19 E, 7 000 ft] [Goto Description]. Holotype: USNM 166352: ad ♂, skull and alcoholic. Collected by: Edmund Heller; collection date: 27 August 1909; original number: 1154. Fisher and Ludwig (2015: 54) indicate that the Field Catalog mentions that this specimen was collected at the Base Camp on the West side of Mount Kenya. Rhinolophus geoffroyi zuluensis: Ortlepp, 18th Report of the Director of Veterinary Sciences and Animal Industry. Union of South Africa. (Name Combination) Rhinolophus acrotis schwarzi Heim de Balsac, Bull. Mus. natn. Hist. nat., Paris, sér. 2, 7: 483. Publication date: November 1934. Type locality: Algeria: Sahara, Tassai des Azdjers: Djanet [24 34 N 09 30 E] [Goto Description]. Holotype: MNHN ♀, alcoholic (skull not removed). Collected by: La Mission Augiéras Draper. Holotype: MNHN ZMMO-2006-246: ♀, skull and alcoholic. Collected by: Dr. Henry Foley; collection date: African Chiroptera Report 2020 1959. 1980. 1993. 2019. ? ? ? ? ? ? ? ? ? ? 281 1934. Paratype: MNHN ZM-MO-2006-247: ♀, skull only. Collected by: Dr. Henry Foley; collection date: 1934. Paratype: MNHN ZM-MO-2006-248: ♀, skull only. Collected by: Dr. Henry Foley; collection date: 1934. Rhinolophus ferrum-equinum keniensis: Harrison, Occ. Pap. Nat. Mus. S. Rhod., 23B: 228.. (Name Combination) Rhinolophus ferrumequinum keniensis: Blackwell, Mycologia, 72 (1): 145. (Name Combination) angur: Koopman, in: Wilson and Reeder, Mammal species of the World (2nd Edition): Chiroptera, 164. (Lapsus) Rhinolophus clivossis: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) Rhinolophus clivosus acrotis: (Name Combination) Rhinolophus clivosus augur: (Name Combination) Rhinolophus clivosus brachygnathus: (Name Combination) Rhinolophus clivosus clivosus: (Name Combination) Rhinolophus clivosus keniensis: (Name Combination) Rhinolophus clivosus zuluensis: (Name Combination) Rhinolophus ferrum-equinum zuluensis: (Name Combination) Rhinolophus geoffroyi augur: (Name Combination) Rhinolophus geoffroyi geoffroyi: (Name Combination) Rhinolophus geoffroyi: (Name Combination, Alternate Spelling) TAXONOMY: Figure 78. Rhinolophus clivosus. ferrumequinum species group (Csorba et al., 2003: 35 - 39; Simmons, 2005: 353). See Csorba et al. (2003: 36 - 37) for review of taxonomy. Roberts (1919: 112) points out that geoffroyii Smith, 1829 antedates augur Andersen, 1904, and later (Roberts, 1951) regards augur as a subspecies of geoffroyii. Allen (1939a) follows Roberts in regarding geoffroyii as the prior name, but Ellerman et al. (1953) reject this, pointing out that the description does not identify geoffroyii and that the type is apparently lost, and therefore propose that the name be discarded as unidentifiable. The next available name would be Rhinolophus augur Andersen, 1904. Stoffberg et al. (2012: 1) indicate that southern African representatives of R. clivosus s.l. are as distinct from more northern African samples as they are from R. ferrumequinum. In southern Africa, they (p. 5) distinguish five geographic groups: Western Cape clade, Knysna region clade, Northern Cape clade, a predominantly KwaZulu-Natal/Mpumalanga mixed group and a Mpumalanga/Limpopo Province clade. About 3.7 Mya, these groups were separated from more northern groups (Kenya, Tanzania and Mozambique) and all of these were separated from the Egyptian populations at about 4.26 Mya. Stoffberg et al. (2012: 2; fig. 1) show the distribution area for four subspecies: geoffroyi in the Western Cape province and the coastal part of the Northern Cape province; zuluensis in the Eastern Cape province, KwaZulu-Natal province, Mpumalange province, southern Limpopo province and Swaziland and Lesotho; augur in eastern Northern Cape province and central Botswana; and zambesiensis in Mozambique, Zambia, Zimbabwe and the extreme southeastern DRC. On the other hand, Benda and Vallo (2012: 83) suggest that all populations occurring in the savanna belt from the Cape to Kenya belong to one taxon (subspecies): augur. Unfortunately, neither Stoffberg et al. (2012) nor Benda and Vallo (2012) provide any information on the validiy of the name "geoffroyii". Therefore, pending further investigation in the taxonomy of this complex, we retain the usage of the name R. clivosus. COMMON NAMES: Afrikaans: Geoffroy se saalneusvlermuis, Geoffroy-saalneusvlermuis, Geoffroyse vlermuis. Arabian: Khafash abu hadweh arabi, Khaffash. Chinese: 佐氏菊头蝠. Czech: vrápenec pouštní, wrápenec pahrbečný, vrapenec pahrbeční. Dutch: Geoffroy's hoefijzerneus. English: Arabian Horseshoe Bat, Cretzschmar's Horseshoe bat, Geoffroy's Horseshoe Bat. French: Rhinolophe de Cretzschmar, Rhinolophe de 282 ISSN 1990-6471 Geoffroy. German: Geoffroys Hufeisennase. Hebrew: ‫הנגב פרסף‬, Parsaf HaNegev. Italian: Rinòlofo di Cretzschrnar, Fèrro di cavàllo di Cretzschmar. Portuguese: Morcego ferradura gigante. ETYMOLOGY OF COMMON NAME: Colloquial name after E. Geoffroy Saint-Hilaire, who collected bats in Egypt during the early part of the nineteenth century (see Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: R. clivosus has been recovered from both the upper Pleistocene and very late Holocene at Border Cave and is possibly present in the late Holocene at Nkupe, South Africa (Avery, 1991: 6). Avery and Avery (2011: 17) report additional Holocene (Blinkklipkop and Wonderwerk), and Pleistocene (Wonderwerk) remains from the Northern Cape province (South Africa). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough globally to qualify for listing in a more threatened category (Kock et al., 2008a; IUCN, 2009; Monadjem et al., 2017cy). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cy). 2008: LC ver 3.1 (2001) (Kock et al., 2008a; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004an; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016d). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Although there are generally considered to be no major threats to the species as a whole, some populations are locally threatened by disturbance of their roosting sites, and indirect poisoning resulting from the use of insecticides, pesticides and similar chemicals (Kock et al., 2008a; IUCN, 2009; Monadjem et al., 2017cy). CONSERVATION ACTIONS: Kock et al. (2008a) [in IUCN (2009)] and Monadjem et al. (2017cy) reported that in view of the species wide range it seems likely that it is present in several protected areas. All bats, including Rhinolophus clivosus, are protected by national legislation in Jordan (Amr pers. comm., 2004). Bats of the genus Rhinolophus are generally susceptible to indirect poisoning through the local use of insecticides; there is a need to evaluate the impact of this threat on populations (especially in southwest Asia), and to investigate alternative methods of insect control (Kock pers. comm., 2004). GENERAL DISTRIBUTION: Rhinolophus clivosus is widespread in North, East and southern Africa, and also in parts of southwest Asia, including western and southeastern areas of the Arabian Peninsula. In North Africa it has been recorded from Algeria, Libya and Egypt; in East Africa, it ranges from Sudan in the north, through all East African countries to Malawi in the south; in southern Africa, it is present in Mozambique and Zambia in the north, ranging southwards into South Africa, Namibia and southern Angola. In addition, there are a number of records from southern and eastern parts of the Democratic Republic of the Congo. In southwest Asia, it ranges from Israel, Jordan and the Sinai Peninsula of Egypt in the north, through the western and southeastern part of the Arabian Peninsula of Saudi Arabia, Yemen and Oman, with a few additional records of the species from central regions of the Arabian Peninsula. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,120,944 km 2. In South Africa, its distributoin is strongly associated with annual precipitation (Babiker Salata, 2012: 49). ZMB has two specimens from the Central African Republic, that seem to be outliers. Native: Afghanistan; Algeria; Angola; Burundi; Cameroon; Chad; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 556); Djibouti (Pearch et al., 2001: 397); Egypt; Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 133); Israel; Jordan; Kenya (Lavrenchenko et al., 2004b: 133); Lesotho (Lynch, 1994: 190; Monadjem et al., 2010d: 556); Liberia; Libyan Arab Jamahiriya; Malawi (Happold et al., 1988; Happold and Happold, 1997b: 818; Monadjem et al., 2010d: 556); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 556; Monadjem et al., 2010c: 377); Namibia (Cotterill, 2004a: 261); Oman; Rwanda; Saudi Arabia; Somalia; South Africa (Monadjem et al., 2010d: 556); Sudan; Swaziland (Monadjem, 2005: 5; Monadjem et al., 2010d: 557); Tanzania (Stanley and Goodman, 2011: 41); Turkmenistan; Uganda (Kityo and Kerbis, 1996: 63); Yemen; Zambia (Ansell, 1978; Monadjem et al., 2010d: 557); Zimbabwe (Cotterill, 2004a: 261; Monadjem et al., 2010d: 557). African Chiroptera Report 2020 Presence Occupied. uncertain: Palestinian Territory, GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: For 5 specimens from the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) reported the following wing parameters: Fa: 53.4 ± 1.5 mm, Wing area: 175.9 ± 5.8 cm 2, Wing span: 33.8 ± 1.1 cm, Wing loading: 9.2 ± 0.8 N/m 2, and aspect ratio: 6.5 ± 0.3. DETAILED MORPHOLOGY: Baculum - See Thomas (1997) and Csorba et al. (2003: 36). Jacobs et al. (2013b: 76) found that R. clivosus has a smaller nasal capsule volume than predicted from its size. FUNCTIONAL MORPHOLOGY: Jacobs et al. (2007a), at De Hoop, South Africa, found R. clivosus to have a higher mean wing loading and aspect ratio than R. capensis, but no differences in their mean shape indices. SEXUAL DIMORPHISM: At De Hoop, South Africa, Jacobs et al. (2007a) found that females were heavier than males, but there were no differences in forearm length, or wingspan. ECHOLOCATION: Finger et al. (2017: 144) found that echolocation signals enables R. clivosus to distinguish the sex of the caller (probably linked with the initial and terminal FM components of the signal) and even its indentity (probably linked with resting frequency). Raw et al. (2018: 1) furthermore found that R. clivosus could discriminate readily between species using echolocation pulses with discrete frequencies. To discriminate between species with overlapping frequencies, additional parameters such as sweep rate of the FM and duty cycle were used. For individuals in South Africa, Taylor (1999b), using a Pettersson D980 detector at a locality in Dundee, reported a maximum frequency of 94.3 (± 0.2) kHz, while Jacobs et al. (2007a) (detector type and locality not reported) reported a peak frequency of 92.1 (± 0.7) kHz. Monadjem et al. (2007a), using an Anabat detector at several different localities in Swaziland, reported a maximum frequency of 91.9 (± 0.65) kHz. From the same country, Monadjem et al. (2017c: 179) reported Fmin: 75.7 ± 3.15 (70.4 - 80.3) kHz, Fknee: 91.5 ± 0.70 (90.0 - 92.3) kHz, Fc: 91.3 ± 1.06 (88.9 - 92.5) kHz and duration: 32.9 ± 5.35 (23.3 - 40.1) msec. Taylor (2000) indicates a constant frequency component of 94 kHz. Monadjem et al. 283 (2010c: 378) reported from Mozambique that the peak echolocation frequencies ranged between 79.8 - 81.0 kHz (ANABAT, Petterson D240x). From Algeria Forest Station (RSA), Schoeman and Jacobs (2002: 157) reported the type of calls to be high duty echolocation dominated by constant frequency calls, with a Fpeak: 92.7 ± 0.5 kHz. Monadjem et al. (2007a) also reported sexual dimorphism in the call with females emitting a higher frequency call than males (females 92.5 (± 0.04) kHz, males 91.6 (± 0.61) kHz). Taylor et al. (2013b: 19) recorded the following parameters for 10 calls from Soutpansberg, South Africa: Fmax: 92.1 ± 0.51 (91.5 - 92.8) kHz, Fmin: 86.7 ± 3.21 (89.6 - 90.9) kHz, Fknee: 90.3 ± 0.42 (89.6 - 90.9) kHz, Fchar: 90.7 ± 0.40 (90.1 - 91.4) kHz, duration: 14.3 ± 3.63 (8.6 - 19.4) msec. Linden et al. (2014: 40) reported the following parameters from the Soutpansberg area (RSA): Fmin: 79 - 90 kHz, Fchar: 90 - 91 kHz, Fknee: 91 - 91 kHz, Slope: 280 - 606 OPS, duration: 9 - 19 msec. Jacobs (2016: 133) and Jacobs et al. (2017b: 3) indicated that the call frequency of about 92 kHz in South Africa is much higher than expected for an animal of that size (which would be something around 66 kHz). They found that climatic factors, especially temperature, play an important role. Eisenring et al. (2016: SI 2) reported the following for 28 calls from the Aberdares Range in Kenya: PF: 84.9 ± 5.0 (66.9 - 88.3), HF: 86.8 ± 4.3 (69.7 89.7), LF: 68.4 ± 6.4 (56.4 - 81.8), DT: 0.4 ± 0.2 (0.0 - 0.6), DF: 18.4 ± 7.3 (4.9 - 32.5), IPI: 0.7 ± 0.7 (0.1 - 3.8). In Jordan, Benda et al. (2010b: 204) recorded the following parameters for 10 calls: Fstart: 73.1 ± 1.9 (71.5 - 77.0) kHz, Fend: 73.8 ± 2.3 (70.0 - 77.0) kHz, Fpeak: 84.4 ± 0.4 (83.5 - 85.0) kHz, Pulse duration: 48.0 ± 4.8 (43.0 - 59.0) msec, and Inter-pulse interval: 39.6 ± 12.9 (27.0 - 59.0) msec. From Israel, Hackett et al. (2016: 223) reported 66 calls with these values: Pulse duration: 54.48 ± 11.22 msec, Fstart: 78.92 ± 5.18 (69.3 - 88.8) kHz, Fend: 78.81 ± 5.26 (68.3 - 88.8) kHz, Fpeak: 85.18 ± 0.57 (83.4 - 88.8) kHz. Luo et al. (2019a: Supp.) reported the following data (Free flying bats): F peak: 85.18 kHz, Fstart: 78.92 kHz, Fend: 78.81 kHz, and duration: 54.48 msec. MOLECULAR BIOLOGY: DNA - Unknown. 284 ISSN 1990-6471 Karyotype - Both Ðulic and Mutere (1974) and Rautenbach et al. (1993) reported 2n = 58. However, Ðulic and Mutere (1974) reported FN = 62, BA = 6, a large acrocentric X chromosome, and a small acrocentric Y chromosome, whereas Rautenbach et al. (1993) reported FN = 60, BA = 4, and submetacentric X and Y chromosomes. Richards et al. (2016d: 187, 192) mentioned 2n = 58 and FN = 60, with primarily acrocentric chromosomes and two small metacentric pairs. Erasmus and Rautenbach (1984: 94) mention 50 acrocentric and six bi-armed autosomal chromosomes, resulting in 62 autosomal arms. Protein / allozyme - Unknown. HABITAT: Sirami et al. (2013: 34) recorded in the Western Cape Province, South Africa that R. clivosus activity was not significantly, nor positively influenced by size of wetland, while habitats 100 m surrounding wetlands were also significantly and positively influenced by the water body. R. clivosus preferentially near trees, orchards surrounding wetlands and over these wetlands (Sirami et al., 2013: 35). HABITS: Jacobs et al. (2007a) at De Hoop, South Africa found that the average esitmated flying height above ground 0.84 ± 0.20 m, n= 6. At the same locality, Thomas and Jacobs (2013: 124) found that R. clivosus emerged earlier than the other species in the area (in order of emergence): Miniopterus natalensis, Rhinolophus capensis, Nycteris thebaica, Myotis tricolor, Neoromicia capensis and Tadarida aegyptiaca. Petersen et al. (2019: 439) studied the behaviour and social calls of captive bats and found four acoustically distinct call types: 1) cascading/rising frequency-modulated (FM) calls, 2) oscillatory FM calls, 3) noisy screech calls and 4) whistle calls. These call types showed only weak associations with specific behaviours, situational categories or functional contexts. ROOST: Avery et al. (1990: 7) indicate that cool and moist caves or cave-like structures are used as daytime roosts. Taylor et al. (1999: 70) reported a roost containing 20 animals in the Shongweni Dam tunnel (RSA). DIET: From the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) reported the following prey groups based on 46 faecal pellets from 5 bats (in volume percent): Lepidoptera (62.6 ± 30.8), Coleoptera (29.4 ± 24.8), Hemiptera (2.0 ± 4.5), Hymenoptera (1.4 ± 3.1), and unknown (4.6 ± 6.4). Jacobs et al. (2007a) at De Hoop, South Africa, found that the diet mainly consisted of Lepidoptera and Coleoptera with lesser amounts of Neuroptera, Hemiptera and Diptera. The mean size of measurable prey taken by R. clivosus was larger than that taken by R. capensis, although the range of prey sizes taken by the two species overlapped (Jacobs et al., 2007a). PREDATORS: Avery et al. (2005: 1054) found this species to be present in pellets from the barn olw (Tyto alba) in South Africa. Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- Rhinolophus clivosus is common in some parts of its range, including South Africa, but is uncommon to rare in other areas, such as in Zimbabwe (Kock et al., 2008a; IUCN, 2009; Monadjem et al., 2017cy). In Swaziland, three separate populations contained over a thousand individuals, while in Jordan forty individuals were observed at a single location (Amr, 2000). Although the population appears to be stable in many regions, in Jordan the population is in decline and this bat may be declining throughout its range in the Arabian Peninsula (Amr pers. comm.) (Kock et al., 2008a; IUCN, 2009). Trend:- 2016: Unknown (Monadjem et al., 2017cy). 2008: Unknown (Kock et al., 2008a; IUCN, 2009). R. clivosus populations within the Asia Minor and Levant region have an unknown population trend, under predictive climate change scenarios (Bilgin et al., 2012: 433). REPRODUCTION AND ONTOGENY: Krutzsch (2000: 116) indicates that spermatogenesis occurs in late spring/summer/early fall. Copulation occurs in fall and is followed by sperm storage in the female until arousal and conception in the following spring. Stanley and Goodman (2011: 41) reported on four females collected in East Usambara (Tanzania), of which two (collected on 17 August 1992) had small teats and two (collected in the first half of August 1991 and 1993) had large, non-lactating teats. Six males collected in the first half of August included three animals with scrotal testes and three with abdominal testes. Four males African Chiroptera Report 2020 collected between July and September 1991 to 1993 in West Usambara had scrotal testes too. PARASITES: Hirst (1923: 989) mentions that Kolenati gave "R. clivosus [= R. blasii]" as host for the mite Periglischrus interruptus Kolenati, but that he corrected this later to R. euryale. Kolenati gave a new name to a mite from a R. clivosus specimen from Egypt: Periglischrus glutinimargo. From Jordan, Benda et al. (2010b: 229) reported the presence of a bat fly: Phthiridium biarticulatum Hermann, 1804 (Nycteribiidae) and a flea Rhinolophopsylla unipectinata unipectinata (Taschenberg, 1880) (Ischnopsyllidae). Additionally, they also mention literature date for Phthiridium integrum from Saudi Arabia and Yemen, Ascodipteron africanum rhinolophi Jobling, 1958 (Streblidae) from Yemen, and five mites from Yemen: Trombicula knighti Radford, 1954, T. filamentosa Radford, 1953, T. brevitarsa Radford, 1952, Brennanella longispina Radford, 1953, and Labidocarpus nasicolus Lawrence, 1938. ACARI Sarcoptidae: Fain (1959a: 340) mentions a paratype of Nycteridocoptes eyndhoveni Fain, 1959, collected from a R. clivosus zuluensis from Lubudi, DRC. Fain (1959c: 237) reported Psorergates (Psorergatoides) rhinolophi Fain from R. clivosus zuluensis from Uvira, Kivu, DRC. Trombiculidae: Trombicula (Grandjeana) reticulata (Vercammen-Grandjean & Nadchatram, 1963) collected from "R. geoffroyi zuluensis" [= R. clivosus zuluensis] from South Africa (Vercammen-Grandjean and Nadchatram, 1963; Stekolnikov, 2018a: 136). Stekolnikov (2018a: 189) also mentioned Willmannium natalense (Lawrence, 1949). DIPTERA: Streblidae: Ascodipteron lophotes Monticelli, 1898 from Eritrea (Haeselbarth et al., 1966: 106), South Africa (Haeselbarth et al., 1966: 106), Yemen (Haeselbarth et al., 1966: 106, host referred to as R. acrotis). Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Nycteribosca africana Walker, 1849 (Zumpt, 1950: 96 - on "R. geoffroyi". Raymondia hardyi Fiedler, 1954 from South Africa (Shapiro et al., 2016: 254). Raymondia intermedia Jobling, 1936 (Shapiro et al., 2016: 255). Raymondia waterstoni Jobling, 1931 (Zumpt, 1950: 96; Haeselbarth et al., 1966: 104). Brachytarsina (Brachytarsina) flavipennis Macquart, 1851 and Raymondia huberi Frauenfeld, 1855 from Algeria (Bendjeddou et al., 2017: 15). 285 Nycteribiidae: Nycteribia schmidlii Schiner, 1853 (Haeselbarth et al., 1966: 108; Bendjeddou et al., 2017: 15). Nycteribia scissa scissa Speiser, 1901 the nominate form found exclusively on R. clivosus from South Africa and Namibia (Haeselbarth et al., 1966: 108). Nycteribia ovalis Theodor, 1957 from Pietermaritzburg, South Africa (Haeselbarth et al., 1966: 110). Penicillidia fulvida (Bigot 1885) (Zumpt, 1950: 97 - on "R. geoffroyi"; Haeselbarth et al., 1966: 114). Nycteribia integra Theodor and Moscona, 1954 (Haeselbarth et al., 1966: 109, host referred to as R. acrotis). Nycteribia scissa Speiser, 1901 (Zumpt, 1950: 97 - on "R. geoffroyi") . Szentiványi et al. (2019: Suppl.) also mentions Penicillidia fulvida bat flies from Kenyan "Rhinolophus ferrumequinum keniensis" which were carrying Arthrorhynchus nycteribiae fungi. Calliphoridae: Zumpt (1950: 88) reported a calyptrate larva of Calliphora croceipalpis Jaennicke, 1867, which was found in decomposing bats ("Rhinolophus geoffroyi") on the bottom of the Sterkfontein caves (RSA). Other magots developed into an unidentified Borboridae and a Phoridae. SIPHONAPTERA Ischnopsyllidae: Rhinolophopsylla ashworthi (Waterston, 1913) known only from a few specimens from the Cape Province (King William’s Town, Willowmore and Oudtshoorn) (Haeselbarth et al., 1966: 191). Rhinolophopsylla capensis Jordan and Rothschild, 1921 (Zumpt, 1950: 97 on "R. geoffroyi"; Haeselbarth et al., 1966: 191, Beaucournu and Kock, 1996). ACARI Spinturnicidae: Spinturnix viduus and Periglischrus africanus were described by Zumpt (1950: 92, 93) from "Rhinolophus geoffroyi" from the Sterkfontein caves (RSA). Ixodidae: ?Ixodes simplex Neumann, 1906 (see Zumpt, 1950: 96); Ixodes pilosus howardi Neumann (on "Rhinolophus augur", see Howard, 1908: 168). Trombiculidae: Trombicula natalensis Lawrence, 1949 (see Zumpt, 1950: 96). Listrophoridae: Alabidocarpus nasicolus Lawrence, 1938 (see Zumpt, 1950: 96). VIRUSES: Bunyaviridae Phlebovirus Rift Valley Fever Virus (RVFV) - Oelofsen and Van der Ryst (1999) tested 40 individuals from eight localities in the Free State Province, South Africa for RVFV antigen using an enzyme linked immunosorbet assay (ELISA), none tested positive. 286 ISSN 1990-6471 Coronaviridae Anthony et al. (2017b: Suppl.) mention the following Alphacoronaviruses: Predict_CoV_70, Human_CoV_229E, Predict_CoV_65, and the Betacoronavirus: Predict_CoV_43 (subgenus Sarbecovirus; see Markotter et al., 2020: 6). Nziza et al. (2019: 157) found three new Coronaviruses in oral and rectal swaps from Rwandan bats: PREDICT_CoV-42, PREDICT_CoV-43 and PREDICT_CoV-44. Gabon, Kenya, Lesotho, Malawi, Mozambique, Namibia, Rwanda, Saudi Arabia, Somalia, South Africa, South Sudan, Sudan, Tanzania, Uganda, Zambia, Zimbabwe. Rhabdoviridae Lyssavirus - Rabies related viruses Rabies: Oelofsen and Smith (1993) tested 85 individual brains, from 14 localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Botswana, Burundi, Cameroon, Central African Republic, Congo (Democratic Republic of the), Egypt, Eritrea, Eswatini, Ethiopia, Figure 79. Distribution of Rhinolophus clivosus Rhinolophus cohenae Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 *2012. Rhinolophus cohenae Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, PLoS ONE, 7(9): 16. Publication date: 12 September 2012. Type locality: South Africa: Mpumalanga: Barberton: Mountainland Nature Reserve [25 43 08 S 31 15 58 E, 690m] [Goto Description]. Holotype: DNSM 8626: ad ♂, skull and alcoholic. Collected by: Lientjie Cohen (Mpumalanga Park). Presented/Donated by: ?: Collector Unknown. Baculum prepared. Skull and mandible in good condition (Taylor et al., 2012c: 16). Paratype: DNSM 11620: ad ♂, skull and alcoholic. Collected by: Dr Samantha Stoffberg and Lientjie Cohen; collection date: 27 September 2009. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 11918: ad ♂, skull and alcoholic. Collected by: Dr Samantha Stoffberg and Lientjie Cohen; collection date: 27 September 2009. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 11919: ad ♂, skull and alcoholic. Collected by: Dr Samantha Stoffberg and Lientjie Cohen; collection date: 27 September 2009. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 7886: ad ♂, skull and alcoholic. Collected by: Lientjie Cohen (Mpumalanga Park); collection date: 27 September 2004. Presented/Donated by: ?: Collector Unknown. Etymology: Recognition of Lientjie Cohen who collected the type specimen and contributed significantly to the conservation of bats in South Africa, particularly in Mpumalanga Province. The species name combines the surname Cohen and genitive singular caseending "ae" indicative of feminine gender (Taylor et al., 2012c: 17). - ZooBank: D0E1C9DD-7D36-4C01-AE0D-60AD474960A0. (Current Combination) TAXONOMY: Based on mitochondrial and nuclear DNA sequences and morphological and morphometric data, Taylor et al. (2012c) found that the Rhinolophus hildebrandtii group includes more species than previously recognized. COMMON NAMES: English: Cohen's Horseshoe Bat. Cohens Hufeisennase. German: CONSERVATION STATUS: Global Justification Cohen et al. (2017) report that this recently described species endemic to South Africa and known from the Mpumalanga Escarpment, and from a few records in the south east of Limpopo Province with an estimated extent of occurrence of 15,640 km2. There are inferred to be fewer than 1,000 mature individuals (and certainly fewer than 10,000) in the population. Colonies are usually small, numbering only a few individuals. The African Chiroptera Report 2020 287 (Driver et al., 2012), and alien invasive plant infestations are causing a decline in available habitat for foraging. Figure 80. Rhinolophus cohenae from Mpumalanga, South Africa. greatest number of mature individuals counted at a single site was ± 40. All recorded colonies are suspected to comprise the same subpopulation. An ongoing decline is inferred to be taking place as a result of loss of habitat due to poor land-use management practices, mining activities, agricultural intensification as well as infestation by alien invasive plant species. Further field surveys and vetting of museum records are needed to more accurately delimit the distribution range of the species. Currently, list the species as Vulnerable C2a(ii) and D1. Assessment History Global 2016: VU C2a(ii); D1 ver 3.1 (2001) (Cohen et al., 2017). Regional South Africa:- 2016: VU C2a(ii), D2 ver 3.1 (2001) (Cohen et al., 2016). MAJOR THREATS: Cohen et al. (2017) reports that the Mpumalanga Tourism and Parks Agency (MTPA) mapped all development applications received at a cadastral scale over a 14-year period (2000-2014), which showed that greatest pressure for land-use change has come from prospecting applications (54% of the land surface area) and mining (25% of land surface area) (Lötter et al., 2014). A major threat within this species’ range is mining (legal, illegal and recommissioning of old mines). Future developments at the above rates or even higher are likely to cause further detriment towards natural ecosystems and processes and in particular, disturb or destroy foraging grounds and roosting and maternity sites, or alter key microclimates needed by the species. Additionally, loss of natural habitat around roost sites through poor land-use management practices, (such as inappropriate burning regimes, overgrazing and alteration of vegetation structure negatively affect foraging areas and prey base), land development activities including agricultural intensification Climate change may also influence micro-climate distribution. This species is very dependent on suitable subterranean environments for roosting and maternity requirements and associated natural habitats for foraging. These sites are limited throughout its distribution range and beyond. The effects of climate change can severely impact on the survival of this species if the above is not provided for and not adequate for habitation any more (Cohen et al., 2017). CONSERVATION ACTIONS: Cohen et al. (2017) reports that the species occurs in the Mariepskop Primary Conservation Area and Barberton Mountainlands Nature Reserve. No specific conservation actions directed towards this species at the moment. The MTPA has developed the Mpumalanga Biodiversity Sector Plan (MBSP) that indicates areas of high conservation value and is based on a systematic conservation plan which considers the distribution of all species and their habitat, sets quantitative targets for these and tries to find the most sufficient selection of areas to meet these targets. A few R. cohenae localities fall within the boundaries of protected areas but most are situated on private land. The MBSP has categorised areas in term of. its biodiversity value and R. cohenae localities located within the Protected Area and Critical Biodiversity and Ecological Support Areas will potentially receive the best protection measures from a land development perspective where certain activities will not be allowed or be restricted. With regards to all other areas, the MBSP land-use guidelines should also be followed and Environmental Impact Assessment legislative tools applied. GENERAL DISTRIBUTION: Known only from three closely-spaced localities in Mpumalanga Province of South Africa in the vicinity of Nelspruit Taylor et al. (2012c: 17). Accurate delimitation of this species’ range is subject to further collecting and reappraisal of existing museum material, previously refered to as R. hildebrandtii. Native: South Africa (Taylor et al., 2012c: 17). ECHOLOCATION: Lowest recorded peak frequency 33.0 kHz in the holotype; mean 32.86 ± 0.24 kHz, n= 7 in type series Taylor et al. (2012c: 16). 288 ISSN 1990-6471 POPULATION: Structure and Density:- Cohen et al. (2017) report that In total, 240 individuals have been counted in surveys but this is an underestimate. The total population is thus inferred to be fewer than 1,000 mature individuals and thus certainly fewer than 10,000 mature individuals, as this species is encountered in small groups of which around 40 individuals was the highest number counted at a single site. All recorded colonies are suspected to be part of one subpopulation. DISTRIBUTION BASED ON VOUCHER SPECIMENS: South Africa. Trend:- Decreasing (Cohen et al., 2017). PARASITES: DIPTERA: Streblidae: Raymondia hardyi Fiedler, 1954 from South Africa (Shapiro et al., 2016: 254; based on a record by Theodor (1968), who mentioned "R. hildebrandtii" as host). ZOOBANK: D0E1C9DD-7D36-4C01-AE0D-60AD474960A0 Figure 81. Distribution of Rhinolophus cohenae Rhinolophus damarensis Roberts, 1946 *1946. Rhinolophus darlingi damarensis Roberts, Ann. Transv. Mus., 20 (4): 303. Type locality: Namibia: Damaraland: Okahandja district: Oserikari [Goto Description]. Holotype: TM 9474: ad ♀, skin and skull. Collected by: Austin Roberts; collection date: 10 August 1941. Note: Caught in bedroom at night. 2013. Rhinolophus d. damarensis: Jacobs, Babiker, Bastian, Kearney, van Eeden and Bishop, PLoS ONE, 8 (12): d82614 (Suppl.). Publication date: 3 December 2013. ? Rhinolophus damarensis: (Name Combination, Current Combination) TAXONOMY: morphology, apart from bacula, were highly convergent. Two lineages which within the R. damarensis clade can be seen: a northern (where the type of damarensis falls) and a southern linage, which diverged about 5 Mya (Jacobs et al., 2013a: 12). R. damarensis is embedded in the fumigatus clade that includes R. darlingi, R. fumigatus, R. eloquens and R. hildebrandtii (Jacobs et al. (2013a: 12). Figure 82. A male Rhinolophus damarensis (TM 48040) caught at Taung, North-West Province, South Africa. Jacobs et al. (2013a: 12) showed that R. darlingi (sensu lato) was polyphyletic comprising two nonsister lineages (R. darlingi (sensu stricto) and R. damarensis), with disjunct distributions in southern Africa, which diverged about 9.7 Mya. They further showed that despite the genetic divergence between the two lineages (R. darlingi (sensu stricto) and R. damarensis) echolocation frequencies as well as skull and post-cranial Based on DNA sequencing, morphometrics, and echolocation data, Jacobs et al. (2013a) concluded that the western population of R. darlingi represents a separate species, which was separated from the eastern population some 9.7 Mya. COMMON NAMES: German: Damara-Hufeisennase. African Chiroptera Report 2020 CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) [as part of R. darlingi] in view of its wide distribution, presumed large population, it occurs in a number of protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008aa; IUCN, 2009). While Monadjem et al. (2017bl) report that with an Extent of Occurrence of 17,3750 km 2 and an estimated population size of 20,000 individuals, this species qualifies as Least Concern; however, the population trend should be closely monitored since there is an ongoing threat of loss of roost sites, as old mines are being re-opened for mining. Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bl). 2008: LC ver 3.1 (2001) [as part of R. darlingi] (Jacobs et al., 2008aa; IUCN, 2009). 2004: LC ver 3.1 (2001) [as part of R. darlingi] (Jacobs et al., 2004v); IUCN, 2004). 1996: LR/lc [as part of R. darlingi] (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016i). 2004: NT ver 3.1 (2001) [as part of R. darlingi] (Friedmann and Daly, 2004). 289 a slight sexual dimorphism: in females the frequencies varied between 84.4 ± 0.7 kHz and 87.6 ± 1.1 kHz in the different localities, and in males between 84.4 kHz ± 0.9 to 86.9 ± 1.1 kHz. The only factor they found to be responsible for the divergence in resting frequency was annual mean temperature. MOLECULAR BIOLOGY: DNA - Jacobs et al. (2013a: 12) sequenced the Cytochrome b gene from the type specimen of R. d. damarensis Roberts, 1924 (TM 9474) (Genbank Nr- KF683224), which grouped distinctly from the eastern populations now referred to as R. darlingi (sensu stricto). Two lineages which within the R. damarensis clade can be seen a northern (where the type of damarensis falls, and a southern linage. POPULATION: Structure and Density:- It occurs in small colonies of less 100 individuals per colony (Monadjem et al., 2017bl). Trend:- Unknown (Monadjem et al., 2017bl). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Congo (Democratic Republic of the), Namibia, South Africa. MAJOR THREATS: The species is potentially threatened by the reopening of mine adits, as well as re-use of old mines (Monadjem et al., 2017bl). CONSERVATION ACTIONS: Monadjem et al. (2017bl) report that there are no direct conservation measures in place. It has however been recorded from formally protected Richtersveld National Park and the Augrabies Falls National Park. ECHOLOCATION: In South Africa, Maluleke et al. (2017: 7347) studied the variation in echolocation calls along a latitudinal gradient between 16 and 32° S. They observed resting frequencies of 85.43 ± 1.3 kHz and average call duration of 31.14 ± 5.9 msec, with Figure 83. Distribution of Rhinolophus damarensis Rhinolophus darlingi K. Andersen, 1905 *1905. Rhinolophus Darlingi K. Andersen, Ann. Mag. nat. Hist., ser. 7, 15 (85): 70. Publication date: 1 January 1905. Type locality: Zimbabwe: Mashonaland: Mazoe [17 30 S 31 03 E, 4 000 ft] [Goto Description]. Holotype: BMNH 1895.8.27.1: ad, skull only. Collected by: J.ff. Darling Esq. Collection date: 13 June 1895; original number: 35. Paratype: BMNH [a]:. Paratype: BMNH [b]:. Paratype: BMNH [c]:. Paratype: BMNH [d]:. Paratype: BMNH [e]:. - Etymology: In honour of J. ff. Darling, a mining engineer, who first collected this species in Mazoe, Zimbabwe, in the early part of the twentieth century (see Smithers, 1983: 126; Taylor, 2005). (Current Combination) 290 ISSN 1990-6471 1924. 2016. ? ? Rhinolophus darlingi barbertonensis Roberts, Ann. Transv. Mus., 10: 59. Publication date: 31 January 1924. Type locality: South Africa: SE Transvaal province: Barberton district: Louw's creek [Goto Description]. Holotype: TM 2476: ad ♀, skin and skull. Collected by: Austin Roberts; collection date: 14 March 1920. Note: Caught in old mine drive. Rhinolophus darling: Hoffmaster and Vonk, Behav. Sci., 6 (25): 1. Publication date: 20 November 2016. (Lapsus) Rhinolophus darlingi darlingi: (Name Combination) Rhinolophus darlingi: (Current Spelling) TAXONOMY: Jacobs et al. (2013a: 12) showed that R. darlingi (sensu lato) was polyphyletic, comprising two nonsister lineages (R. darlingi (sensu stricto) and R. damarensis), with disjunct distributions in southern Africa, which diverged about 9.7 Mya. They further showed that despite the genetic divergence between the two lineages (R. darlingi (sensu stricto) and R. damarensis) echolocation frequencies as well as skull and post-cranial morphology, except for the bacula, were highly convergent. R. darlingi is embedded in the fumigatus clade (Csorba et al., 2003: 39 - 40; Simmons (2005: 354) that includes – R. damarensis, R. fumigatus, R. eloquens and R. hildebrandtii (Jacobs et al. (2013a: 12). R. darlingi includes R. d. barbertonensis Roberts, 1924. COMMON NAMES: Afrikaans: Darling se saalneusvlermuis, Darlingsaalneusvlermuis. Czech: vrápenec Darlingův. English: Darling's Horseshoe Bat. French: Fer-àCheval de Darling, Rhinolophe de Darling. German: Darlings Hufeisennase. Portuguese: Morcego ferradura de Darling. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: R. darlingi is represented intermittently in the upper Pleistocene and in the very late Holocene at Border Cave (Avery, 1991: 6). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in a number of protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008aa; IUCN, 2009; Monadjem et al., 2017cv). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cv). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008aa; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004v); IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016j). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Jacobs et al. (2008aa) [in IUCN (2009)] and Monadjem et al. (2017cv) reported that there appear to be no major threats to this species as a whole. CONSERVATION ACTIONS: This species is present within several protected areas. Further taxonomic studies are needed for populations recorded outside of southern Africa. No direct conservation measures are currently needed for the species (Jacobs et al., 2008aa; IUCN, 2009; Monadjem et al., 2017cv). GENERAL DISTRIBUTION: Rhinolophus darlingi is largely distributed in southern Africa with some additional records outside of this area. Skinner and Chimimba (2005) report that it has been recorded from Namibia and northeastern Botswana; that it is widely distributed in Zimbabwe; in South Africa, where its distribution is best predicted by geology (Babiker Salata, 2012: 50), it is known from Limpopo Province, eastern Mpumalanga, northern Gauteng, in parts of the Eastern Cape and the Northern Cape, and from KwaZulu-Natal. In much of eastern Mozambique; and in the lowveld and Lubombo regions of Swaziland. It is unclear if the species is present in Lesotho. Outside of southern Africa, it has been recorded from Benguela in Angola, Banagi in Tanzania, and possibly from Nigeria suggesting a wider distribution than is currently known (Skinner and Chimimba, 2005). Herkt et al. (2017: Appendix S9), however, found that there are substantial differences in terms of occupied ecological niches between the populations from southern Africa and those from Nigeria, which support the statement in IUCN (2013) that they may represent a separate species. African Chiroptera Report 2020 For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,217,047 km2. CAS has 5 specimens from the Pyramids of Giza, Egypt, which may be possible misidentifications as these seem to be well outside of the distribution range of the species. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 557); Botswana (Monadjem et al., 2010d: 557); Burundi; Congo (The Democratic Republic of the); Kenya; Malawi (Happold et al., 1988; Monadjem et al., 2010d: 557); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 557); Namibia (Monadjem et al., 2010d: 557); South Africa (Monadjem et al., 2010d: 557); Swaziland (Monadjem et al., 2010d: 558); Tanzania; Zambia (Cotterill, 2004a: 261; Monadjem et al., 2010d: 558); Zimbabwe (Monadjem et al., 2010d: 558). Presence uncertain: Lesotho; Nigeria. ECHOLOCATION: Jacobs et al. (2007a) reported a peak frequency of 88.1 (± 2.0) kHz for individuals from South Africa (detector type not reported). For individuals from Swaziland Taylor (1999b) using a Petterssen D980 at Mlawula reported a maximum frequency of 86.2 (± 0.1) kHz, while Monadjem et al. (2007a) using an Anabat detector at a number of localities reported a maximum frequency of 85.8 (± 0.36) kHz. Taylor (2000) reported a constant frequency component of 86 kHz. Monadjem et al. (2017c: 179) registered the following parameters: Fmin: 74.3 ± 4.28 (70.8 - 79.1) kHz, Fknee: 85.7 ± 0.46 (85.2 - 86.1) kHz, Fc: 85.6 ± 0.38 (85.2 - 85.8) kHz and duration: 26.5 ± 3.74 (22.6 - 30.0) msec. They also indicated that the maximum detection distance was 2 m. One call from Waterberg, South Africa had the following parameters: Fmax: 87.4 kHz, Fmin: 84.2 kHz, Fknee: 85.6 kHz, Fchar: 85.1 kHz, duration: 28.0 msec (Taylor et al., 2013b: 19). These authors also reported an additional four calls from Swaziland: Fmax: 85.8 ± 0.25 (85.5 - 86.0) kHz, Fmin: 74.5 ± 4.54 (72.3 - 76.1) kHz, Fknee: 84.8 ± 0.52 (84.1 - 85.2) kHz, Fchar: 85.2 ± 0.54 (84.5 85.6) kHz, duration: 17.8 ± 3.39 (13.3 - 21.5) msec. Jacobs (2016: 119) indicates that the second harmonic of this bat coincides with the fourth harmonic of R. hildebrandtii, which possibly might illustrate "harmonic hopping". MOLECULAR BIOLOGY: DNA - Jacobs et al. (2013a: 12) sequenced the Cytochrome b gene from the type specimen of R. 291 d. barbetonensis Roberts 1924 (TM 2476) (Genbank Nr- KF683217), which grouped with specimens collected from the type locality of the type of R. darlingi (Mazoe, Zimbabwe). Karyotype - Both Peterson and Nagorsen (1975) and Rautenbach (1986) reported, for specimens from Zimbabwe and South Africa respectively, 2n = 58, FN = 60, BA = 4, and subtelocentric X and Y chromosomes. Erasmus and Rautenbach (1984: 94) mention 52 acrocentric and four bi-armed autosomal chromosomes, resulting in 60 autosomal arms. Protein / allozyme - Unknown. POPULATION: Structure and Density:- Rhinolophus darlingi is a locally common species that, while usually found in small numbers, can be represented by hundreds of bats in a colony (Skinner and Chimimba, 2005) (Jacobs et al., 2008aa; IUCN, 2009; Monadjem et al., 2017cv). Trend:- 2016: Unknown (Monadjem et al., 2017cv). 2008: Unknown (Jacobs et al., 2008aa; IUCN, 2009). POSTNATAL DEVELOPMENT: Hoffmaster and Vonk (2016: 1) refer to Carter and Wilkinson (2013), who suggest that R. darlingi mothers, cooperating to keep all of the offspring warm, increase the probability of their own offspring’s survival. PARASITES: Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Nycteribiidae: Nycteribia scissa rhodesiensis Theodor, 1957 from Zimbabwe and Malawi (Haeselbarth et al., 1966: 108). Nycteribia tecta Theodor,1957 from Zimbabwe and South Africa, but Haeselbarth et al. (1966: 110) suggests these may represent a distinct subspecies. SIPHONAPTERA Pulicidae: Echidnophaga aethiops Jordan and Rothschild, 1906 locality not mentioned (Haeselbarth et al., 1966: 135). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV: Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in one individual from Limpopo Province, South Africa, none were tested positive (0/1). 292 ISSN 1990-6471 Rhabdoviridae Lyssavirus - Rabies related viruses Rabies: Oelofsen and Smith (1993) tested five individuals from three localities using "Trousse Platelia Rage" ELISA kit (Diagnostic Pasteur), none tested positive for antibodies to rabies virus glycoprotein G. Oelofsen and Smith (1993) tested five individual brains, from three localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Eswatini, Kenya, Lesotho, Malawi, Mozambique, Namibia, Nigeria, South Africa, Tanzania, Togo, Uganda, Zambia, Zimbabwe. Figure 84. Distribution of Rhinolophus darlingi Rhinolophus deckenii Peters, 1868 *1868. Rhinolophus Deckenii Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 705 (for 1867). Type locality: Tanzania: Zanzibar coast (mainland opposite Zanzibar) [Goto Description]. Holotype: ZMB 3269: ad ♀, alcoholic (skull missing). Collected by: Baron Karl Klaus von der Decken; collection date: undated. - Etymology: In honour of baron Karl Klaus Von der Decken (1833 - 1865), collector of the type specimen. (Current Combination) ? Rhinolophus deckeni: (Alternate Spelling) ? Rhinolophus deckenii: (Current Spelling) TAXONOMY: Treated as a subspecies of clivosus by Hayman and Hill (1971: 23); but see Koopman (1975: 386). ferrumequinum species group (Csorba et al., 2003: 41 - 42; Simmons, 2005: 354). See Csorba et al. (2003: 41) for remarks on taxonomy. Assessment History Global 2008: NT ver 3.1 (2001) (Jacobs et al., 2008ab; IUCN, 2009). 2004: DD ver 3.1 (2001) (Jacobs et al., 2004p; IUCN, 2004). 1996: DD (Baillie and Groombridge, 1996). COMMON NAMES: Chinese: 德 氏 菊 头 蝠 . Czech: vrápenec východoafrický. English: Decken's Horseshoe Bat, Eastern Africa horseshoe bat. French: Ferà-Cheval d'Afrique orientale, Rhinolophe de Decken, Rhinolophe d'Afrique orientale. German: Deckens Hufeisennase. Italian: Rinòlofo di Dècken, Fèrro di cavàllo di Dècken. Regional None known. CONSERVATION STATUS: Global Justification Listed as Near Threatened (NT ver 3.1 (2001)) because this species is probably in significant decline (but probably at a rate of less than, though close to, 30 % over ten years) because of deforestation through much of its range, thus making the species close to qualifying for Vulnerable A2c (Jacobs et al., 2008ab; IUCN, 2009). CONSERVATION ACTIONS: There appear to be no direct conservation measures in place. It has been recorded from some National Parks and Forest Reserves in Tanzania. There is a need to maintain areas of suitable forest habitat for this species. Further research is needed into the species taxonomy, biology and overall natural history (Jacobs et al., 2008ab; IUCN, 2009). MAJOR THREATS: Jacobs et al. (2008ab) [in IUCN (2009)] reported that this species is presumably threatened by the logging and conversion of forest land to agricultural use, especially in coastal areas. GENERAL DISTRIBUTION: Rhinolophus deckenii, a little-known East African species has been recorded from Uganda, Kenya, African Chiroptera Report 2020 293 and Tanzania (including the islands of Pemba, and Unguja [=Zanzibar] [see O'Brien, 2011: 286]). It has recently been found in Mozambique. Nycteribiidae: Nycteribia tecta Theodor 1957 from Kenya, Uganda, Tanzania and the Sudan (Haeselbarth et al., 1966: 110). Native: Kenya; Mozambique (Monadjem et al., 2010d: 558; Monadjem et al., 2010c: 378); Tanzania (including the islands of Zanzibar and Pemba) (Stanley and Goodman, 2011: 42); Uganda. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, Malawi, Mozambique, Tanzania. ECHOLOCATION: Monadjem et al. (2010c: 378) reported from Mozambique that the peak echolocation frequency of a single male was recorded at 72 kHz (ANABAT). POPULATION: Structure and density:- Small colonies of less than 20 individuals. It is reported to be locally common in some areas, such as the Manga Forest Reserve, Tanzania (Doggart et al., 1999b) (Jacobs et al., 2008ab; IUCN, 2009). Trend:- 2008: Decreasing (Jacobs et al., 2008ab; IUCN, 2009). Figure 85. Distribution of Rhinolophus deckenii PARASITES: Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Rhinolophus denti Thomas, 1904 *1904. Rhinolophus Denti Thomas, Ann. Mag. nat. Hist., ser. 7, 13 (77): 386. Publication date: 1 May 1904. Type locality: South Africa: Northern Cape province: [formerly Bechuanaland]: Kuruman [26 28 S 22 27 E, 1 300 m] [Goto Description]. Holotype: BMNH 1904.4.8.2: ♂. Collected by: Captain R.E. Dent; collection date: 24 January 1904; original number: 7. - Etymology: In honour of R.E. Dent (Smithers, 1983: 130; Skinner and Smithers, 1990; Taylor (2005), the collector of the type specimen. (Current Combination) 1960. Rhinolophus denti knorri Eisentraut, Stuttg. Beitr. Naturk., 39: 3. Publication date: 1 June 1960. Type locality: Guinea: 12 km W Kolenté, Salung Plateau: Nyembaro [ca. 10 05 N 12 30 W] [Goto Description]. Holotype: SMNS 6111: ad ♂, skin and skull. Collected by: Hans Knorr; collection date: 27 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 293) mention the locality for this specimen as "SalungPlateau near Nyembaro-Nerebili, 10 km western of Kolenté, 400 m, Guinea". Paratype: SMNS 6112: ad ♀, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Paratype: SMNS 6112a: ad ♀, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Paratype: ZFMK MAM-1959.0174: ♂, skin and skull. Collected by: Hans Knorr; collection date: 15 November 1956; original number: 90. See Hutterer and Peters (2010: 11). Paratype: ZFMK MAM-1959.0175: ♂, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. See Hutterer (1984: 30), Hutterer and Peters (2010: 11).. - Comments: Considered a valid subspecies by Grubb et al. (1998: 78). Type specimen in the SMNS. ? Rhinolophus denti denti: (Name Combination) ? Rhinolophus denti: (Current Spelling) 294 ISSN 1990-6471 TAXONOMY: because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008ak; IUCN, 2009; Monadjem et al., 2017bd). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bd). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008ak; IUCN, 2009). 2004: DD ver 3.1 (2001) (Jacobs et al., 2004ak; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Figure 86. A female Rhinolophus denti (TM 48035) caught at Taung, North-West Province, South Africa. Placed within the capensis-group (Csorba et al., 2003: 9 - 10; Simmons, 2005: 354). Possibly includes R. swinnyi (see Roberts, 1914; Csorba et al., 2003: 9 - 10). Two subspecies are currently recognised, besides the nominate form described from the Northern Cape Province of South Africa (and including Namibia, Botswana and Zimbabwe), also R. d. knorri Eisentraut, 1960 from Guinea (and including Ghana) (Csorba et al., 2003: 9 - 10; Simmons, 2005: 354). R. d. knorri is slightly smaller than the nominate form especially in the length of tibia and ear (Csorba et al., 2003: 9 - 10). See Csorba et al. (2003: 9 - 10) for further remarks on taxonomy. COMMON NAMES: Afrikaans: Dent se saalneusvlermuis, Dentsaalneusvlermuis, Dentse Vlermuis. Czech: vrápenec Dentův. English: Dent's Horseshoe Bat. French: Rhinolophe de Dent. German: Dents Hufeisennase. Portuguese: Morcego ferradura de Dent. SIMILAR SPECIES: R. swinnyi (Shortridge, 1934; Roberts, 1914; 1946: 304). Shortridge (1934) was incorrect though, in indicating that Hewitt (1913) described a new subspecies of R. denti, from Pirie in the Eastern Cape province of South Africa, when it was actually R. swinnyi piriensis (Hewitt, 1913: 402). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Avery and Avery (2011: 16) indicate that material from the eastern part of the Northern Cape province in South Africa, that was previously assigned to R. capensis should probably be assigned to R. denti. CONSERVATION STATUS: Global Justification Although this species is known mainly from isolated records from a large area, it is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and Regional South Africa:- 2016: NT D1 ver 3.1 (2001) (Schoeman et al., 2016a). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: Jacobs et al. (2008ak) [in IUCN (2009)] and Monadjem et al. (2017bd) reported there appear to be no major threats to this species. It might be locally threatened in parts of its range by disturbance to roosting caves. CONSERVATION ACTIONS: There appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further studies are needed into the distribution of this species (particularly in West and Central Africa) (Jacobs et al., 2008ak; IUCN, 2009; Monadjem et al., 2017bd). GENERAL DISTRIBUTION: Rhinolophus denti is widely, but patchily, recorded in West and southern Africa. It ranges from southeastern Senegal, through northern parts of West Africa to northeastern Ghana; in Central Africa it appears to have only been recorded from eastern Congo and southern Angola; in southern Africa, it is present in Namibia, northwestern and southwestern Botswana and northern parts of South Africa, where the species' distribution is linked with land use/land cover (Babiker Salata, 2012: 50). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 781,091 km2. A record from Gambia is doubtful according to Kock et al. (2002: 85), since it is possibly of an adolescent R. landeri, as is the record from Côte d'Ivoire (Csorba et al., 2003: 10) that is also apparently a R. landeri (see Simmons, 2005: 354). Records have been reported from Zimbabwe (Smithers and Wilson, 1979: 57; Fenton and Bell, 1981). However, Smithers (1983) later indicated African Chiroptera Report 2020 three records from northeastern Zimbabwe in Smithers and Wilson (1979) and Csorba et al. (2003: 10) are misidentifications. Herkt et al. (2017: Appendix S9) found that there are major differences between the occupied ecological niches for denti from southwestern Africa and knorri from West Africa. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 558); Botswana (Smithers, 1971; Csorba et al., 2003: 10; Monadjem et al., 2010d: 558); Ghana (Csorba et al., 2003: 10); Guinea (Csorba et al., 2003: 10); Guinea-Bissau (Rainho and Ranco, 2001: 48; Simmons, 2005: 354); Mozambique (Smithers and Lobão Tello, 1976; Csorba et al., 2003: 10; Smithers and Lobão Tello, 1976); Namibia (Ellerman et al., 1953; Kock and Howell, 1988; Csorba et al., 2003: 10; Monadjem et al., 2010d: 558); Senegal; South Africa (Northern Cape Province (Ellerman et al., 1953; Herselman and Norton, 1985; Rautenbach, 1986; Monadjem et al., 2010d: 558; Csorba et al., 2003: 10), North West Province (James, 1986; Csorba et al., 2003: 10), Free State Province (Watson, 1998; Monadjem et al., 2010d: 558)). Bates et al. (2013: 338) reject the presence of R. denti in the Republic of Congo. GEOGRAPHIC VARIATION: See "Taxonomy" section. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Smithers (1983), Skinner and Smithers (1990), Taylor (2000), Csorba et al. (2003: 9 - 10), Taylor (2005). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Smithers (1983), Skinner and Smithers (1990), Taylor (2000), Csorba et al. (2003: 9 - 10), Taylor (2005). DETAILED MORPHOLOGY: Baculum - Unknown Of the skull, teeth and noseleaves see Csorba et al. (2003: 9 - 10). SEXUAL DIMORPHISM: Five mensural characters (forearm length, fourth metacarpal length, width across the upper canines, rostral width, and length of the mandibular toothrow) showed significant variation between the sexes in an analysis of specimens from Koegelbeen Cave in the Northern Cape Province of South Africa (Rautenbach, 1986). 295 ECHOLOCATION: For individuals at Sengwa in Zimbabwe Jacobs (1996) and Fenton and Bell (1981) reported a maximum frequency of 110 kHz. Jacobs et al. (2007a) reported a similar peak frequency of 110.9 (± 1.7) kHz for individuals in South Africa. Jacobs (2016: 119) indicates that the second harmonic of this bat coincides with the fourth harmonic of R. capensis, which possibly might illustrate "harmonic hopping". MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Peterson and Nagorsen (1975) reported the diploid number as 2n = 58, aFN = 62, BA = 6, X = ST, Y =?, based on specimens from Zimbabwe. These results were supported by Rautenbach (1986) who also reported the Y chromosome to be metacentric. Karyologically identical to R. swinnyi (Csorba et al. (2003: 9 - 10). Protein / allozyme - Unknown. HABITAT: Found in relatively arid regions (Csorba et al., 2003: 9 - 10). Even the most south easterly record in Africa comes from the drier southwestern part of the Free State Province of South Africa (Watson, 1998). HABITS: In north western Namibia a specimen shot by Shortridge (1934: 52) at "Rua Cana Falls" had a slow fluttering Kerivoula-like flight, and several other individuals were observed "circling low among trees and bushes near the Cunene River on bright moonlight nights". Found roosting at Drotsky's Cave in Namibia throughout the year (Smithers, 1983; Skinner and Smithers, 1990; Taylor, 2005). ROOST: Shortridge (1934: 52) records finding a single individual, at Louisvale in the Northern Cape Province of South Africa, inside a low water culvert occupied by a small colony of Nycteris. Shortridge, 1934: 52) also records finding two individuals "hanging from the thatched roof inside a small native house at Kaoko-Otavi" in the north western part of Namibia. Also found in caves (Smithers, 1983; Skinner and Smithers, 1990; Taylor, 2005). At Drotsky's Cave in Namibia the bats were "clinging to the sides of stalactites…hanging freely from the sides in open clusters", and were found in semi-dark 90 m from the cave entrance and in total darkness deep in the cave (Smithers, 1983; Skinner and Smithers, 1990; Taylor, 2005). Notes on an index card for specimens in the Transvaal Museum, from a cave near Warrenton in the Northern Cape Province of 296 ISSN 1990-6471 South Africa, indicate the cave was wet with water on the floor and dripping from stalactites, and that the bats congregated in separate groups. James (1986: 218) recorded a single individual at a disused quarry near Buxton in the North West Province, roosting on the ceiling of a "ruined mine structure which formed a 2 x 2 x 20 m cement, semi-dark room in a hillside". Weber and Fahr (2006: 4) indicate that, in Guinea, R. denti largely or exclusively depends on the availability of caves as day roosts. MIGRATION: May not migrate as R. denti has been found roosting at Drotsky's Cave in Namibia throughout the year (Smithers, 1983; Skinner and Smithers, 1990; Taylor, 2005). DIET: Insectivorous (Smithers, 1983; Skinner Smithers, 1990; Taylor, 2000; 2005). ACTIVITY AND BEHAVIOUR: Notes on an index card for specimens in the Transvaal Museum, from a cave near Warrenton in the Northern Cape Province of South Africa, indicate that on 14th of June 1983 the bats were in torpor. PARASITES: Brennan et al. (2015: 145) found no antibodies against Pseudogymnoascus destructans (Whitenose syndrome) in blood samples from bats from Botswana. VIRUSES: Paramyxoviridae Mortlock et al. (2015: 1841) reported that two out of three examined South African R. denti specimens tested positive for Paramyxovirus sequences. and POPULATION: Structure The species is known from fewer than 100 in colonies in West Africa, and fewer than 200 colonies in southern Africa (Jacobs et al., 2008ak; IUCN, 2009; Monadjem et al., 2017bd). Density "Sparsely distributed", with no records of large colonies (Shortridge, 1934:52). At Drotsky's Cave in Namibia the bats were observed "clinging to the sides of stalactites in dozens" (Smithers, 1983; Skinner and Smithers, 1990; Taylor, 2005). Herselman and Norton (1985: 92) indicated that this species had not been collected from the Northern Cape province in the "last half century" and could be considered as "extremely rare, possibly extinct, in the Cape Province". Yet, specimens in the Transvaal Museum, collected from the Northern Cape Province of South Africa between 1983 and 1988, indicate that at three different visits; 12, 71 and 15 specimens were taken from Koegelbeen Cave, 12 specimens were taken on a single date from a cave near Warrenton, and 11 specimens were taken on a single date from a locality near Postmasberg. Rhabdoviridae Lyssavirus - Rabies related viruses Rabies: Oelofsen and Smith (1993) tested 12 individuals from three localities using "Trousse Platelia Rage" ELISA kit (Diagnostic Pasteur), none tested positive for antibodies to rabies virus glycoprotein G. Oelofsen and Smith (1993) tested 15 individual brains, from two localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Botswana, Cameroon, Congo (Democratic Republic of the), Gabon, Namibia, South Africa. Trend 2016: Unknown (Monadjem et al., 2017bd). 2008: Unkown (Jacobs et al., 2008ak; IUCN, 2009). Figure 87. Distribution of Rhinolophus denti Rhinolophus eloquens K. Andersen, 1905 *1905. Rhinolophus Hildebrandti eloquens K. Andersen, Ann. Mag. nat. Hist., ser. 7, 15 (85): 74. Publication date: 1 January 1905. Type locality: Uganda: Entebbe [00 04 N 32 28 E, 3 African Chiroptera Report 2020 1922. 2016. ? ? ? ? 297 800 ft] [Goto Description]. Holotype: BMNH 1899.8.4.4: ad, skull only. Collected by: Sir Frederic John Jackson. See Andersen (1905a: 75). Topotype: ROM 68683: ad ♀, alcoholic (skull not removed). Collected by: Theo Simpson Jones; collection date: 4 July 1963. Presented/Donated by: ?: Collector Unknown. - Etymology: From the Latin masculine adjective èloquens. Andersen (1905a) provides no etymological information; however, it is possible that this Rhinolophus is "eloquent" from a phylogenetic and zoogeographical point of view, as Andersen (1905a: 75) writes that "The present form is of great interest from phylogenetic no less than from a zoogeographical point of view" (see Lanza et al., 2015: 93). Rhinolophus hildebrandti perauritus de Beaux, Atti Soc. ital. Sci. nat., 61: 22. Publication date: February 1922. Type locality: Somalia: Territory of the Rahanuin [Goto Description]. Rhinolophus eloquensis: Amori, Segniagbeto, Decher, Assou, Gippoliti and Luiselli, Zoosystema, 38 (2): 219. Publication date: 24 June 2016. (Lapsus) Rhinolophus aethiops eloquens: (Name Combination) Rhinolophus eloquens eloquens: (Name Combination) Rhinolophus eloquens: (Name Combination, Current Combination) Rhinolophus fumigatus eloquens: (Name Combination) TAXONOMY: Included as subspecies in aethiops by Kock (1969a: 121). Includes perauritus; see Koopman (1975: 389). fumigatus species group (Csorba et al., 2003: 47 - 48; Simmons, 2005: 354). See Csorba et al. (2003: 48) for remarks on taxonomy. COMMON NAMES: Chinese: 乌 干 达 菊 头 蝠 . Czech: vrápenec ugandský. English: Eloquent Horseshoe Bat. French: Rhinolophe éloquent. German: Andersens Hufeisennase. Italian: Rinòlofo eloquènte, Fèrro di cavàllo eloquènte. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008ac; IUCN, 2009; Monadjem et al., 2017cj). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cj). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008ac; IUCN, 2009). 2004: DD ver 3.1 (2001) (Jacobs et al., 2004u; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Rhinolophus eloquens is threatened by the general conversion of its habitat to agricultural use (Jacobs et al., 2008ac; IUCN, 2009; Monadjem et al., 2017cj). CONSERVATION ACTIONS: Jacobs et al. (2008ac) [in IUCN (2009)] and Monadjem et al. (2017cj) reported there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further research needed into the distribution, natural history and threats to this bat. GENERAL DISTRIBUTION: Rhinolophus eloquens is endemic to Central and Eastern Africa. It has been recorded from southern Sudan, eastern Democratic Republic of the Congo, Rwanda, Uganda, Kenya, Somalia and the islands of Pemba, Unguja [=Zanzibar], and Mafia in Tanzania (see O'Brien, 2011: 286). Native: Congo, (Democratic Republic of) (Hayman et al., 1966; Monadjem et al., 2010d: 558); Ethiopia (Lavrenchenko et al., 2004b: 147); Kenya; Rwanda; Somalia; Sudan; Tanzania (United Republic of); Uganda. POPULATION: Structure and Density:- Little information is available on the population abundance or size of this species (Jacobs et al., 2008ac; IUCN, 2009; Monadjem et al., 2017cj). Trend:- 2016: Unknown (Monadjem et al., 2017cj). 2008: Unknown (Jacobs et al., 2008ac; IUCN, 2009). PARASITES: HEMIPTERA: Polyctenidae: Androctenes horvathi Jordan 1912 recorded from Kenya and Sudan (Haeselbarth et al., 1966: 17). 298 ISSN 1990-6471 DIPTERA: Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in subsaharan Africa (Haeselbarth et al., 1966: 100). Jobling (1954) as mentions Raymondia intermedia Jobling, 1936 as possible parasite of R. eloquens (see also Shapiro et al., 2016: 255). Raymondia seminuda Jobling, 1954 (Shapiro et al., 2016: 255). Nycteribiidae: Nycteribia hoogstraali Theodor, 1957 from the Torit district in the Sudan (Zumpt 1966: 109). Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114). Mount Elgon bat virus is reported by Luis et al. (2013: suppl.) although they remark that the bat might have been a R. hildebrandti. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Ethiopia, Kenya, Rwanda, South Sudan, Sudan, Tanzania, Uganda. ACARI: Trombiculidae: Stekolnikov (2018a: 50) reported the presence of Whartonia oweni VercammenGrandjean and Brennan, 1957. VIRUSES: Filoviridae - Filoviruses Marburgvirus Kuzmin et al. (2010b: 353) detected MARV RNA in 3.6 % of the examined R. eloquens in the DRC. Luis et al. (2013: suppl.) report the presence of Lake Victoria marburg virus. Rhabdoviridae Vesiculovirus Figure 88. Distribution of Rhinolophus eloquens Rhinolophus euryale Blasius, 1853 *1853. Rhinolophus euryale Blasius, Arch. Naturgesch. Berlin, 19 (1): 49, 52, 54. Publication date: 1853. Type locality: Italy: Milan. - Etymology: Referring to "Euryale", one of the Gorgons (see Palmer, 1904: 279). (Current Combination) 1867. Rhinolophus algirus Loche, Exploration scientifique de l'Algerie pendant les années 1840, 1841, 1842. Mammifères, Cheiroptères, 83. Publication date: 1867. Type locality: Algeria: "Jardin Marengo". - Comments: Allen (1939a: 74) states that this name is not certainly identifiable, but may possible prove to supersede barbarus if the types can be found. Ellerman and Morrison-Scott (1951: 120) include it with a question mark in the synonymy of R. euryale barbarus. 1896. Hipposiderus euryale: Thomas, Ann. Mus. civ. Stor. nat. Genova, (2) 17 (37): 105. (Name Combination) 1904. E[uryalus] atlanticus K. Andersen and Matschie, Sber. Ges. naturf. Freunde Berlin, 77. Type locality: France: Indre-et-Loire: St. Paterna. Holotype: ZMB 12963: ♂, alcoholic (skull not removed). Collected by: Nitsche; collection date: 6 April 1893; original number: (formerly ZMB 6705). See Turni and Kock (2008: 28). Paratype: MNHN ZM-MO-18792030: alcoholic (skull not removed). Collected by: The Lataste collection. See Rode (1941: 238). 1904. E[uryalus] Cabreræ K. Andersen and Matschie, Sber. Ges. naturf. Freunde Berlin, 78. Type locality: Spain: Madrid: Alcata de Henares [40 25 N 03 43 W]. Holotype: ZMB 12964: ♀, alcoholic (skull not removed). Original number: ex. Museum Madrid. See Turni and Kock (2008: 28). - Comments: Mentioned as cabrerai by Trouessart, 1910 (see Cabrera, 1914: 83). 1970. R[hinolophus] euryole: Rushton, Ann. Génét. Sél. anim., 2 (4): 463. (Lapsus) 1970. Rhinolophus euroyle: Rushton, Ann. Génét. Sél. anim., 2 (4): 466. (Lapsus) TAXONOMY: Revised by DeBlase (1972). Gaisler (2001: 68) does not recognises subspecies. euryale species group (Csorba et al., 2003: 14 - 16; Simmons, 2005: 354 - 355). See Csorba et al. (2003: 16) for remarks on taxonomy. African Chiroptera Report 2020 COMMON NAMES: Albanian: Lakuriq nate hundëpatkua i Mesdheut. Arabian: Khaffash. Armenian: Հարավային պայտաքիթ. Azerbaijani: Cənub nalburunu. Basque: Ferra-saguzar mediterraneo. Belarusian: Падкаванос міжземнаморскі. Bosnian: Mediteranski potkovasti šišmiš. Breton: Frigribell greizdouarel. Bulgarian: Южен подковонос. Castilian (Spain): Murciélago mediterráneo de herradura, Rinolofo mediterraneao de herradura, Murciélago de herradura mediterráneo. Catalan (Spain): Ratpenat de ferradura mediterrani, Rat penat mediterrani de ferradura, Ratapinyada mediterrània de ferradura. Croatian: Južni potkovnjak, Mediteranski potkovasti šišmiš. Czech: Vrápenec jižní, Vrápenec středozemský. Danish: Middelhavehestekonæse. Dutch: Paarse hoefijzerneus. English: Mediterranean Horseshoe Bat. Estonian: Vahemere sagarnina. Finnish: Välimerenherkko. French: Rhinolophe euryale. Frisian: Pearse hoefizernoas. Galician (Spain): Morcego de ferradura mediterráneo, Morcego mediterráneo de ferradura. Georgian: სამხრეთული ცხვირნალა. German: Mittelmeerhufeisennase, Mittelmeer-Hufeisennase, Gleichsatteligen Kammnase. Greek: Μεσορινόλοφος. Hebrew: ‫בהיר פרסף‬, Parsaf Bahir. Hungarian: Kereknyergű patkósdenevér, Kereknyergû patkósorrú denevér, Rinolophos i mesogiaki. Italian: Rinolofo mediterraneo, Rinolofo Eurìale, Ferro di cavallo Eurìale, ferro di cavallo mediterraneo. Irish Gaelic: Crú-ialtóg Mheánmhuirí. Latvian: Vidusjuras pakavdegunis. Lithuanian: Pietinis pasagnosis. Luxembourgish: Mëttelmier-Huffeisennues. Macedonian: Медитерански потковичар, Juzhen Potkovichar. Maltese: Rinolofu tal-Mediterran. Montenegrin: Južni potkovičar. Norwegian: Middelhavshesteskonese. Polish: Podkowiec śródziem-nomorski. Portuguese: Morcego-de-ferraduramediterrânico. Rhaeto-Romance: Rinolof mediterran. Romanian: Liliacul mediteranean cu potcoavă, Liliac-sudic. Russian: Подковонос южный, Южный подковонос [= Yuzhnyj podkovonos]. Serbian: Средоземни потковичар [= Sredozemni potkovičar]. Scottish Gaelic: Crudh-ialtag Meadhan-thìreach. Slovak: Podkovár južný. Slovenian: Južni podkovnjak. Swedish: Medelhavshästskonäsa. Turkish: Akdeniz Nalburunlu Yarasası. Ukrainian: Підковик (Підковоніс) південний. Welsh: Ystlum pedol Môr y Canoldir. SIMILAR SPECIES: Puechmaille and Teeling (2013: 252) point out that visual identification of hibernating bats might be problematic in the case of R. euryale and R. mehelyi, as bat they originally identified as R. 299 mehelyi turned out to belong to the other species after using non-invasive genetic tests on the animals' droppings. CONSERVATION STATUS: Global Justification Many range states have reported that populations have declined, and colonies have disappeared over the last 27 years (=3 generations). It is inferred that overall population decline has approached 30 % over that period (although the population is now stable and or even increasing in some areas, e.g. France), so the species is assessed as Near Threatened (approaching A2c) (NT ver 3.1 (2001)) (Hutson et al., 2008o; IUCN, 2009; Juste and Alcaldé, 2016a). Assessment History Global 2016: NT ver 3.1 (2001) (Juste and Alcaldé, 2016a). 2008: NT ver 3.1 (2001) (Hutson et al., 2008o; IUCN, 2009). 1996: VU A2c ver 2.3 (1994). Regional None known. MAJOR THREATS: Threats include loss of foraging habitat, and disturbance and loss of underground habitats. On a landscape scale, fragmentation and loss of linear elements such as hedgerows and riparian vegetation is a problem because such elements are used as landscape references for commuting. The species' strong dependence upon caves for roosting makes it particularly sensitive to cave disturbance, such as that from caving or tourism. Tourist disturbance of caves affects the species in several range states. The use of organochlorine pesticides is believed to have contributed to the earlier dramatic decline of the species in France (Brosset et al., 1988). In North Africa, threats include habitat loss due to agriculture (livestock) and human disturbance (Hutson et al., 2008o; IUCN, 2009; Juste and Alcaldé, 2016a). Aerial hawking, water stress, and its small distribution range are believed to be the most important risk factors for this species linked with climatic change (Sherwin et al. (2012: 174). CONSERVATION ACTIONS: Juste and Alcaldé (2016a) supports Hutson et al. (2008o) [in IUCN (2009)] who reported Rhinolophus euryale is protected by national legislation in most range states. There are also international legal obligations for protection of this species through the Bonn Convention (Eurobats) and Bern Convention, where these apply. It is included in Annex II (and IV) of EU Habitats and 300 ISSN 1990-6471 Species Directive, and hence requires special measures for conservation including designation of Special Areas for Conservation. There is some habitat protection through Natura 2000, and some roosts are already protected by national legislation). The species is directly or indirectly benefiting from EU LIFEfunded projects in France, Spain and Italy. No specific measures are in place in North Africa. GENERAL DISTRIBUTION: Rhinolophus euryale is a western Palaearctic species, occurring in southern Europe, north-west Africa (known range extends across northern Morocco, Algeria, and Tunisia), and the Near East. There is only a single record from Cyprus, but this is regarded by most authors to be R. mehelyi. It is widely distributed over its range, and is found from sea level to 1,000 m. Native: Abkhasia; Albania; Algeria (Csorba et al., 2003: 16); Andorra; Armenia; Azerbaijan; Bosnia and Herzegovina; Bulgaria; Croatia; France (Corse) (Csorba et al., 2003: 16); Georgia; Gibraltar; Greece (East Aegean Is.); Holy See (Vatican City State); Hungary; Iran (Csorba et al., 2003: 16), Islamic Republic of; Iraq; Israel (Csorba et al., 2003: 16); Italy (Sardegna, Sicilia); Jordan; Lebanon [see Horácek et al. (2008: 55)] (Csorba et al., 2003: 16); Macedonia, the former Yugoslav Republic of; Monaco; Montenegro; Morocco (Csorba et al., 2003: 16; El Ibrahimi and Rguibi Idrissi, 2015: 359); Palestinian Territory, Occupied; Portugal (Csorba et al., 2003: 16); Romania; Russian Federation; San Marino; Serbia; Slovakia; Slovenia; Spain; Syrian Arab Republic (Csorba et al., 2003: 16); Tunisia (Csorba et al., 2003: 16; Dalhoumi et al., 2016b: 867); Turkey (Csorba et al., 2003: 16). Regionally extinct: Switzerland. Presence uncertain: Cyprus [see Benda et al. (2007: 71)]; Egypt; Turkmenistan. DETAILED MORPHOLOGY: Baculum - See Csorba et al. (2003: 15). In their study on take-off performance, Gardiner et al. (2014: 1059) determined a wing beat frequency of 11.48 ± 0.50 Hz. ECHOLOCATION: The data for 38 animals from Greece were the following: Fstart: 91.8 ± 5.00 kHz, Fend: 88.7 ± 3.49 kHz, Fpeak: 104.8 ± 0.96 kHz, bandwidth: 3.5 ± 1.05 kHz, duration 53.8 ± 20.42 msec and interpulse interval: 84.5 ± 30.01 msec (Papadatou et al., 2008b: 132). Walters et al. (2012: suppl.) report the following figures for 26 calls (15 sequences) from French and Italian bats: duration: 35.23 ± 18.06 msec, Fmax: 102.28 ± 1.20 kHz, Fmin: 88.22 ± 4.30 kHz, bandwidth: 14.06 ± 3.69 kHz, Fpeak: 101.68 ± 2.18 kHz. Animals from Tunisia emitted sounds at 102.39 ± 1.01 kHz (105.0 - 106.5 kHz; Dalhoumi et al., 2016b: 866). Schuchmann et al. (2012: 161) found that both sexes were able to recognize the sex of conspecifics from their echolocation calls. Luo et al. (2019a: Supp.) reported the following data (for two calls): Fpeak: 104.8, 102.4 kHz, Fstart: 91.8, 93.8 kHz, Fend: 88.7, 89.1 kHz, and duration: 53.8, 40.6 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Both Bovey (1949) and Capanna and Manfredi Romanini (1971) reported 2n = 58, FN = 60. Rushton (1970: 463) also reported 2n = 58, FN=60, with two (sub)metacentric, 24 acrocentrics, and 2 minute pairs or acrocentric chromosomes. The X chromosome is reported as submetacentric and the Y chromosome as minute. Zima et al. (1992a) reported 2n=58, FN=60, BA=4, a submetacentric X chromosome and a metacentric Y chromosome for a specimen from Greece. Capanna and Civitelli (1964a: 371) reported 2n = 58, including 24 acrocentric pairs, 2 metacentric pairs and two pairs of very small chromosomes. The X chromosome was reported as a large submetacentric and the Y as a very small one. In their study of mitochondrial DNA, Budinski et al. (2019: 770, 772) found that northern African R. euryale clustered together with the samples from Europe and Turkey in Clade III, indicating that these orignated from the same source population. Protein / allozyme - Unknown. HABITAT: Russo et al. (2005: 327) and Goiti et al. (2003: 75; 2008: 498 - 499) indicate that, in Spain, R. euryale always foraged in areas where there were trees or hedegrows, and never over open meadows, pastures, heath, gorse- or fern-covered areas (or any other open habitats). About 96 % of the bats they tagged, used hedgerows to forage in. The second most frequented habitat (90 %) was broadleaved woodland, whereas isolated trees were visited between 1 and 12 %. Exotic tree plantations (e.g. Eucalyptus formed the least preferred habitat. Furthermore, the breeding colony size was found to be positively correlated African Chiroptera Report 2020 with the availability of edges around breeding sites. HABITS: From the Iberian Peninsula, Goiti et al. (2006: 141) reported that during pre-breeding foraging occurred on average within 1.3 km, and at most 4.2 km from the roost. For lactating females, the average distance was 4.3 km, and a maximum distance from the roost of 9.2 km was observed. In the same period, males foraged closer (mean 1.9 km). During the post-lactation period, adults travelled a mean of 4.6 km, whereas juveniles flew on average only 2.6 km. ROOST: Based on data from Sardinia, Russo et al. (2014: 9) rule out that R. euryale and R. mehelyi compete for roosting sites. DIET: In northern Spain, Goiti et al. (2008: 496 - 497) examined 810 feces from 168 bats. These showed that the bat's diet included 10 prey categories, of which the Lepidoptera (moths) formed the staple diet (between 68 and 99 % of the diet). Moths are also the most important prey during lactation and post-lactation. Another important prey item, although very seasonal and especially prebreeding, was the beetle Rhizotrogus (Scarabaeidae). Other food items (in descending rank) included lacewings (Neuroptera, Chrysopidae and Hemerobiidae), craneflies (Diptera, Tipulidae), brachyceran flies, Psocoptera, and Hymenoptera. Diets were similar for adults and juveniles. In the same area, Aldasoro et al. (2019: 9) found the following prey groups (in descending order of occurrence): Lepidoptera (100 %), Diptera (94 %), Neuroptera (22 %), Ephemenoptera (17 %), Hemiptera (11 %), Hymentoptera (6 %) and Trichoptera (6 %). Arrizabalaga-Escudero et al. (2019: 1) found that seasonality was the most important factor explaining the dietary variation in adult bats. They also found that young bats ate slower, smaller and lighter moths, suggesting that they still need to acquire the skills to capture the same prey items as adults. In the southwestern Iberian peninsula, Arrizabalaga-Escudero et al. (2018: Suppl.) identified the following food items from faecal samples of R. euryale (food items marked with an asterisk were also found in faecal samples of R. mehelyi): Coleoptera (Cerambycidae: Arhopalus ferus (Mulsant, 1839)*); Diptera (Tachinidae: unidentified*: Phryno vetula (Meigen, 1824); Muscidae: Polietes lardarius (Fabricius, 1781)); Lepidoptera (Crambidae: Calamotropha paludella (Hübner, 1824)*; Metasia ophialis (Treitschke, 301 1829); Ostrinia nubilalis (Hübner, 1796)*; Erebidae Catocala nymphaea (Esper, 1787); Catocala nymphagoga (Esper, 1787)*; Lygephila lusoria (Linnaeus, 1758); Gelechiidae: Teleiopsis sp.*; Geometridae: Aplasta ononaria (Füssli, 1783); Camptogramma bilineata (Linnaeus, 1758)*; Ennomos quercaria (Hübner, 1813)*; Gymnoscelis rufifasciata (Haworth, 1809); Idaea ochrata/rufaria; Pachycnemia hippocastanaria (Hübner, 1799)*; Rhoptria asperaria (Hübner, 1817)*; Noctuidae: Acronicta rumicis (Linnaeus, 1758); Agrotis segetum/trux*; Agrotis ipsilon (Hufnagel, 1766)*; Amphipyra tragopoginis (Clerck, 1759); Apamea monoglypha/sicula; Apamea arabs Oberthur, 1881; Autographa gamma/pulchrina*; Calophasia platyptera (Esper, [1788])*; Cosmia trapezina (Linnaeus, 1758); Mythimna albipuncta (Denis & Schiffermüller, 1775)*; Mythimna loreyi/sicula*; Peridroma saucia (Hübner, 1808)*; Polyphaenis sericata (Esper, 1787)*; Sesamia nonagrioides (Lefèbvre, 1827)*; Nolidae: Meganola strigula (Denis & Schiffermüller, 1775); Praydidae: Prays fraxinella Bjerkander, 1784*; Pyralidae: Ephestia mistralella (Millière, 1874); Pempelia palumbella (Denis & Schiffermüller, 1775)*; Phycita sp.; Tortricidae: Cnephasia sp. 1*; Cnephasia sp. 2*); Neuroptera (Chrysopidae: Chrysoperla sp.*; Cunctochrysa albolineata (Killington, 1935); Pseudomallada flavifrons (Brauer, 1851); Myrmeleontidae: Distoleon tetragrammicus (Fabricius, 1798)*); Orthoptera (Tettigoniidae: Tessellana tessellata (Charpentier 1825)). Of the 62 prey taxa, 32 (representing over 75 % of all food items) were consumed by both bat species. Andreas et al. (2013) using faecal analysis of R. euryale from southern Slovakia, showed that this species mainly fed on medium-sized moths (25 40 mm) (%f = 60; %vol = 74) and the diet of R. euryale contained Neuroptera (Hemerobiidae) (%f = 2.2; %vol = 0.3), nematoceran Diptera (%f = 13.3; %vol = 9.1) and Trichoptera (%f = 6.7; %vol = 3.6). In the same area (and northern Hungaria), Miková et al. (2013: 848) investigated the relation between food and ambient temperature during winter. Ahmim and Moali (2013: 175) examined droppings from northern Algeria, which contained mainly Insecta (96.19%) and Chilopoda (3.81%). Diptera accounted for 29.00 % (Culicidae: 14.29 %, Chironomidae/Ceratopogonidae: 7.14 %, and Tipulidae: 5.71 %. Lepidoptera were the second most important order and accounted for 19.08 %. Laurent (1944a: 13) tried to (force) feed captive animals with mealworms (Tenebrio molitor), but these were rejected. 302 ISSN 1990-6471 Gardiner et al. (2014: 1058) assessed its foraging strategy as "low altitude slow aerial hawking", with no observations of gleaning. In Hungary, Maxinová et al. (2017a: 98) examined "non-consumptive faeces" (produced during hibernation after no feeding activity) and found these not to contain any insect DNA, but rather bat DNA as well as peptides belonging to other craniates and eukaryotes. They also found bacterial peptides and one Nematode. Furthermore, the high concentration of inorganic material suggest that drinking as well as direct sediment consumption occur inside the cave environment during the hibernation period. They conclude that winter arousals are primarily aimed at regulating the water balance. Maxinová et al. (2017b: 93) also found that the bat's digestive enzymes (amylase, chitobiase, endochitinase and glukosaminidase) remained active during the entire hibernation period. POPULATION: Structure and Density:- According to Hutson et al. (2008o) [in IUCN (2009)] supported by Juste and Alcaldé (2016a), R. euryale is an infrequent species. Summer colonies number ca. 50 - 1,500 individuals (S. Aulagnier pers. comm., 2007). Winter clusters typically number up to 2,000 animals. It occurs in large vulnerable colonies and is considered threatened in many range states. Large population declines have been reported in several European countries, including Spain (Palomo and Gisbert, 2002). In France, the population declined by ca. 70 % between 1940 and 1980, although subsequently the trend appears to have stabilised (Brosset et al., 1988; S. Aulagnier pers. comm., 2007). Apart from R. blasii, which may have gone extinct in the country, R. euryale is probably the rarest rhinolophid in Italy, and anecdotal evidence suggests that a number of colonies have declined in the past few decades (D. Russo pers. comm., 2006). The species has a very small and declining population in Portugal (Rodrigues et al., 2003; Cabral et al., 2005). It is stable and common in the central and eastern Balkans (M. Paunovic pers. comm., 2007). There is little information on population trends outside Europe, although it is suspected that continuing declines have also occurred in at least parts of the non-European range. For example, in Iran the species is no longer found in caves which 30 years ago held 20,000 individuals of different species (M. Sharifi pers. comm., 2005). Trend:- 2016: Decreasing (Juste and Alcaldé, 2016a). 2008: Decreasing (Hutson et al., 2008o; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 13 years, and Ibáñez et al. (2018: 80) recaptured a male bat in La Rioja (Spain) that was banded 21 years before. REPRODUCTION AND ONTOGENY: Tuttle and Stevenson (1982: 121) indicate that males need about 27 months to reach sexual maturity, whereas females need between 15 and 27 months. Szekely et al. (2015: Suppl.) mention that both males and females become sexually mature after 821 days (= about 27 months). The gestation period takes 90 days, after which a young is born with a weight of 3.89 g (adult: 8.2 g). POSTNATAL DEVELOPMENT: In Iran, Eghbali et al. (2017: 276) found that the young's body mass increased linearly until day 23, when it attained 74.29 % of the mean mass of adult females. Its body mass increased (from 3.58 g at birth) by 0.36 g per day and its forearm length increased (from 20.10 mm at birth) by 1.41 mm/day. By day 70, the epiphysal gap was closed. The wing load decreased by 0.09 Nm -2/ day until day 36, and subsequently increased again to a maximum of 6.56 ± 0.30 Nm -2 by day 80. PARASITES: Hirst (1923: 989) mentions that Kolenati described a mite (Periglischrus interruptus) from a bat, he originally identified as "R. clivosus [= R. blasii]", but that he corrected this later to R. euryale. Ševcík et al. (2012: 35) report on the presence of the bat flies Nycteribia schmidlii schmidlii Schiner, 1853 and Phthiridium biarticulatum Hermann, 1804 on bats from Cyprus. The bat flies Brachytarsina flavipennis Macquart, 1851 (Streblidae) and Phthiridium biarticulatum Hermann, 1804 (Nycteribiidae) were reported from Jordan by Benda et al. (2010b: 238). Benda et al. (2010b: 238) also refer to previous reports from Turkey mentioning the following parasites: Nycteribia pedicularia Latreille, 1805, N. schmidlii Schiner, 1853, N. kolenatii Theodor and Moscona, 1954, N. vexata Westwood, 1835, Brachytarsina flavipennis Macquart, 1851, and Rhinolophopsylla u. unipectinata (Taschenberg, 1880) and in Palestine Penicillidia conspicua Speiser, 1901 and P. dufourii (Westwood, 1835). Bendjeddou et al. (2017: 15) reported the following ectoparasites from Algeria: Nycteribiidae: Nycteribia (Nycteribia) pedicularia Latreille, 1805, Penicillidia (Penicillidia) dufourii Westwood, 1835 African Chiroptera Report 2020 and Phthiridium biarticulatum Hermann, 1804; and Siphonaptera: Rhinolophopsylla unipectinata arabs Jordan and Rothschild, 1921. From Tunisia, Vermeil (1960: 738) reported the presence of the Nycteribiids Nycteribia latreillii Leach, 1817 and Nycteribia vexata vexata Westwood, 1835. From non-specified areas, Haelewaters et al. (2018: 974) reported the Nycteribiid Penicilidia conspicua Speiser, 1901, which might be a host for the fungus Arthrorhynchus nycteribiae (Peyr.) Thaxt. Malinicová et al. (2017: 211) investigated the gut microflora of Hungarian R. euryale during hibernation and found that the number of enterobacteria and enterococci decreased during hibernation and were restored at the end of hibernation. 303 Orthomyxoviridae Despite sampling and testing of 2 individuals of this species, no evidence of influenza A-like viruses (Orthomyxoviridae) were obtained (Fereidouni et al., 2015). Picornaviridae The presence of Picornavirus was mentioned by Nieto-Rabiela et al. (2019: Suppl.). Rhabdoviridae Lyssavirus Serra-Cobo et al. (2018: 2) found one out of 31 Algerian bats testing seropositive for European bat lyssavirus1 and none of the twelve Moroccan bats. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Egypt, France, Morocco, Tunisia. VIRUSES: Adenoviridae Mastadenovirus This virus was mentioned by Nieto-Rabiela et al. (2019: Suppl.). Coronaviridae - Coronaviruses Luis et al. (2013: suppl.) mention the presence of a SARS coronavirus-related virus. Alphacoronavirus Reported from Germany by Kohl and Kurth (2014: 3112). Ar Gouilh et al. (2018: 90) reported the EPI 10 virus from an R. euryale from Morocco. Figure 89. Distribution of Rhinolophus euryale Betacoronavirus Reported from Bulgaria and Germany by Kohl and Kurth (2014: 3112). Rhinolophus ferrumequinum (Schreber, 1774) *1774. Vespertilio ferrum-equinum Schreber, Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen, 1 (8): pl. 62, upper fig + 1 (9): 174. Publication date: 1774. Type locality: France: "France". - Etymology: From the Latin "ferrum" meaning iron and "equus" meaning horse. 1792. Vespertilio ferrum-equinum major Kerr, Animal Kingdom, 1 (1): xvii, 99. Type locality: France. - Comments: This might be an African form, but we lack further information. 1803. major E. Geoffroy Saint-Hilaire, Catalogue des Mammifères du Muséum national d'Histoire naturelle, Paris, ???. - Comments: This might be an African form, but we lack further information. 1961. Rhinolophus ferrumequinum millali: Lavocat, Notes Mém. Serv. Mines Carte geol. Maroc, 155: ?. Type locality: Morocco. - Comments: This is a fossil form from the MiocenePliocene. (Name Combination) ? Rhinolophus ferrumequinum ferrumequinum: (Name Combination) ? Rhinolophus ferrumequinum: (Name Combination, Current Combination) 304 ISSN 1990-6471 TAXONOMY: Taxonomic revisions by Felten et al. (1977), Strelkov et al. (1978), Kryštufek (1993) and Thomas (1997). Thomas (1997) recognized seven subspecies based on morphological characters (R. f. ferrumequinum in Europe and northwest Africa, R. f. creticum in Crete, R. f. irani in Iran, Iraq and Turkmenistan, R. f. proximus from Afghanistan and Pakistan to India, R. f. tragatus in north India and China, R. f. korai in Korea and R. f. nippon in Japan). ferrumequinum species group (Csorba et al., 2003: 42 - 45; Simmons, 2005: 355). See Csorba et al., 2003: 44 - 45) for further remarks on the taxonomy. Based on cytochrome b analyses, Koh et al. (2014: 97) indicate that northern African populations form one clade with specimens from Europe and western Asia, which is different from three other clades: far-eastern Asia, eastern China and central China. COMMON NAMES: Albanian: Lakuriq nate hundëpatkua i madh. Arabian: Khafash abu hadwa kabir, Khaffash. Armenian: Մեծ պայտաքիթ. Azerbaijani: Böyük nalburun. Basque: Ferra-saguzar handi. Belarusian: Падкаванос вялікі. Bosnian: Veliki potkovasti šišmiš. Breton: Frigribell vras. Bulgarian: Голям подковонос. Castilian (Spain): Murciélago grande de herradura, Rinolofo grande de herradura. Catalan (Spain): Rat penat gran de ferradura, Ratapinyada gran de ferradura, Ratpenat de ferradura gran. Croatian: Veliki potkovnjak, Veliki potkovasti šišmiš. Czech: Vrápenec velký, wrápenec podkowní, podkowáček, podkováček, obecný, vrápenec podkovní, vrápenec podkovníček, vrápenec veliký, veliký vrápenec podkovní, vrápeník podkovní, vrápenec podkrovní. Danish: Stor hestekonæse. Dutch: Grote hoefijzerneus. English: Larger Horseshoe Bat, Greater Horseshoe Bat. Estonian: Suur-sagarnina. Finnish: Euroopanisoherkko. French: le grand rhinolophe fer à cheval, Grand rhinolophe, Grand Fer-àCheval. Frisian: Grutte hoefizernoas. Galician (Spain): Morcego de ferradura grande. Georgian: დიდი ცხვირნალა. German: Große Hufeisennase, Großhufeisennase, Hufeisennase. Greek: Τρανορινόλοφος. Hebrew: ‫גדול פרסף‬, parsaf gadol. Hungarian: Nagy patkósdenevér nagy patkósorrú denevér, Rhinolophos i megali. Irish Gaelic: Crú-ialtóg mhór. Italian: Rinolofo maggiore, Ferro di Cavallo Maggiore. Latvian: Lielais pakavdegunis. Lithuanian: Didysis pasagnosis. Luxembourgish: Grouss Huffeisennues. Macedonian: Голем потковичар [= Golem Potkovichar]. Maltese: Rinolofu Kbir, Farfett il-Lejl tan-Nala kbir. Montenegrin: Veliki potkovičar. Nepali: Thulo Ghodnale Chamero. Norwegian: Storhesteskonese, Stor hesteskonese. Polish: Podkowiec duŻy. Portuguese: Morcego-de-ferradura-grande. Rhaeto-Romance: Rinolof grond, Nas fier-chaval grond. Romanian: Liliacul mare cu potcoavă, Liliac-mare-cupotcoavã. Russian: Подковонос большой, Болъшой подковонос [= Bolshoj podkovonos]. Serbian: Велики потковичар [= Veliki potkovičar]. Scottish Gaelic: Crudh-ialtag mhòr. Slovak: Podkovár velký. Slovenian: Veliki podkovnjak, mali podkovnjak. Swedish: Större hästskonäsa. Turkish: Büyük Nalburunlu Yarasa. Ukrainian: Підковик (Підковоніс) великий [=Pidkovyk velykyy]. Welsh: Ystlum pedol mwyaf. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Lavocat (1961) described a new subspecies found in deposits of the Mio-Pliocene, at Beni Mellal. See Butler (1978). CONSERVATION STATUS: Global Justification This species has a large range. Although there have been marked and well-documented declines in some areas, the species remains widespread, abundant, and apparently stable in other areas. Assessed as Least Concern (LC ver 3.1 (2001)) (Aulagnier et al., 2008a; IUCN, 2009; Piraccini, 2016b). Assessment History Global 2016: LC ver 3.1 (2001) (Piraccini, 2016b). 2008: LC ver 3.1 (2001) (Aulagnier et al., 2008a; IUCN, 2009). 2000: LR/nt. 1996: LR/nt ver 2.3 (1994). Regional None known. MAJOR THREATS: Piraccini (2016b) supportsAulagnier et al. (2008a) [in IUCN (2009)] who reported the main threats are fragmentation and isolation of habitats, change of management regime of deciduous forests and agricultural areas, loss of insects due to pesticide use, and disturbance and loss of underground habitats and attics. In northwest Europe, habitat change is likely to have been amongst the major causes of declines, the conversion of woodland and small-field landscape to large-scale agricultural land being particularly damaging. While declines elsewhere, particularly in eastern Europe, may currently not be so marked, the loss of cultural landscapes in those countries as they move towards western-style economies may have significant effects in the near future. The use of pesticides has been a recognized threat to the insect food, particularly where these have been directed against the larvae of favoured food items, African Chiroptera Report 2020 such as melolonthid beetles, larvae of noctuid moths or crane-flies. Favoured prey may be affected secondarily by pesticide use, such as the loss of dung fauna from the use of persistent antiparasitic drugs (avermectins) on cattle. Populations in caves and other underground habitats have suffered from increased disturbance (for example by tourist visits to caves). In buildings, colonies may be affected by human intolerance, renovation work or the application of pesticides, such as some of those used for the remedial treatment of timbers (Hutson et al., 2001). In South Asia, this species is threatened by deforestation, generally resulting from logging operations and the conversion of land for agricultural and other uses. Disturbance to roosting sites is likely to be a potential threat to the populations of this species (Molur et al., 2002). Concerning climatic change, the most important risk factors identified by Sherwin et al. (2012: 174), are water stress, aerial hawking and long-distance dispersal. CONSERVATION ACTIONS: Piraccini (2016b) supports Aulagnier et al. (2008a) [in IUCN (2009)] who reported Rhinolophus ferrumequinum has been the subject of widespread conservation activity, especially in Europe. Until recently, this has concentrated on roosts in buildings and caves. Many buildings used as roosts have management agreements and many underground sites have been protected. Nevertheless, sites continue to be lost or damaged. More recently, attention has turned to identifying more precisely the food and foraging requirements. A European meeting (Germany, May 1995) discussed the status and conservation needs for the species on a pan-European scale. The Bern Convention has commissioned a Europe-wide Species Action Plan under the PanEuropean Biological and Landscape Diversity Strategy (Ransome and Hutson, 2000). It is protected by national legislation in some range states. There are international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention in parts of its range where these apply. It is included in Annex II (and IV) of the European Union Habitats Directive, and hence requires special measures for conservation including designation of Special Areas for Conservation. There is some habitat protection through Natura 2000 (some roosts are already protected by national legislation). Flanders and Jones (2009: 894) point out that, in the U.K., management of broad-leaved woodland areas in close proximity (within 4 km) of transitional roosts (used in spring and autumn) should be enhanced. 305 There are no specific conservation measures in place for the species in North Africa or South Asia. Populations should be monitored to record changes in abundance and distribution (Molur et al., 2002). GENERAL DISTRIBUTION: Rhinolophus ferrumequinum has a wide range in the Palaearctic, occuring from North Africa and southern Europe through south-west Asia, the Caucasus, Iran, Afghanistan, Pakistan and the Himalayas to south-eastern China, Korea, and Japan (Csorba et al., 2003; Abe et al., 2005). It usually occurs below 800 m asl, but can be found up to 3,000 m asl in the Caucasus depending on roost availability and humidity (K. Tsytsulina pers. comm., 2005). Native: Abkhasia; Afghanistan; Albania; Algeria; Andorra; Armenia; Austria; Azerbaijan; Bangladesh; Bhutan; Bosnia and Herzegovina; Bulgaria; China; Croatia; Cyprus; Czech Republic; France (Corse); Georgia; Germany; Gibraltar; Greece (Kriti); Hungary; India; Iran, Islamic Republic of; Iraq; Israel; Italy (Sardegna, Sicilia); Japan; Jordan; Kazakhstan; Korea, Democratic People's Republic of; Korea, Republic of; Kyrgyzstan; Lebanon; Liechtenstein; Luxembourg; Macedonia, the former Yugoslav Republic of; Moldova; Monaco; Montenegro; Morocco (El Ibrahimi and Rguibi Idrissi, 2015: 358); Nepal; Pakistan; Palestinian Territory, Occupied; Poland; Portugal; Romania; Russian Federation; San Marino; Saudi Arabia; Serbia; Slovakia; Slovenia; Spain (Baleares); Switzerland; Syrian Arab Republic; Tajikistan; Tunisia (Dalhoumi et al., 2014: 53, 2016b: "3"; 2019b: 26); Turkey; Turkmenistan; Ukraine; United Kingdom; Uzbekistan. Possibly extinct: Malta. Regionally extinct: Belgium; Netherlands. BIOGEOGRAPHY: See Rossiter et al. (2007). GEOGRAPHIC VARIATION: Benda et al. (2014c: 16) mention that the Libyan R. ferrumequinum specimens are very small in size, and even the smallest within the Mediterranean range of this species, which led to their confusion with R. clivosus. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Lucati and López-Baucells (2016: Suppl.) refers to an albino specimen that was reported from "South Africa?" by Allen (1939b). DETAILED MORPHOLOGY: Baculum - See Csorba et al. (2003: 43). 306 ISSN 1990-6471 Forman (1973) [in Scillitani et al. (2005: 302)] regards the stomach of this bat as primitive for the order and probably represents the ancestral condition of lower Eutheria, because it presents a number of features in common with Insectivora. R. ferrumequinum is one of the few bat species, where the females can have a baubellum (os clitoris) (Lough-Stevens et al. (2017: 1078). FUNCTIONAL MORPHOLOGY: Kobayashi (2018: 69) found that the bat's M. quadriceps femoris has a short muscle belly attached to the proximal portion of the femur and that the insertional tendon of this muscle and its patellar ligament are very thin. These features indicate that R. ferrumequinum cannot exert a strong and long-distance extension of its knee joints, which results in the inability to crawl. In their study - covering the entire distribution range of R. ferrumequinum - Jiang et al. (2019: 1) found that populations with long forearms in both sexes occurred in areas with higher mean temperatures in the warmest quarter and lower mean temperatures in the coldest quarter. ECHOLOCATION: Fukui et al. (2011: 1548) report that, in Japan, the detection range for Rhinolophus ferrumequinum calls is less than 5 metres. Jones and Siemers (2010: 450) indicate that juveniles emit lower frequencies than adults, and Jones and Ransome (1993: 125) found that the young's resting constant-frequency (RF) values are at least partially determined by their mother's RF. Jones and Ransome (1993: 125) furthermore report that calls of R. ferrumequinum specimens aged between 1 and 28 years vary seasonally and over a lifetime in a predictable manner. For 28 Greek specimens, Papadatou et al. (2008b: 132) report a start frequency of 66.5 ± 3.23 kHz, a terminating frequency of 64.9 ± 4.33 kHz, and a bandwidth of 3.9 ± 1.34 kHz. The duration of the call is 53.8 ± 10.50 msec, and the interpulse interval is 92.1 ± 24.94 msec. Andrews et al. (2006: 199) analyzed 500 echolocation calls from UK bats, and found an average frequency of 83.8 ± 0.4 kHz and 21.0 ± 6.4 ms duration. Another set of 500 calls had an average frequency of 84.0 ± 0.5 kHz and a duration of 31.9 ± 9.5 ms. Social calls were emitted at frequencies between 15 and 29 kHz and their duration was between 4 and 49 ms (p. 201). Walters et al. (2012: suppl.) report the following figures for 30 calls from bats from France, Switzerland and the UK: duration: 50.08 ± 13.52 msec, Fmax: 81.66 ± 1.68 kHz, Fmin: 70.29 ± 3.62 kHz, bandwidth: 11.37 ± 3.19 kHz, Fpeak: 81.46 ± 1.65 kHz. Dalhoumi et al. (2016b: 866) reported a long constant frequency part between 85 and 88 kHz for Tunisian animals. Luo et al. (2019a: Supp.) reported the following data (for two calls): Fpeak: 82.3, 78.7 kHz, Fstart: 69.3, 66.5 kHz, Fend: 70.3, 64.9 kHz, and duration: 49.4, 53.8 msec. From experiments in a flight tunnel, Koselj and Siemers (2013: 81) found that R. ferrumequinum bats can extract information from echolocation calls produced by conspecifics. This could facilitate sensory cooperation among echolocating bats. In China, Sun et al. (2013: 1) found regional differences in the resting frequency of echolocation calls of R. ferrumequinum which were significantly correlated with geographic distance and mean annual temperature. Besides the ultrasound calls used in echolocation, Andrews and Andrews (2003: 221) also report on twelve types of ultrasound social calls, which were in the range of 20 to 29 kHz and had a duration of 1 to 49 ms. Furthermore, there were also low frequency (audible) social calls (1 - 10 kHz). For Morocco, Disca et al. (2014: 226) indicate that the type of call is FM-FC-FM, with Fpeak: 84.6 ± 0.8 kHz and duration: 30.3 ± 7.6 msec. MOLECULAR BIOLOGY: DNA - See Rossiter et al. (2007). Karyotype - Makino (1948), Capanna and Civitelli (1964a: 363), Rushton (1970: 463), Baker et al. (1974; 1975) and Qumsiyeh et al. (1986) all reported 2n = 58, FN = 60, a submetacentric X chromosome, and an acrocentric Y chromosome. Volleth and Eick (2012: 167) reported 2n = 58 and 28 as segment number. In Turkey, Arslan and Zima (2014: 7) found 2n = 58, FNa = 60-62, FN = 64-66, a metacentric X and an acro-/metacentric Y. The autosomal chromosomes consist of two medium-sized bi-armed pairs and 26 acrocentric pairs, but they indicate that one of the acrocentric pairs is considered by some other authors to be biarmed, leading to the different FN and FNa values. Capanna and Civitelli (1964a: 371) mention 24 pairs of acrocentric chromosomes, 2 metacentric pairs, and 2 pairs of very small chromosomes; the X chromosome was reported as a large submetacentric and the Y as a very small one. Protein / allozyme: Unknown. African Chiroptera Report 2020 HABITAT: In the U.K., Flanders and Jones (2009: 891) reported that on an overal area of about 632 ha used by a colony of R. ferrumequinum specimens, 28.7 % consisted of pasture, 26.6 % of broadleaved woodland, 16.1 % of arable, 8.2 % of mixed woodland, 6.7 % urban, 5.8 % scrub, 4.3% grassland, 1.6 % heathland, 1.4 % water, and 0.7 % coniferous woodland. They also reported the following habitat preferences (high to low): pasture, broad-leaved woodland, mixed woodland, water, scrub, grassland, urban, coniferous woodland, heathland, and arable. Fukui et al. (2011: 1551) indicate that, in Japan, this species prefers natural forest gaps over artificially made gaps as in the latter debris is generally removed, leaving no tree branches for it to hang from while scanning for prey. HABITS: Park et al. (2000) [in Sherwin et al. (2012: 172)] found that R. ferrumequinum was longer active when the temperatures were higher than 10 °C, which they considered to be a thermal threshold for insect activity. DIET: Flanders and Jones (2009: 892) examined 1,580 bat droppings in the U.K. and found that Lepidoptera (moths) were the most abundant prey. In spring, Melolontha melolontha (May bugs), beetles of the genus Geotrupes (dor beetles), Trichoptera (caddis flies), Tipulidae (crane flies), Ichneumonidae (ichneumonid wasps), and Lepidoptera were contributing to the diet. In autumn, the two beetle species were replaced by representatives of the genus Aphodius (dung beetles), and additional to the other groups of insects, Diptera (mainly dung flies) were also part of the main prey types. Andreas et al. (2013) using faecal analysis of R. euryale from southern Slovakia, showed that this species fed mainly on large moths (>40mm) (%f = 54; %vol = 89) and comprised also Coleoptera (%f = 19.5; %vol = 4.82), represented by Scarabaeidae, Elateridae and Curculionoidea. In northern Spain, Aldasoro et al. (2019: 9) found the following prey groups (in descending order of occurrence): Diptera, Lepidoptera, Coleoptera, Neuroptera, Hemiptera and Trichoptera. In northern Algeria, Ahmim and Moali (2013: "2") found that R. ferrumequinum fed on three groups of arthropods: Insecta (95.31 % [Diptera: 34.56 % and Lepidoptera: 24.13 %]), Chilopoda (4.49 %) and Araneida (0.20 %). The largest groups within the Diptera were Culicidae (10.40 %), 307 Chironomidae/Ceratopogonidae (10.94 %) and Tipulidae (4.28 %). In Libya, Benda et al. (2014c: 19) examined faecal pellets and found one set to contain 84 % by volume larger moths, 9 % ants and 7 % beetles, whereas a second set contained only Lepidoptera with wingspans of ca. 40 mm. POPULATION: Structure and Density:Rhinolophus ferrumequinum is an infrequent species in most parts of its range, although in at least parts of southwest Asia and the Caucasus it is abundant and widespread (it is the most frequently reported species in Turkey: A. Karatas pers. comm., 2005), with populations in Iran and Turkey considered to be stable (A. Karatas, M. Sharifi and K. Tsytsulina pers. comm., 2005), although it may be decreasing in Russian parts of the Caucasus (S. Kruskop pers. comm., 2005). Summer colonies of c. 30 200 individuals (up to 400 animals), and winter clusters of up to 500 animals are typical. In Europe, the two most widespread Rhinolophus species, R. ferrumequinum and R. hipposideros, are of particular conservation concern and are the subject of considerable research and monitoring. R. ferrumequinum has shown marked declines in range in northwest Europe within the last 100 years (e.g. United Kingdom, Germany, Austria), and has gone extinct in some countries (eg. Belgium, Netherlands). However, there are signs of stabilisation and/or recovery in some northwest European countries (Hutson et al., 2001). For example, in the UK the species declined massively in the past, but it is now stable at a low population level (around 5,000 individuals) (Ransome and Hutson, 2000). However, in Austria declines continue, with population reductions of 70 % in the last 10 years (from 100 to 30 breeding individuals: Spitzenberger (2002); F. Spitzenberger pers. comm., 2006). In other parts of Europe, trends vary and are generally less well known: in Malta the species has gone extinct, in Portugal and Spain the trend is not known (although some colonies have disappeared in Spain) (Palomo and Gisbert, 2002; Cabral et al., 2005), in Croatia the population is thought to be stable (N. Tvrtkovic pers. comm.), and in Romania the population has been slowly increasing since 1989 due to reduced use of pesticides and a return to traditional agriculture with colonies of up to 800 individuals. In Switzerland the species is very rare (3 maternity roosts with some 200 individuals), but the population trend appears stable (H. Kraettli pers. comm., 2006). In its north African and south Asian range the population size and trends are unknown (Aulagnier et al., 2008a; IUCN, 2009; Piraccini, 2016b). 308 ISSN 1990-6471 Trend:- 2016: Decreasing (Piraccini, 2016b). 2008: Decreasing (Aulagnier et al., 2008a; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 30.5 years. ACTIVITY AND BEHAVIOUR: In the U.K., Flanders and Jones (2009: 891) reported a maximum nightly foraging range of between 2.17 and 2.44 km, and the foraging area averaged 148.1 ha. Fukui et al. (2011: 1551) [referring to Jones and Rayner (1989)] indicate that R. ferrumequinum is a "flycatcher" species, meaning that it hangs from tree branches while scanning for its prey. Smotherman et al. (2016: 537) refer to Ma et al. (2006) in describing the song-like structure for this species as consisting of different syllable types in series. REPRODUCTION AND ONTOGENY: Tuttle and Stevenson (1982: 121) mention that males attain sexual maturity between 15 and 51 months, whereas this for females is between 15 and 39 months. Szekely et al. (2015: Suppl.) report 730 days (about 24 months) for both sexes. Gharaibeh (1997: 55) mentions lactating females and juveniles from the cave at Djebel Oust (Tunisia) on 29 July 1996. Benda et al. (2014c: 16) collected a pregnant female on 28 May at Nanatalah (Libya). Kurta and Kunz (1987: 82) report that, at birth, the young weights about 29.3 % of the mother's body mass (5.8 versus 19.8 g), and that its forearm length is about 36.0 % of the mother's length (25.3 versus 70.3 mm). The gestation period takes 80 days (with the young having a birth weight of 5.8 g - 25.4 % of the adult), and weaning occurs after 60 days (Szekely et al., 2015: Suppl.). MATING: Fenton (1984) [in Keeley and Keeley (2004: 117)] reports the existence of copulatory plugs. For R. ferrumequinum, these plugs are formed from male secretions, and probably serve to prevent further insemination by other males, rather than preventing sperm leakage from the female's reproductive trace (in that case the plugs would probably be made by the females). However, Rossiter et al. (2000b) mention that these plugs can be ejected by the females, which would allow additional matings. In Britain, Rossiter et al. (2000a: 545) found that mating was polygynous, but six females gave birth to young fathered by the same male in different years. This seems to indicate fidelity for either mating sites or individuals or from sperm competition. Females start visiting males for mating in autumn, so breeding partnerships are primarily determined by the female. Mating occurs in territory sites occupied by single males, which can be born both within and outside the female's own natal colony. Relatedness between parents, however, was no less than the average recorded for male/female pairs. Gene flow between colonies is likely to be primarily mediated by both female and male dispersal during the mating period rather than more permanent movements. Liu et al. (2013a: 2179) describe the vocal communications during mating in China. They suggest these calls can serve as an indicator for the female bats to recognize the mating male in order to maintain mate fidelity as greater horseshoe bats are sexually segregated before mating. PARASITES: Beaucournu and Kock (1996) mentioned Rhinolophopsylla unipectinata arabs Jordan and Rothschild, 1921 on bats from Algeria. In Europe Genov et al. (1992) reported the presence of the following Nematodes: Molinostrongylus skrjabini Skarbilovitch, 1934 and M. alatus (Ortlepp, 1932). Ševcík et al. (2012: 35) report on the presence of the bat flies Phthiridium biarticulatum Hermann, 1804, Phthiridium biarticulatum Hermann, 1804, and Brachytarsina flavipennis Macquart, 1851 on bats from Crete and Cyprus. Phthiridium biarticulatum was also reported on R. ferrumequinum from Jordan by Benda et al. (2010b: 226), and from Algeria by Bendjeddou et al. (2013: 325). The latter authors also found Brachytarsina flavipennis and Penicillidia dufouri (Westwood, 1825) (Diptera: Nycteribiidae). Bendjeddou et al. (2017: 15) added two additional Nycteribiidae: Nycteribia (Listropoda) schmidlii schmidlii Schiner, 1853 and Phthiridium biarticulatum Hermann, 1804 and one Siphonaptera: Rhinolophopsylla unipectinata arabs Jordan and Rothschild, 1921. Haelewaters et al. (2018: 794) also reported Penicillidia conspicua Speiser, 1901 and P. fulvida, Bigot, 1885, Phthiridium biarticulatum Hermann, 1804, which all might be hosts for the fungus Arthrorhynchus nycteribiae. ACARI Nycteribiidae: Nycteribia kolenati Theodor & Moscona, 1954 was reported by Haelewaters et al. (2017: 1) from R. ferrumequinum from Hungary. African Chiroptera Report 2020 Vermeil (1960) reported two bat flies from Tunisian R. ferrumequinum: Nycteribia biarticulata Hermann, 1804 and Nycteribosca kollari Frauenfeld, 1855. Trombiculidae: Stekolnikov (2018a: 186) mentioned Sasatrombicula cherrata (Taufflieb, 1960) from Oued Cherrat (Morocco). VIRUSES: In Italy, Balboni et al. (2010: 216) reported two R. ferrumequinum specimens, which tested positive for a fragment of RNA-dependent RNA polymerase (RdRp) gene of viruses related to Coronavirus. Phylogenetic analysis showed a close correlation between one of these and SARSrelated CoV belonging to the genus Betacoronavirus (see also de Jong et al., 2011: 10). Nieto-Rabiela et al. (2019: Siuppl.) mentioned the following list of viruses: Adeno associated virus, Astrovirus, Banna virus isolate, Bat calicivirus, Bat circovirus, Bat coronavirus, Bat gammaretrovirus, Bat hepevirus, Bat herpesvirus, Bat mastadenovirus, Bat paramyxovirus, Bat parvovirus, Bat polyomavirus, Bat rhabdovirus, Betacoronavirus, European bat lyssavirus, Hantavirus, Issyk Kul virus, Picornavirus, Puumala virus, Rhinolophus ferrumequinum herpesvirus 1, Rhinolophus ferrumequinum papillomavirus, Rhinolophus ferrumequinum paramyxovirus, Roseolovirus, SARS. Adenoviridae Mastadenovirus Kohl and Kurth (2014: 3111) report this virus from Hungary. Betacoronavirus This virus was reported from bats from Germany and Bulgaria by Kohl and Kurth (2014: 3112). Flaviviridae Flavivirus Japanese encephalitis virus was reported by Luis et al. (2013: suppl.). Orthomyxoviridae Despite sampling and testing of 2 individuals of this species, no evidence of influenza A-like viruses (Orthomyxoviridae) were obtained (Fereidouni et al., 2015). Papillomaviridae Papillomavirus This virus was reported from Spanish bats by Kohl and Kurth (2014: 3113). Rhabdoviridae Lyssavirus - Rabies related viruses de Jong et al. (2011: 9) report the presence of European bat lyssavirus 2. Serra-Cobo et al. (2018: 2) found one seropositive bat among the 47 tested for European bat lyssavirus 1 in Algeria and eight ouf of 67 tested in Morocco. "Various European bat lyssaviruses" were reported by Kohl and Kurth (2014: 3114) from various European countries. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Libya, Morocco, Tunisia. Astroviridae Luis et al. (2013: suppl.) report on the presence of an unassigned astrovirus. Bunyaviridae Hantavirus de Jong et al. (2011: 9), Luis et al. (2013: suppl.) report Hantaan virus occurring on this species, as well as an unassigned Issyk-kul virus (Keterah virus). Coronaviridae - Coronaviruses Luis et al. (2013: suppl.) mentions the presence of Coronavirus BatCoV BB98-15 and SARS coronavirus. Alphacoronavirus Kohl and Kurth (2014: 3112) report this virus to be present in German bats. 309 Figure 90. Distribution of Rhinolophus ferrumequinum 310 ISSN 1990-6471 Rhinolophus fumigatus Rüppell, 1842 *1842. Rhinolophus fumigatus Rüppell, Mus. Senckenberg., 3 (2): 132. Type locality: Ethiopia: Shoa province: Shoa province [ca. 09 00 N 39 00 E] [Goto Description]. Lectotype: SMF 4372: ad, mounted skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell; collection date: 1841. Presented/Donated by: Wilhem Peter Edward Simon Rüppell. See: 451). Mertens (1925: 20). Turni and Kock (2008) mention this specimen (old catalog SMF II.F.7.a) as Lectotype. Cotype: SMF II.F.7b:; collection date: 1841. Presented/Donated by: Wilhem Peter Edward Simon Rüppell. See Andersen (1904c: 451). - Etymology: From the passive Latin perfect participle fumigàtus meaning "smoked", referring to the fur colour of the species (see Lanza et al., 2015: 96). (Current Combination) 1869. Rhinolophus æthiops Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 637 (for 1868). Type locality: Namibia: Damaraland: Otjimbingue [22 15 S 16 10 E]. Syntype: ZMB 3295: ♂, skull and alcoholic. Collected by: Missionary Hahn. See Turni and Kock (2008). Syntype: ZMB 50073: ♀, alcoholic (skull not removed). Collected by: Missionary Hahn. Formerly second specimen with number ZMB 3295; see Turni and Kock (2008: 30). Comments: Turni and Kock (2008: 30) referring to Csorba et al. (2003: 50) mention that ZMB 3295 might possibly represent a species distinct from fumigatus. 1877. Rhinolophus macrocephalus Heuglin, Reise in Nordost Afrika, 2: 22. Publication date: 1877. Type locality: Ethiopia: Tigre province: As(s)am river: Adowa [14 10 N 38 54 E, 1 900 m]. Syntype: SMNS 1059: imm ♀, alcoholic (skull not removed). Collected by: Martin Theodor von Heuglin; collection date: 1862. See Andersen (1904c: 451), Dieterlen et al. (2013: 294). Syntype: SMNS 1059a: imm ♀. Collected by: Martin Theodor von Heuglin. Presented/Donated by: ?: Collector Unknown. Syntype: SMNS 1059b: ad ♂, alcoholic (skull not removed). Collected by: Martin Theodor von Heuglin; collection date: 1862. See Andersen (1904c: 451), Dieterlen et al. (2013 294). 1885. Rhinolophus antinorii Dobson, Ann. Mus. civ. Stor. nat. Genova, (2) 2: 16. Publication date: 22 May 1885. Type locality: Ethiopia: Shoa province: Daimbi. 1905. Rhinolophus fumigatus exsul K. Andersen, Ann. Mag. nat. Hist., ser. 7, 15 (85): 74. Publication date: 1 January 1905. Type locality: Kenya: Kitui [01 22 S 38 01 E, 3 500 ft] [Goto Description]. Holotype: BMNH 1901.5.6.3: ad, skin only. Collected by: Mrs. Hildegarde Hinde; collection date: 3 January 1901; original number: 68. See Andersen (1905a: 74). - Comments: Considered a valid subspecies by Ansell and Dowsett (1988: 32). 1913. Rhinolophus foxi Thomas, Ann. Mag. nat. Hist., ser. 8, 11 (63): 314. Publication date: 1 March 1913. Type locality: Nigeria: Northern Nigeria, Bauchi plateau: Kabwir [09 24 N 09 34 E, 2 500 ft] [Goto Description]. Holotype: BMNH 1913.2.5.1: ad ♂. Collected by: Mr. J.C. Fox and The Cambridge University Mission; collection date: 14 November 1912; original number: 45. Presented/Donated by: Mr. J.C. Fox. 1914. acrotis G.M. Allen, Bull. Mus. comp. Zool., 58 (7): ???. - Comments: Preoccupied by acrotis Heuglin, 1861. 1917. Rhinolophus abæ J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 428. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Aba [03 53 N 30 17 E] [Goto Description]. Holotype: AMNH 49113: ad ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 15 December 1911; original number: 1756. 1939. Rhinolophus aethiops diversus Sanborn, Field Mus. Nat. Hist., Zool. Ser., 24 (5): 42. Publication date: 19 September 1939. Type locality: Senegal: Bakel [14 54 N 12 26 W] [Goto Description]. Holotype: BMNH 1919.7.7.2774: ad ♀, alcoholic (skull not removed). Collection date: 27 September 1887. Formerly in the Lataste collection (see: 42). Comments: Considered a valid subspecies by Grubb et al. (1998: 77), but they also remark that it might be a subspecies of aethiops. ? Rhinolophus abae: (Alternate Spelling) ? Rhinolophus aethiops: (Alternate Spelling) ? Rhinolophus fumigatus abae: (Name Combination) ? Rhinolophus fumigatus aethiops: (Name Combination) ? Rhinolophus fumigatus foxi: (Name Combination) ? Rhinolophus fumigatus fumigatus: (Name Combination) African Chiroptera Report 2020 311 TAXONOMY: Does not include eloquens or perauritus, but does include aethiops; see Koopman (1975: 389 - 390). fumigatus species group (Csorba et al., 2003: 48 51; Simmons, 2005: 356). See Csorba et al. (2003: 49 - 50) for remarks on taxonomy, e.g. that fumigatus and aethiops are probably different species. CONSERVATION ACTIONS: Jacobs et al. (2008al) [in IUCN (2009)] and Monadjem et al. (2017be) reported that in view of the species wide range in East and southern Africa, it seems probable that it is present in several protected areas. No direct conservation measures are currently needed for this species as a whole. Jacobs et al. (2016g: 94) refer to Dool et al. (2016b), who found that R. fumigatus from the western part of Southern Africa seems to be a distinct but sister lineage to R. fumigatus from the eastern half of Africa. GENERAL DISTRIBUTION: Rhinolophus fumigatus is widely, but patchily, recorded over much of sub-Saharan Africa. It ranges from Senegal and The Gambia in the east, through West and Central Africa to Ethiopia and Eritrea in the east, and from here through East and southern Africa as far south as Namibia and northeastern South Africa. COMMON NAMES: Afrikaans: Rüppell se saalneusvlermuis, Rüppellsaalneusvlermuis. Chinese: 达 马 拉 菊 头 蝠 . Czech: vrápenec kouřový. English: Rüppell's Horseshoe Bat, Smoky Horseshoe Bat, Abyssinian Horseshoe Bat. French: Rhinolophe de Rüppell. German: Rüppells Hufeisennase, Rüppells Hufeisennase, Shoa Kamnase. Italian: Rinòlofo affumicàto, Fèrro di cavàllo affumicàto. Portuguese: Morcego ferradura da Damarlandia. ETYMOLOGY OF COMMON NAME: The colloquial name refers to the describer of the species (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008al; IUCN, 2009; Monadjem et al., 2017be). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017be). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008al; IUCN, 2009). 2004: LC ver 3.1 (2001) (IUCN, 2004; Jacobs et al., 2004aj). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016k). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: There appear to be no major threats to Rhinolophus fumigatus as a whole (Jacobs et al., 2008al; IUCN, 2009; Monadjem et al., 2017be). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,246,343 km 2. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 558; Taylor et al., 2018b: 62); Benin (Capo-Chichi et al., 2004: 162); Botswana (Cotterill, 2004a: 261); Burkina Faso (Kangoyé et al., 2015a: 610); Burundi; Cameroon; Central African Republic; Chad; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 558); Côte d'Ivoire; Eritrea; Ethiopia; Gabon; Gambia; Ghana; Guinea; Kenya; Liberia; Malawi (Happold et al., 1988; Monadjem et al., 2010d: 558); Mali (Meinig, 2000: 105); Mauritania; Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 558; Monadjem et al., 2010c: 378); Namibia (Monadjem et al., 2010d: 559); Niger; Nigeria; Rwanda; Senegal; Sierra Leone; Somalia; South Africa (Monadjem et al., 2010d: 559); Sudan; Tanzania (Stanley and Goodman, 2011: 42); Togo; Uganda; Zambia (Ansell, 1978; Monadjem et al., 2010d: 559); Zimbabwe (Monadjem et al., 2010d: 559). SEXUAL DIMORPHISM: Kangoyé et al. (2015a: 610) found no differences in body measurement of males and females from Burkina Faso, but the cranial measurements of the males were on average larger than those of the females. ECHOLOCATION: Jacobs et al. (2007a) reported a peak frequency of 53.8 (± 0.2) kHz for individuals South Africa. Monadjem et al. (2010c: 378) reported from Mozambique the peak echolocation frequencies of a single male was 54 kHz (ANABAT). 1 call of tye FM/CF/FM was recorded by Manga Mongombe (2012: 79) from Maroua (Cameroon) 312 ISSN 1990-6471 with the following parameters: Fmax: 62.6 kHz, Fmin: 61.6 kHz, Fmean: 61.9 kHz, Fknee: 61.7 kHZ, Fchar: 61.6 kHz, and duration: 0.57 msec. Kangoyé et al. (2015a: 610) reported that five males from Burkina Faso called at 54.2 ± 0.4 (53.454.4) kHz. In Sierra Leone, a flying bat was recorded by Weber et al. (2019: 21) at 51.5 kHz. For two calls from the Aberdares Range in Kenya, Eisenring et al. (2016: SI 2) gave the following values: PF: 57.9 ± 1.1 (57.2 - 58.7), HF: 106.7 ± 4.3 (103.7 - 109.8), LF: 54.3 ± 0.3 (54.0 - 54.5), DT: 0.0 ± 0.0 (0.0 - 0.1), DF: 52.5 ± 4.7 (49.2 55.8), IPI: 0.7 ± 0.1 (0.6 - 0.8). For a bat from the Kalahari Desert, Adams and Kwiecinski (2018: 4) reported: Fchar: 53.9 kHz, Fmax: 54.4 kHz, Fmin: 53.6 kHz, and duration: 40.1 msec. Weier et al. (2020: Suppl.) reported on four calls from the Okavango River Basin with the following characteristics: Fmax: 53.19 ± 0.60 kHz, Fmin: 51.44 ± 1.67 kHz, Fknee: 52.59 ± 0.78 kHz, Fchar: 52.74 ± 0.66 kHz, slope: 0.31 ± 5.49 Sc, duration: 10.24 ± 5.89 msec. Jacobs (2016: 119) indicates that the second harmonic of this bat coincides with the third harmonic of R. hildebrandtii, which possibly might illustrate "harmonic hopping". MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach (1986) reported 2n = 58, FN = 60, BA = 4, a subtelocentric X chromosome, and a metacentric Y. Koubínová et al. (2010b: 394) reported for Senegal similar results to Rautenbach (1986), but there were only two pairs of small meta- and submetacentric autosomes, and the remaining 26 pairs were acrocentric, a medium sized acrocentric (no. 15) had an achromatic gap near the centromere, while the X was a medium sized subtelocentric, and the Y was dot-like. Koubínová (2013: 7) reported the karyotype as: 2n = 58, FNa = 60, FN = 64. Protein / allozyme: Unknown. ROOST: Lelant and Chenaval (2011d: 13) report on a colony of over 1,000 animals inhabiting a very old Baobab tree (Adansonia digitata) in Fadgal, Senegal. POPULATION: Structure and Density:- It is a locally common species that can be encountered as large colonies. Colonies in southern Africa tend to be smaller (Jacobs et al., 2008al; IUCN, 2009; Monadjem et al., 2017be). Trend:- 2016: Unknown (Monadjem et al., 2017be). 2008: Unknown (Jacobs et al., 2008al; IUCN, 2009). REPRODUCTION AND ONTOGENY: Happold and Happold (1990b: 566) reported births in Malawi early in the wet season, and females were in "fairly close reproductive synchrony". A pregnant female was collected in August 1992 in the East Usambara Mountains (Tanzania) (Stanley and Goodman, 2011: 42). PARASITES: HAEMOSPORIDA Lutz et al. (2016: 9) examined 5 R. fumigatus specimens from East Africa and found two of them infected with Nycteria sp. Rosskopf et al. (2018: Suppl.) mentioned the (same?) record from Malawi. Fain (1959a: 340) mentions that one of the paratypes of Nycteridocoptes eyndhoveni Fain, 1959 (Acari: Sarcoptidae) was collected from a "R. aethiops" specimen Kambisa, Angola. HEMIPTERA: Polyctenidae: Androctenes horvathi Jordan, 1912 recorded from Transvaal, South Africa (Haeselbarth et al., 1966: 17, host referred to as R. eloquens). DIPTERA: Streblidae: Ascodipteron brevior Maa, 1965, female cysts found at the base of the ears in Congo, Namibia and South Africa (Haeselbarth et al., 1966: 106). Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100, host refered to as R. eloquens). Raymondia intermedia Jobling, 1936 (Shapiro et al., 2016: 255). Nycteribiidae: Nycteribia scissa sudanica Theodor, 1957 from Ethiopia, the Sudan and the Congo (Haeselbarth et al., 1966: 108). Penicillidia fulvida (Bigot ,1885) (Haeselbarth et al., 1966: 114, host referred to as R. foxi). SIPHONAPTERA: Ischnopsyllidae: Rhinolophopsylla ectopa (Jordan, 1937) was reported on "R. aethiops" from Kenya by Beaucournu and Kock (1996). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Rhinolophus fumigatus: Coronaviruses and Paramyxoviruses. African Chiroptera Report 2020 313 Coronaviridae - Coronaviruses Alphacoronavirus SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in 204 individuals from Oriental Province, DRC, one tested positive (1/204, 0.5 %). UTILISATION: Lelant and Chenaval (2012: 15) report on the use of R. fumigatus specimens in traditional medicine in Senegal. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Burkina Faso, Burundi, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Eritrea, Ethiopia, Gabon, Ghana, Guinea, Kenya, Malawi, Mauritania, Morocco, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, South Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. Figure 91. Distribution of Rhinolophus fumigatus Rhinolophus gorongosae Taylor, MacDonald, Goodman, Kearney, Cotterill, Stoffberg, Monadjem, Schoeman, Guyton, Naskrecki and Richards, 2018 *2018. Rhinolophus gorongosae Taylor, MacDonald, Goodman, Kearney, Cotterill, Stoffberg, Monadjem, Schoeman, Guyton, Naskrecki and Richards, Zool. J. Linn. Soc., 184 (1): 1249, 1262, figs 6, 8, 9, 10, 12. Publication date: 24 April 2018. Type locality: Mozambique: Sofala province: Gorongosa National Park: Bunga Inselberg [18 35 56 S 34 20 35 E, 212 m] [Goto Description]. Holotype: DNSM 14820: ad ♂, skull and alcoholic. Collected by: Jennifer Anna Guyton; collection date: 25 April 2015; original number: JAG 196. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14815: ♀. Collected by: Jennifer Anna Guyton; collection date: 24-25 April 2015; original number: JAG 188. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14816: Collected by: Jennifer Anna Guyton; collection date: 24-25 April 2015. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14817: Collected by: Jennifer Anna Guyton; collection date: 24-25 April 2015. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14818: Collected by: Jennifer Anna Guyton; collection date: 24-25 April 2015. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14819: Collected by: Jennifer Anna Guyton; collection date: 24-25 April 2015. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14828: Collected by: Jennifer Anna Guyton; collection date: 2 May 2015. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14843: ♀. Collected by: Jennifer Anna Guyton; collection date: 5 November 2015; original number: JAG 228. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 14865: Collected by: Jennifer Anna Guyton; collection date: 22 July 2015. Presented/Donated by: ?: Collector Unknown. - Etymology: The name refers to the Gorongosa district (Mozambique) and the Gorongose National Park, where the type specimen was collected (Taylor et al. (2018a: 14). TAXONOMY: Demos et al. (2019a: 11) resequenced the cyt-b of some of the specimens used by Taylor et al. (2018a) in the description of the species and found that this diverged only between 1 and 1.4 % from that of R. simulator and R. rhodesiae. This "strongly suggests" that the genetic arguments for the newly described R. gorongosae were based on sequencing errors. COMMON NAMES: English: Least horseshoe Gorongosa-Hufeisennase. bat. German: 314 ISSN 1990-6471 GENERAL DISTRIBUTION: This species has only been found in the Gorongosa National Park (Mozambique). Taylor et al. (2018a: 17) tentatively assigned a specimen from Mount Inago (Mozambique) to this species, but this specimen needs to be examined further. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Taylor et al. (2018a: 14) indicate that R. gorongosae is the smallest rhinolophid in the Ethiopian region, even smaller than R. denti. ECHOLOCATION: Taylor et al. (2018a: 17) report a echolocation CF of 106 (104 - 108) kHz. peak DISTRIBUTION BASED ON VOUCHER SPECIMENS: Mozambique. Figure 92. Distribution of Rhinolophus gorongosae Rhinolophus guineensis Eisentraut, 1960 *1960. Rhinolophus landeri guineensis Eisentraut, Stuttg. Beitr. Naturk., 39: 1, fig. 1. Publication date: 1 June 1960. Type locality: Guinea: foot of Kelesi Plateau: Tahiré [ca. 10 05 N 12 30 W, 500 m] [Goto Description]. Holotype: SMNS 6103: ad ♂, skin and skull. Collected by: Hans Knorr; collection date: 5 October 1956. Presented/Donated by: ?: Collector Unknown. Paratype: SMNS 6104: ad, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: SMNS 6105: ad, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: SMNS 6106: ad, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: SMNS 6107: ad, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: SMNS 6108: ad, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: SMNS 6109: ad, alcoholic (skull not removed). Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: SMNS 6113: ad, skin and skull. Collected by: Hans Knorr; collection date: 19 November 1956. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) mention Salung Plateau near Nyembaro, 10 km W of Kolenté, Guinea as locality. Paratype: ZFMK MAM-1959.0176: ♀, skin and skull. Collected by: Hans Knorr; collection date: 5 October 1956; original number: 29. Presented/Donated by: ?: Collector Unknown. Paratype: ZFMK MAM-1959.0177: ♂, skin and skull. Collected by: Hans Knorr; collection date: 23 November 1956; original number: 137. See Hutterer (1984: 30), Hutterer and Peters (2010: 11). - Etymology: Refers to country from which the type specimen was collected. ? Rhinolophus guineensis: (Name Combination, Current Combination) TAXONOMY: Meester et al. (1986) state that Rosevear (1965) and Kock (1969a) regard guineensis Eisentraut, 1960, from Guinea, as subspecifically distinct from landeri. Hayman and Hill (1971), on the other hand, consider it a synonym, while Böhme and African Chiroptera Report 2020 Hutterer (1979), Csorba et al. (2003: 59 - 61) and Simmons (2005: 356) regard it as a distinct species, sympatric with landeri and somewhat larger. 315 Senegal, Guinea, Sierra Leone, Liberia and Côte d'Ivoire. It is a highland species recorded at elevations of 1,400 m asl and over. It may be present in Gambia in remnant patches of forest (Grubb et al., 1998). COMMON NAMES: Czech: vrápenec guinejský. English: Guinean Horseshoe Bat (Csorba et al., 2003: 59 - 61; Simmons, 2005), Senegal Horseshoe Bat. French: Rhinolophe de Guinée, Rhinolophe du Sénégal. German: Guinea-Hufeisennase. Native: Côte d'Ivoire; Guinea (Denys et al., 2013: 281; Decher et al., 2016: 264); Liberia (Simmons, 2005); Senegal; Sierra Leone (Csorba et al., 2003). Presence uncertain: Gambia (Grubb et al., 1998). SIMILAR SPECIES: See Csorba et al. (2003). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Csorba et al. (2003). CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B2ab(ii,iii) ver 3.1 (2001)) because its area of occupancy is probably less than 2,000 km2 (roosting caves), its distribution is severely fragmented, and there is continuing decline in the extent and quality of its forest and cave habitats (Fahr, 2008j; IUCN, 2009). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Csorba et al. (2003). Assessment History Global 2008: VU B2ab(ii,iii) ver 3.1 (2001) (Fahr, 2008j; IUCN, 2009). 2004: VU B2ab(iii) ver 3.1 (2001) (Fahr, 2004g; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). ECHOLOCATION: In Guinea, Fahr and Ebigbo (2003) reported the frequency of the CF-component at 85.3 - 85.4 kHz. Regional None known. HABITAT: Monadjem et al. (2016y: 366) recorded this species from higher altitudes (above 900 m) in the Mount Nimba area. MAJOR THREATS: Major threats to Rhinolophus guineensis include deforestation resulting from logging operations, the conversion of land to agricultural use, and mining activities. There is also a limited threat of overhunting for the bushmeat trade (Fahr, 2008j; IUCN, 2009). Sagot and Chaverri (2015: 1670) mention roost loss or disturbance, habitat degradation or loss, and hunting as major threats for this species. CONSERVATION ACTIONS: Fahr (2008j) [in IUCN (2009)] reported Rhinolophus guineensis has been recorded from the Mount Nimba World Heritage Site and the "Massif du Ziama" Biosphere Reserve, both in Guinea. It has also been recorded from a few state forests ("Forets Classees") in Guinea. There is a need to conserve remaining areas of suitable habitat for this species. Further studies are needed to better determine the species range. GENERAL DISTRIBUTION: Rhinolophus guineensis is a West African bat that has been patchily recorded from southern DETAILED MORPHOLOGY: Baculum - Unknown For a description of the crania, teeth, and noseleaves see Csorba et al. (2003). In Sierra Leone, one female called with a CFcomponent at 82.0 kHz (Weber et al., 2019: 21). See also Csorba et al. (2003). HABITS: See Csorba et al. (2003). ROOST: Weber and Fahr (2006: 4) indicate that R. guineensis largely or exclusively depends on the availability of caves as day roosts in Guinea. POPULATION: Structure and Density:- Rhinolophus guineensis is possibly a rare species overall, although little information is available on the population abundance of this bat (Fahr, 2008j; IUCN, 2009). Trend:- 2008: Unknown (Fahr, 2008j; IUCN, 2009). REPRODUCTION AND ONTOGENY: Weber et al. (2019: 21) recorded a pregnant female on 30 March in Sierra Leone. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Guinea, Liberia, Nigeria. 316 ISSN 1990-6471 Figure 93. Distribution of Rhinolophus guineensis Rhinolophus hildebrandtii Peters, 1878 *1878. Rhinolophus Hildebrandtii Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 195, pl. 1, fig. 1, 1a. Type locality: Kenya: Taita district: Ndi [03 14 S 38 30 E, 2 000 m] [Goto Description]. Syntype: BMNH 1879.1.21.1: ad. Turni and Kock (2008: 33). Reject the holotype status for this specimen and consider this also as a possible syntype. Syntype: ZMB 5267: ad ♀, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008: 33). Syntype: ZMB 5378: ♂, skin and skull. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008: 33). Syntype: ZMB 5379: ♂, skin and skull. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008: 33). Comments: Turni and Kock (2008: 33) reject the holotype status for the BMNH specimen and consider this also as a syntype. - Etymology: In honour of Johannes Maria Hildebrandt (1847 - 1881), a German naturalist, explorer and linguist, who collected a large number of plants and animals in eastern Africa, Madagascar, Comoro islands and Arabia (see Pearl, 1994: 3; Taylor, 2005, Lanza et al., 2015: 101). (Current Combination) 1991. Rhinolophus hildebrantii: Avery, Durban Mus. Novit., 16: 1. (Lapsus) ? Rhinolophus hildebrandti hildebrandti: (Name Combination, Alternate Spelling) ? Rhinolophus hildebrandti: (Name Combination) ? Rhinolophus hildebrandtii: (Current Spelling) TAXONOMY: See Pearl (1994). Within the fumigatus species group (Csorba et al., 2003: 51 - 53; Simmons, 2005: 356). See Csorba et al., 2003: 52 ) for a review of the taxonomy. Taylor et al. (2010: 297) indicate that cryptic species might be present in the southern African R. hildebrandtii complex. These species probably diverged during the late Neogene (PlioPleistocene) when more open woodlands replaced the seasonal woodland/forest habitats (Schoeman and Monadjem (2018: 4). COMMON NAMES: Afrikaans: Hildebrandt se saalneusvlermuis, Hildebrandt-saalneusvlermuis, Hildebrandtse Vlermuis. Chinese: 希 氏 菊 头 蝠 . Czech: vrápenec Hildebrandtův. English: Hildebrandt's Horseshoe Bat. French: Rhinolophe d'Hildebrandt, Fer à cheval d'Hildebrant, Fer à cheval du Kenya. German: Hildebrandts Hufeisennase. Italian: Rinòlofo di Hildebrandt, Fèrro di cavàllo di Hildebrandt. Kiluba (DRC): Kasusu. Portuguese: Morcego ferradura de Hildebrandt. Yao (Malawi): Lichinji (applied to all largish bats). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: R. hildebrandtii was found in the earlier half of the upper Pleistocene sequence at Border Cave, KwaZulu-Natal, South Africa (Avery, 1991: 6; Taylor, 1998: 35). African Chiroptera Report 2020 CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs, 2008h; IUCN, 2009; Monadjem and Jacobs, 2017c). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Jacobs, 2017c). 2008: LC ver 3.1 (2001) assessed as Rhinolophus hildebrandti (Jacobs, 2008h; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs, 2004i; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Jacobs (2008h) [in IUCN (2009)] and Monadjem and Jacobs (2017c) reported there appear to be no major threats to Rhinolophus hildebrantii as a whole. CONSERVATION ACTIONS: Jacobs (2008h) [in IUCN (2009)] and Monadjem and Jacobs (2017c) reported there appear to be no direct conservation measures in place. In view of the species wide East African and southern African range, it seems probable that the species is present in some protected areas. GENERAL DISTRIBUTION: Rhinolophus hildebrantii is largely endemic to East Africa, except for a Central African records from Rwanda and southern Democratic Republic of the Congo, and a single questionable West African record from Nigeria. In East Africa it ranges from Ethiopia and northern Sudan in the north, through Uganda, Kenya and Tanzania (including the island of Pemba [see O'Brien, 2011: 286]), into southern Africa of Zambia, Malawi, Mozambique, Zimbabwe and Botswana, being found as far south as northern South Africa. Taylor (1998: 35) mentioned Pleistocene deposits of this species reported by Avery (1991) from Border Cave, northern Maputaland, KwaZulu-Natal, South Africa. In South Africa, its distribution is mostly affected by precipitation seasonality (Babiker Salata, 2012: 49). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 946,716 km2. 317 Native: Botswana (Monadjem et al., 2010d: 559); Burundi; Congo (The Democratic Republic of the) (Hayman et al., 1966; Ansell, 1974; Monadjem et al., 2010d: 559); Ethiopia (Kruskop et al., 2014: 102 report the first record North of the Rift Valley); Kenya; Malawi (Ansell, 1974; Happold et al., 1988; Happold and Happold, 1997b: 818; Monadjem et al., 2010d: 559); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 559; Monadjem et al., 2010c: 379); Rwanda; Somalia; South Africa (Monadjem et al., 2010d: 559); Sudan; Tanzania (Stanley and Goodman, 2011: 42), United Republic of; Uganda (Kityo and Kerbis, 1996: 61); Zambia (Ansell, 1969; Monadjem et al., 2010d: 559); Zimbabwe (Monadjem et al., 2010d: 559). Presence uncertain: Namibia (Cotterill, 2004a: 261); Nigeria. FUNCTIONAL MORPHOLOGY: Aylward et al. (2019: ) refer to Makanya and Maina (1994), who reported honeycomb ridge-like villi in the proximal part of the bat's intestine. ECHOLOCATION: Taylor et al. (2005), using a Pettersson D980, reported three, slightly different maximum frequencies (42.3 (± 0.2) kHz, 43.2 (± 0.1) kHz, 42.4 (0) kHz) from three different localities in Kenya for individuals of R. cf. hildebrandtii. Jacobs (1996) reported a maximum frequency of 37 - 46 kHz for individuals at Sengwa in Zimbabwe. Aldridge and Rautenbach (1987) reported a maximum frequency of c. 40 kHz for individuals from Pafuri in South Africa. Jacobs et al. (2007a) reported a peak frequency of 33.0 ± 0.9 kHz for individuals in South Africa (detector type and localities not reported). From Mozambique Monadjem et al. (2010c: 379) reported that the peak echolocation frequencies ranged between 35 - 40 kHz (ANABAT, Petterson D240x). At Uzungwa Scarp Forest Reserve, Tanzania, Trentin and Rovero (2011: 56) describe the calls as starting at 28.5 kHz, increasing in .5 msecs to 39.8 kHz, and after 5 msecs decreasing to 26.7 kHz. The maximum intensity is at 35.5 kHz. Luo et al. (2019a: Supp.) reported the following data (Hand released bats, two calls): Fpeak: 42.8, 42.1 kHz and duration: 57.7, 45.2 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Both Peterson and Nagorsen (1975) and Rautenbach (1986) reported 2n = 58, FN = 60, BA = 4, and subtelocentric X and Y chromosomes. Protein / allozyme - Unknown. 318 ISSN 1990-6471 PREDATORS: Schätti (1984: 337) refers to Broadley (in lit.) who mentions a R. hildebrandtii specimen having been swallowed by an African housesnake (Lamprophis fuliginosus (Boie, 1827)). Theodor (1968) recorded Raymondia setiloba (Jobling, 1954) from "R. hildebrandtii" in the DRC, but Shapiro et al. (2016: 256) indicate that the identify of the host is uncertain, and therefore report it as Rhinolophus sp. POPULATION: Structure and Density:- This species is quite common in Zambia (Jacobs, 2008h; IUCN, 2009; Monadjem and Jacobs, 2017c). ACARI Sarcoptidae: Fain (1959a: 341) reports on Nycteridocoptes eyndhoveni Fain, 1959 collected on a bat of this species captured in Lubudi, DRC. Trend:- 2016: Unknown (Monadjem and Jacobs, 2017c). 2008: Unknown (Jacobs, 2008h; IUCN, 2009). Trombiculidae: Stekolnikov (2018a: 116) found Trisetica aethiopica (Hirst, 1926). Stekolnikov and Quetglas (2019: 8) indicate this bat was captured in South Sudan. REPRODUCTION AND ONTOGENY: A male with abdominal testes was collected in the West Usambara Mountains (Tanzania) on 15 July 1991 (Stanley and Goodman, 2011: 42). Trentin and Rovero (2011: 50) reported two lactating females, and six females with single youngs, netted in December 2005 at Uzungwa Scarp Forest Reserve, Tanzania. In Malawi, young are born around June (dry season) (Happold and Happold, 1990b: 566). PARASITES: HAEMOSPORIDA Schaer et al. (2015: 381) found Nycteria sp. parasites in R. hildebrandtii from Kenya. Lutz et al. (2016: 9) found eight out of 14 specimens from East Africa to be infected by Nycteria sp. Perkins and Schaer (2016: Suppl.) report on the presence of Nycteria congolensis (Garnham, 1966) on bats from the DRC and East Africa as well as of Nycteria sp. from Kenya and Malawi or Mozambique. VIRUSES: Coronaviridae 31.3 % (5 out of 16) of the Kenyan bats tested by Tao et al. (2017: 3) were positive for CoV (genus Betacoronavirus, subgenus Sarbecovirus, see Markotter et al., 2020: 6). Rhabdoviridae Vesiculovirus Metselaar et al. (1969: 183) extracted a new virus (the "Mount Elgon Bat Virus") from the salivary glands of R. hildebrandtii eloquens, collected at the Kiminimi caves on the slopes of Mount Elgon, Kenya. Willoughby et al. (2017: Suppl.) mentions this virus as "Mount Elgon bat ledantevirus". DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Burundi, Congo (Democratic Republic of the), Ethiopia, Kenya, Nigeria, South Sudan, Sudan, Tanzania, Uganda. DIPTERA Nycteribiidae: Nycteribia scissa Speiser; Nycteribia rotundata Theodor; Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114). Streblidae: Ascodipteron brevior Maa 1965, female cysts found at the base of the ears in Congo (Haeselbarth et al., 1966: 106). Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Raymondia intermedia Jobling, 1936 in the Congo (Haeselbarth et al., 1966: 102). Raymondia waterstoni Jobling, 1931 (Haeselbarth et al., 1966: 104), these records may possibly also be associated with R. smithersii (see Shapiro et al., 2016: 256). Raymondia intermedia Jobling, 1936 (Shapiro et al., 2016: 255). Figure 94. Distribution of Rhinolophus hildebrandtii African Chiroptera Report 2020 319 Rhinolophus hilli Aellen, 1973 *1973. Rhinolophus hilli Aellen, Period. biol., 75 (1): 101. Type locality: Rwanda: Cyangugu province: Uwinka [02 29 S 29 12 E, 2 300 m] [Goto Description]. Holotype: ZMZ 126639: ad ♀, skull and alcoholic. Collected by: U. Goepel; collection date: 25 August 1964. (Current Combination) TAXONOMY: Previously recognized as a synonym of R. ruwenzorii J. Eric Hill, 1942 but recognized as a separate species by Fahr et al. (2002: 108) and Simmons (2005). COMMON NAMES: Czech: vrápenec Hillův. English: Hill's Horseshoe Bat. French: Rhinolophe de Hill. German: Hills Hufeisennase. CONSERVATION STATUS: Global Justification Listed as Critically Endangered (CR B1ab(iii,v)+2ab(iii,v) ver 3.1 (2001)) because its extent of occurrence appears to be less than 100 km2 and its area of occupancy (presumably cave roosting sites) is probably less than 10 km 2 (field surveys in the Albertine rift have not found the species elsewhere), all individuals are in a single location (Nyungwe National Park), there is continuing decline in the extent and quality of its habitat, and there is believed to be an ongoing population decline related to over-harvesting for food (Fahr, 2008k; IUCN, 2009; Fahr, 2010). Assessment History Global 2010: CE B1ab(iii,v)+2ab(iii,v) ver 3.1 (2001) (Fahr, 2010). 2008: CR B1ab(iii,v)+2ab(iii,v) ver 3.1 (2001) (Fahr, 2008k; IUCN, 2009). 2004: CR (Fahr, 2004f; IUCN, 2004). needed to conserve the habitat of this species. There is a need for bat awareness programmes for local people to prevent over-harvesting of the species. Further field surveys are also needed to locate important roosts and to learn more about the natural history of this poorly known species. GENERAL DISTRIBUTION: Rhinolophus hilli is known only within the Nyunwe National Park, Rwanda. It has been identified from only two close localities (approximately 8 km apart). Field surveys in surrounding areas of potentially suitable habitat have not found additional populations. It has an elevational range of between 1,750 and 2,512 m asl (Fahr et al., 2002). Native: Rwanda. POPULATION: Structure and Density:- There is little information available on the abundance of Rhinolophus hilli; colonies of this species have yet to be found but are likely to be small (Fahr et al., 2002; Fahr, 2010). Trend:- 2010: Decreasing (Fahr, 2010). 2008: Decreasing (Fahr, 2008k; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Rwanda. Regional None known. MAJOR THREATS: Fahr et al. (2002) indicate that while there is little information available on this seemingly limited range species, it is probably threatened by habitat destruction (presumably by logging operations and the conversion of land to agricultural use) and direct exploitation (for subsistence food) in their day roosts (see Fahr, 2008k) [in IUCN (2009)]; Fahr, 2010). CONSERVATION ACTIONS: Fahr (2010) repreats what stated by Fahr (2008k) [in IUCN (2009)] who reported that although Rhinolophus hilli is present within the Nyungwe National Park, it seems additional measures are Figure 95. Distribution of Rhinolophus hilli 320 ISSN 1990-6471 Rhinolophus hillorum Koopman, 1989 *1989. Rhinolophus clivosus hillorum Koopman, Am. Mus. Novit., 2946: 4. Publication date: 28 June 1989. Type locality: Liberia: Lofa county: ca 2 mi SW Voinjama, near Zozoma: John Hegbe farm: (the type lable mentions John's Town) [08 25 N 09 35 W, 500 m] [Goto Description]. Holotype: AMNH 257044: ad ♀, skull and alcoholic. Collected by: Andrew S. Voros; collection date: early October; original number: 5924. Type lable mentions 3 September 1983. - Etymology: In honour of the two (often confused) eminent mammalogists named John E. Hill: J. Eric (1907-1947), and J. Edwards (1928-1997) (see Koopman, 1989b: 5). ? Rhinolophus hillorum: (Name Combination, Current Combination) TAXONOMY: Considered a synonym of clivosus by Koopman (1993a: 164), but see Simmons (2005: 356). See Csorba et al. (2003: 131) for remarks on taxonomy. COMMON NAMES: Czech: vrápenec západoafrický. English: Upland Horseshoe bat, Hill's Horseshoe bat. French: Rhinolophe de Lofa. German: HochlandHufeisennase. CONSERVATION STATUS: Global Justification Listed as Near Threatened since although its Extent of Occurrence is probably less than 20,000 kmsuper 2 and its habitat is declining from the loss of montane habitat, especially in northern Liberia and southern Guinea, making the species close to qualifying for Vulnerable (Jacobs et al., 2010). Assessment History Global 2010: NT ver 3.1 (2001) (Jacobs et al., 2010). 2008: VU A2c ver 3.1 (2001) (Jacobs et al., 2008ad; IUCN, 2009). 2004: VU A4c; B2ab(iii) ver 3.1 (2001) (Jacobs et al., 2004s; IUCN, 2004). Regional None known. MAJOR THREATS: Colonies are threatened by general deforestation, often resulting from logging and mining operations, and overharvesting for the bushmeat trade (Jacobs et al., 2008ad; IUCN, 2009; Jacobs et al., 2010). CONSERVATION ACTIONS: Jacobs et al. (2010) reprats what is stated by Jacobs et al. (2008ad) [in IUCN (2009)] who reported there appear to be no direct conservation measures in place for Rhinolophus hillorum. It is not known if the species is present in any protected areas, but it has been recorded from the Sapoba Forest Reserve in Nigeria and the Bali Forest Reserve in Cameroon (Cotterill, 2002a). There is a need to identify and protect important areas for this species. Further research is needed into the species distribution, including the location of any additional colonies. GENERAL DISTRIBUTION: Rhinolphus hillorum is largely endemic to West and Central Africa, with a single questionable record from northern Uganda. It has been recorded from a few small, disjunct, colonies in Liberia (including Voinjama; Tokadeh; and the Wonegizi Mountains), Guinea, Nigeria (Sapoba Forest Reserve) and Cameroon (including Lake Manenguba and the Bali Forest Reserve). Native: Cameroon; Guinea (Fahr et al., 2006a: 72); Liberia; Nigeria. Presence uncertain: Uganda. HABITAT: Monadjem et al. (2016y: 366) recorded five specimens from forested habitats on the Liberian side of Mount Nimba, on altitudes ranging from 780 to 1,200 m. ROOST: Monadjem et al. (2016y: 366) found a roost in an old mine adit at Mount Tokadeh, Liberia. POPULATION: Structure and Density:- The species appears to have a small population, but further research is needed to confirm this (Jacobs et al., 2008ad; IUCN, 2009; R,6206). 2008: Decreasing (Jacobs et al., 2008ad; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Guinea, Kenya, Liberia, Nigeria. African Chiroptera Report 2020 321 Figure 96. Distribution of Rhinolophus hillorum Rhinolophus hipposideros (Bechstein, 1800) 1792. minor Kerr, in: Linnaeus, Animal Kingdom, 1 (1): xvii, 91. Publication date: February 1792. Type locality: France: "France". - Comments: Preoccupied by minor Kerr, 1792: 97 (two pages before this) (see Simmons, 2005). *1800. Vespertilio hipposideros Bechstein, Allgemeine Uebersicht Vierfüssige Thiere, 2: 629. Type locality: France: "France". - Etymology: from the masculine Greek substantive "ἴππος" (híppos), meaning "horse" and from the neuter Greek substantive "σίδηρος" (síderos), meaning "iron", referring to the occurrence of a fleshy horseshoe-shaped expansion on the muzzle in this species as in all other species of the genus (see Lanza et al., 2015: 104). 1861. Rhinolophus miminus Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 6. Publication date: 1861. Type locality: Eritrea: Kérén [ca. 15 45 N 38 20 E]. Holotype: SMNS 987: sad ♂, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Andersen (1904c: 455) mentions it as an immature, but full-grown specimen and 1862? as collection date. Dieterlen et al. (2013: 294) mention "skull and specimen in alcohol" and mention 1861 as collection date. 1870. Rhinolophus Eggenhöffner Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 62 (1): 151. - Comments: This might be an African form, but we do not have any further information. 1905. Rhinolophus hipposiderus typicus K. Andersen, Proc. zool. Soc. Lond., 1905, II, I (10): 141. Publication date: 7 October 1905. - Comments: This might be an African form, but we lack any further information. 1918. [Rhinolophus hipposideros] escaleræ K. Andersen, Ann. Mag. nat. Hist., ser. 9, 2 (10): 378. Publication date: 1 October 1918. Type locality: Morocco: Mogador [=Essaouira]: Ha-ha [ca. 31 31 N 09 46 W] [Goto Description]. Holotype: BMNH 1910.11.24.2: ♀. Collected by: D. Manuel M. de la Escalera. Presented/Donated by: Michael Rodgers Oldfield Thomas. See Andersen (1918: 378). 1937. Rhinolophus hipposideros vespa Laurent, Bull. Soc. Hist. nat. Afr. Nord, 28: 157, text-fig. 1 - 2. Publication date: February 1937. Type locality: Morocco: near Rabat: Oued Korifla [ca. 34 02 N 06 50 W]. Holotype: MNHN 919: ♂, alcoholic (skull not removed). Collected by: Prof. Dr. R. Laurent; collection date: 8 August 1927. Number 181 in Rode (1941: 238). Paratype: MNHN 833: ♂, alcoholic (skull missing). Collected by: Prof. Dr. R. Laurent. Number 181a in Rode (1941: 238). 1978. eggenhoffner: Corbet, Mammals Palaearctic region, 43. - Comments: Possibly extralimital. (Lapsus) ? Rhinolophus hipposideros minimus: (Name Combination) ? Rhinolophus hipposideros: (Name Combination, Current Combination) 322 ISSN 1990-6471 TAXONOMY: Revised by Felten et al. (1977). The status of populations around the Mediterranean, on Mediterranean Islands, and on both sides of the Red Sea is unclear (see Horácek et al., 2000: 101). hipposideros species group (Csorba et al., 2003: 53 - 55; Simmons, 2005: 356 - 357). See Csorba et al. (2003: 54) for review of taxonomy. Kozhurina (2009: 72) points out that Borkhausen (1797 - Deutsche Fauna, oder kurzgefaßte Naturgeschichte der Thiere Deutschlands. Erster Theil: Säugethiere und Vögel. Frankfurt am Main: 85) was the first to give a scientific name to this species: Noctilio Hipposideros. COMMON NAMES: Albanian: Lakuriq nate hundpatkua i vogël. Arabian: Khafash abu hadwa saghir, Khaffash. Armenian: Փոքր պայտաքիթ. Azerbaijani: Kiçik nalburun. Basque: Ferra-saguzar txikia. Belarusian: Падкаванос малы. Bosnian: Mali potkovasti šišmiš. Breton: Frigribell vihan. Bulgarian: Малък подковонос. Castilian (Spain): Murciélago pequeño de herradura, Rinolofo pequeño de herradura. Catalan (Spain): Ratpenat de ferradura petit, Rat penat petit de ferradura, Ratapinyada petita de ferradura. Croatian: Mali potkovnjak, Mali potkovasti šišmiš. Czech: Vrápenec malý, wrápenec menší, vrápenec menší, malý vrápenec podkovní, vrápeník malý, vrápenec moravský. Danish: Lille hestekonæse, Dvaerghesteskonaes. Dutch: Kleine hoefijzerneus. English: Lesser Horseshoe Bat, Smaller Horseshoe Bat. Estonian: Väikesagarnina. Finnish: Euroopanpikkuherkko. French: Petit rhinolophe, Petit rhinolophe fer à cheval, Petit Fer-à-Cheval. Frisian: Lytse hoefizernoas. Galician (Spain): Morcego de ferradura pequeño, Morcego pequeño de ferradura. Georgian: მცირე ცხვირნალა. German: Kleine Hufeisennase, Kleinhufeisennase. Greek: Μικρορινόλοφος. Hebrew: ‫גמדי פרסף‬, Parsaf Gamadi. Hungarian: Kis patkósdenevér, Kis patkósorrú denevér, Rhinolophos i mikra. Irish Gaelic: Crú-ialtóg bheag. Italian: Rinolofo minore, Ferro di Cavallo Minore. Latvian: Mazais pakavdegunis. Lithuanian: Mažasis pasagnosis. Luxembourgish: Kleng Huffeisennues. Macedonian: Мал потковичар [= Mal Potkovichar]. Maltese: Rinolofu Żgħir, Farfett ilLejl tan-Nala. Montenegrin: Mali potkovičar. Norwegian: Dverghesteskonese, Liten hesteskonese. Polish: Podkowiec mały. Portuguese: Morcego-de-ferradura-pequeno. Rhaeto-Romance: Rinolof pitschen, Nas fierchaval pitschen. Romanian: Liliacul mic cu potcoavă, Liliac-mic-cupotcoavã. Russian: Подковонос малый [= Podkovonos malyj. Serbian: Мали потковичар [= Mali potkovičar]. Scottish Gaelic: Crudh-ialtag bheag. Slovak: Podkovár malý. Slovenian: Mali podkovnjak. Swedish: Dvärghästskonäsa. Turkish: Küçük Nalburunlu Yarasa. Ukrainian: Підковик (Підковоніс) малий [= Pidkovyk malyy]. Welsh: Ystlum pedol lleiaf. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: R. hipposideros is recorded from the Guenfouda Cave (syn. Ghar Zebouj) in eastern Morocco, dating from the beginning of the Holocene (Neolithic) period (López-García et al., 2012: 51). Dool et al. (2013: 4067) indicate that during the Pleistocene, R. hipposideros utilized multiple refugia across the Mediterranean. Dool et al. (2016a: 206) indicate that R. hipposideros represents the oldest lineage of the genus and diverged fromthe other species around 16 Mya. CONSERVATION STATUS: Global Justification This species has a large range. Although there have been marked and well-documented declines in some areas, the species remains widespread, fairly common, and apparently stable in other areas. Assessed as Least Concern (LC ver 3.1 (2001)) (Jacobs et al., 2008aq; IUCN, 2009; Taylor, 2016d). Assessment History Global 2016: LC ver 3.1 (2001) (Taylor, 2016d). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008aq; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004af; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Threats include disturbance and loss of underground habitats and attics (conversion of attics in human habitat), agricultural intensification, fragmentation and isolation of habitats, and the use of pesticides in agricultural areas (Jacobs et al., 2008aq; IUCN, 2009; Taylor, 2016d). In Austria, Reiter et al. (2012: 283) studied the influence of landscape fragmentation on R. hipposideros and found that conservation measures for colonies of lesser horseshoe bats should be undertaken within 2.5 km of the nursery roost. As these bats seem to be unable to adapt their spatial foraging behaviour by shifting from woodland to a degraded landscape, any loss of African Chiroptera Report 2020 woodland near the colonial roosts will negatively influence the colony. R. hipposideros's aerial hawking and its longdistance disperal are considered to be major risk factors in view of climatic change (Sherwin et al., 2012: 174). CONSERVATION ACTIONS: According to Jacobs et al. (2008aq) [in IUCN (2009)] and supported by Taylor (2016d), Rhinolophus hipposideros is protected by national legislation in all European range states. There are international legal obligations for protection through Bonn Convention (Eurobats) and Bern Convention, where those apply. Included in Annex II (and IV) of EU Habitats and Species Directive and hence requiring special measures for conservation including designation of Special Areas for Conservation. Some habitat protection through Natura 2000. Recommended conservation measures include protecting maternity roosting sites, hibernation caves and foraging habitats. No specific conservation measures apply in South Asia; more research and monitoring are needed. GENERAL DISTRIBUTION: Rhinolophus hipposideros is widely distributed in the western and central Palaearctic. It is found in all the European countries (including the islands) of the Mediterranean region. In North Africa it is recorded from Morocco, Algeria, Tunisia and the eastern part of the Sinai (to Egypt); also occurs in eastern Africa. Also recorded from Anatolia and the countries of the Levant. It occurs from sea level to 2,000 m. Native: Afghanistan; Albania; Algeria; Andorra; Armenia; Austria; Azerbaijan; Belgium; Bosnia and Herzegovina; Bulgaria; China; Croatia; Cyprus (Benda et al., 2007: 71); Czech Republic; Djibouti (Pearch et al., 2001: 388); Egypt (Sinai); Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 147); France [Corse]; Georgia; Germany; Gibraltar; Greece [Kriti]; Holy See [Vatican City State]; Hungary; India; Iran; Ireland; Islamic Republic of; Iraq; Ireland; Israel; Italy [Sardegna, Sicilia]; Jordan; Kazakhstan; Kyrgyzstan; Lebanon; Luxembourg; Macedonia, the former Yugoslav Republic of; Malta; Moldova; Monaco; Montenegro; Morocco (Benda et al., 2010a: 157, El Ibrahimi and Rguibi Idrissi, 2015: 359); Pakistan; Palestinian Territory, Occupied (Shehab et al., 2006: 161); Poland; Portugal; Romania; Russian Federation; San Marino; Saudi Arabia; Serbia; Slovakia; Slovenia; Spain [Baleares]; Sudan; Switzerland; Syrian Arab Republic; Tajikistan; Tunisia (Dalhoumi et al., 2014: 53; 2016b: 867; 2019b: 26); 323 Turkey; Turkmenistan; Ukraine; United Kingdom; Uzbekistan. Regionally extinct: Liechtenstein; Netherlands. DETAILED MORPHOLOGY: Baculum - See Csorba et al. (2003: 54). ECHOLOCATION: Jones and Siemers (2010: 449 - 450) indicated that females emit pulses with a higher frequency than males, and also that juveniles emit lower frequencies than adults. Papadatou et al. (2008b: 132) report the following data for 5 Greek specimens: Fstart: 96.6 ± 10.25 kHz, Fend: 84.8 ± 4.74 kHz, Fpeak: 110.6 ± 3.93 kHz, bandwidth: 4.8 ± 1.61 kHz, duration 45.2 ± 6.43 msec and interpulse interval: 98.2 ± 29.10 msec. For 52 calls of bats from France, Switzerland and the UK, Walters et al. (2012: suppl.) report the following figures: duration: 42.47 ± 12.66 msec, Fmax: 109.76 ± 2.35 kHz, Fmin: 94.35 ± 3.95 kHz, bandwidth: 15.41 ± 4.26 kHz, Fpeak: 109.58 ± 2.36 kHz. The following data for 18 Iranian calls were reported by Benda et al. (2012a: 182): Fstart: 111.2 ± 0.8 (109.9 - 112.2) kHz, Fend: 108.5 ± 1.4 (106.2 - 110.7) kHz, Fpeak: 110.3 ± 0.8 (109.0 - 111.1) kHz, duration: 49.9 ± 1.5 (47.8 - 52.0) msec, interpulse interval: 41.9 ± 3.8 (36.1 - 48.7) msec. Disca et al. (2014: 226) indicate that the type of call is FM-FC-FM in Morocco, with Fpeak: 116.9 ± 0.2 kHz and duration: 39.7 ± 13.2 msec, whereas Dalhoumi et al. (2016b: 866) report a frequency "> 112 kHz" (111.5 - 117.5 kHz). 57 calls were recorded by Hackett et al. (2016: 223) from Israel: Pulse duration: 42.28 ± 12.09 msec, Fstart: 92.54 ± 6.79 (83.9 - 109.3) kHz, Fend: 93.37 ± 8.61 (80 - 114.2) kHz, Fpeak: 107.58 ± 0.49 (103.5 - 109.3) kHz. Luo et al. (2019a: Supp.) reported the following data (for two calls): Fpeak: 111.1, 109 kHz, Fstart: 99, 98.2 kHz, Fend: 96.6, 96.1 kHz, and duration: 43.6, 45.7 msec. Dool et al. (2016b: Suppl.) showed that there is a sexual dimorphism in the peak frequencies in Irish bats, with males call on average 4 kHz lower. MOLECULAR BIOLOGY: DNA - See Puerma et al. (2007). Karyotype - Capanna et al. (1967) reported 2n = 56, FN = 60. However, Rushton (1970: 463) reported 2n = 58, FN = 60, a metacentric X, and a minute Y. For a female from Jordan, Qumsiyeh and Baker (1985) reported 2n = 58, FN = 60, BA = 324 ISSN 1990-6471 4. More recently, Puerma et al. (2007) reported 2n = 54, FN = 62, BA = 10, a submetacentric X and an acrocentric Y for specimens from Spain. Volleth et al. (2013: 55) and Arslan and Zima (2014: 8) indicate that three diploid chromosome numbers are present in Europe: 54, 56, and 58. The specimens with 58 chromosomes occur in Asia Minor and the Middle East, those with 56 chromosomes in the Czech Republic, Slovakia, Italy and Greece, and those with 54 chromosomes in Spain, Germany and (possibly) Switzerland. Protein / allozyme - Unknown. DIET: Andreas et al. (2013) using faecal analysis of R. hipposideros from southern Slovakia, showed that this species fed only on the small moths (<25mm) (%f = 59; %vol = 87) and this prey category was supplemented with Neuroptera (Hemerobiidae) (%f = 13.8; %vol = 1.5), nematoceran (%f = 20.7; %vol = 10.0) and brachyceran (%f = 6.9; %vol = 1.2) Diptera. In northern Spain, Aldasoro et al. (2019: 9) found the following prey groups (in descending order of occurrence): Diptera (100 %), Lepidoptera (71 %), Neuroptera (48 %), Hemiptera and Trichoptera (both 12 %), Coleoptera (10 %), Aranea and Hymenoptera (6 %) and Psocoptera (3 %). Of the 55 prey species, identified by Baroja et al. (2019: 5-6), 25 cause significant agricultural damage and eight are pests for grapevines. Droppings from northern Algeria were examined by Ahmim and Moali (2013: 175), who found the following prey items: Insecta (93.46 %) and Chilopoda (6.54 %). Among the Insecta, Diptera were mostly encountered (41.58 %), more specifically: Culicidae (15.59%), Chironomidae/Ceratopogonidae (9.68 %) and Tipulidae (6.45 %). Lepidoptera formed the second most important insect order (21.1 %), followed by Hemiptera (11.68 %). POPULATION: Structure and Density:- According to Taylor, 2016d) and Jacobs et al. (2008aq) [in IUCN (2009)], Rhinolophus hipposideros) is an infrequent species in the northern part of its range. In Europe the species forms summer colonies of 10 - 50 individuals (up to 1,500 animals), whereas it is solitary in winter or forms loose aggregations up to 500 animals per roost. Since the 1950s the northern border of the range in western and central Europe has moved to the south. In the Netherlands, northern Belgium and Germany (except for a few colonies in Bavaria, Thüringen, Sachsen and Sachsen-Anhalt) the species went extinct (Fairon et al., 1982; Schofield, 1999). It disappeared from northern and western parts of Bohemia, and much of Poland where 87 % of the hibernating population was lost between 1950 and 1990 (Urbanczyk, 1994; Ohlendorf, 1997). In Switzerland and Austria the distribution became fragmented, as colonies remained only at higher elevations (>400 m) (Stutz and Haffner, 1984; Spitzenberger, 2002), although in Switzerland at least the population has started to slowly recover over the last 10 years (increasing from 2,200 to 2,500 adults counted in maternity roosts: H. Kraettli pers. comm., 2006). In Spain some colonies have disappeared due to the restoration of buildings, but there are no data on population trend (J. Juste and T. Alcalde pers. comm., 2006), and in France there have been some declines in the north, although large populations in the south are thought to be more stable (EMA Workshop, 2006). In the southwest Asian part of the range it gathers in wintering colonies of up to 40 animals, although it is mainly solitary (K. Tsytsulina pers. comm., 2005). In Turkey it is a commonly reported species, and the population is stable (A. Karatash pers. comm., 2005). It is common in Iran although encountered less frequently than R. ferrumequinum (M. Sharifi pers. comm., 2005). It is not known how abundant this species is in Jordan and Syria but it may be more common than the collection reports indicate (Amr, 2000). Size and trends within Africa and South Asia are unknown. Trend:- 2016: Decreasing (Taylor, 2016d). 2008: Decreasing (Jacobs et al., 2008aq; IUCN, 2009). LIFESPAN: Lagunas-Rangel (2019: 2) reports a longevity of 29.4 years. REPRODUCTION AND ONTOGENY: Tuttle and Stevenson (1982: 120) indicate that some females might attain sexual maturity in as little as three months, whereas other females and males might need 15 months. At birth, the young has about 34.4 % of the mother's body mass (4.2 g versus 12.3 g) (see Kurta and Kunz, 1987: 82). POSTNATAL DEVELOPMENT: In Austria, Reiter (2004: 234) found that the average forearm length at one day of age was 17.5 mm and the average weight 2.0 g. He also found that the growth of both the forearm and weight was almost linear during the first 14 days. In some cases, a loss of weight could be observed between day 15 and 20, which occurred a few days before the first flights took place (at between 18 and 20 days of age). At the time of their first flight, the African Chiroptera Report 2020 325 forearm was 92 % of that the adult female, and weight was 70 % of the adult's. This virus was mentioned by Nieto-Rabiela et al. (2019: Suppl.). PARASITES: Lainson and Naiff (2000: 128) repored R. hipposideros as host for Eimeria hessei Lavier, 1924 (Apicomplexa). Paramyxoviridae Nieto-Rabiela et al. (2019: Suppl.) mention presence of Bat paramyxovirus. In Italy, Voyron et al. (2011: 193) reported the following fungal entitiy from R. hipposideros carcasses: Mortierella gamsii Miko, 1974. Genov et al. (1992) reported the presence of the Nematode Molinostrongylus alatus (Ortlepp, 1932) in Moldova. Benda et al. (2010b: 234) found the bat fly Phthiridium integrum (Theodor and Moscona, 1954) (Nycteribiidae) on R. hipposideros in Jordan. They also indicate that Phthiridium bilobum (Theodor and Moscona, 1954) was reported before from Palestine, and the fleas Ischnopsyllus octactenus (Kolenati, 1856) and Rhinolophopsylla u. unipectinata (Taschenberg, 1880) from Turkey. From Algeria, Bendjeddou et al. (2013: 325) reported infestations by Phthiridium biarticulatum Hermann, 1804 (Diptera: Nycteribiidae) and Cimex lectularius (Linnaeus, 1758) (Hemiptera: Cimicidae). Another Algerian ectoparasite was reported by Bendjeddou et al. (2017: 15): Arachnida: Ixodes ricinus (Linnaeus, 1758). Picornaviridae Nieto-Rabiela et al. (2019: Suppl.) mention presence of Picornavirus. Reoviridae Orthoreovirus Reported from Italy by Kohl and Kurth (2014: 3113). Rotavirus Rotavirus A was mentioned by Nieto-Rabiela et al. (2019: Suppl.). Orthomyxoviridae Despite sampling and testing of 6 individuals of this species, no evidence of influenza A-like viruses (Orthomyxoviridae) were obtained (Fereidouni et al., 2015). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Angola, Cameroon, Egypt, Eritrea, Ethiopia, Liberia, Morocco, Sudan, Tunisia. VIRUSES: Circoviridae Bat Circovirus was mentioned by Nieto-Rabiela et al. (2019: Suppl.). Coronaviridae Alphacoronavirus This virus was reported from Italy by Kohl and Kurth (2014: 3112). Betacoronavirus Kohl and Kurth (2014: 3112) reported this virus from Bulgaria, Germany, Slovenia, Italy. Herpesviridae Rosoleovirus Figure 97. Distribution of Rhinolophus hipposideros Rhinolophus horaceki Benda and Vallo, 2012 *2012. Rhinolophus horaceki Benda and Vallo [Goto Description]. Holotype: NMP 49880: ad ♂, skin and skull. Collected by: Michal Andreas, Petr Benda, Vladimir Hanák, Antonin Reiter and Marcel Uhrin; collection date: 15 May 2002; original number: pb2124. Paratype: NMP 49861: ad ♂, skull and alcoholic. Collected by: Michal Andreas, Petr Benda, Vladimir Hanák, Antonin Reiter and Marcel Uhrin; collection date: 12 May 2002; original number: pb2104. Paratype: NMP 49879: ad ♂, skull and alcoholic. Collected by: Michal Andreas, Petr Benda, Vladimir Hanák, Antonin Reiter and Marcel Uhrin; 326 ISSN 1990-6471 collection date: 15 May 2002; original number: pb2123. Paratype: NMP 49882: ad ♀, skull and alcoholic. Collected by: Michal Andreas, Petr Benda, Vladimir Hanák, Antonin Reiter and Marcel Uhrin; collection date: 16 May 2002; original number: pb2127. Paratype: NMP 49915: ad ♀, skull and alcoholic. Collected by: Michal Andreas, Petr Benda, Vladimir Hanák, Antonin Reiter and Marcel Uhrin; collection date: 20 May 2002; original number: pb2163. (Current Combination) TAXONOMY: Based on cytochrome b sequences, Benda and Vallo (2012) were able to distinguish a new species of Rhinolophus from northern Libya (Cyrenaica). The cytochrome b shares unique base positions with the ferrumequinum/clivosus complex, but also with the fumigatus group. Benda et al. (2014c: 24) indicate that specimens of this species were reported as R. clivosus by Qumsiyeh and Schlitter (1982). COMMON NAMES: Dutch: Horacekcs hoefijzerneus. Lybische Hufeisennase. German: GENERAL DISTRIBUTION: Benda et al. (2014c: 22) indicate that R. horaceki is an endemic of the Mediterranean part of Cyrenaica, streching about 360 km along the coast, and covering no more than 4,000 km 2. ROOST: Benda et al. (2014c: 22) found a group of four bats in a cave situated in the northern slope of the side valley of the western part of Wadi Al Kuf (Libya). DIET: The diet of this species was determined by Benda et al. (2014c: 26). One group of faecal pellets contained 99.8 % by volume medium-sized moths and 0.2 % small nematoceran Diptera. A second group of pellets contained 54 % of medium-size Lepidoptera, 38 % of Blattoidea, 4 % of Formicoidea, 2 % of Auchenorrhyncha and 2 % of Coleoptera. REPRODUCTION AND ONTOGENY: Two pregnant females were collected by Benda et al. (2014c: 24) on 16 and 20 May 2002. Rhinolophus kahuzi Fahr and Kerbis Peterhans, 2013 *2013. Rhinolophus kahuzi Fahr and Kerbis Peterhans, in: Kerbis Peterhans, Fahr, Huhndorf, Kaleme, Plumptre, Marks and Kizungu, Bonn. Zool. Bull., 62 (2): 192, 194, figs 5, 6. Publication date: November 2013. Type locality: Congo (Democratic Republic of the): Western slope of Mt. Kahuzi [02 15 09 S 28 40 09 E, 2 600 m] [Goto Description]. Holotype: FMNH 219793: ad ♂, skull and alcoholic. Collected by: R. Kizungu; collection date: 28 July 2007; original number: JCK 5406. Presented/Donated by: ?: Collector Unknown. - Etymology: The name refers to the type locality and is used as a noun in apposition (see Kerbis Peterhans et al., 2013: 194). (Current Combination) COMMON NAMES: English: Kahuzi horseshoe bat. German: KahuziHufeisennase. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the). Figure 98. Distribution of Rhinolophus kahuzi African Chiroptera Report 2020 327 Rhinolophus landeri Martin, 1838 *1838. Rhinolophus landeri Martin, Proc. zool. Soc. Lond., 1837, V (lviii): 101. Publication date: 25 May 1838. Type locality: Equatorial Guinea: Bioko [formerly Fernando Po] [ca. 03 21 N 08 40 E]. Holotype: BMNH 1855.12.26.250:. - Etymology: Named by Martin after Richard Lemon Lander (1804 - 1834), an English explorer of western Africa (see Rosevear, 1965; Taylor, 2005; Lanza et al., 2015 109). (Current Combination) 1904. Rhinolophus Dobsoni Thomas, Ann. Mag. nat. Hist., ser. 7, 14 (80): 156, footnote. Publication date: 1 August 1904. Type locality: Sudan: Kordofan [ca. 13 00 N 30 00 E] [Goto Description]. Holotype: BMNH 1847.5.7.49: ♀. 1917. Rhinolophus axillaris Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 429. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Aba [03 53 N 30 17 E] [Goto Description]. Holotype: AMNH 49175: ad ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 17 December 1911; original number: 1807. The museum labels indicate it as a female. ? Rhinolophus landeri dobsoni: (Name Combination) ? Rhinolophus landeri landeri: (Name Combination) TAXONOMY: Meester et al. (1986) state that Rosevear (1965) and Kock (1969a) regard guineensis Eisentraut, 1960, from Guinea, as subspecifically distinct from landeri. Hayman and Hill (1971), on the other hand, consider it a synonym, while Böhme and Hutterer (1979, Csorba et al. (2003: 59 - 61) and Simmons (2005: 356) regard it as a distinct species, sympatric with landeri and somewhat larger. Other names associated with landeri are dobsoni Thomas, 1904, which is either a synonym (Kock, 1969a) or a valid subspecies (Koopman, 1975). See Csorba et al. (2003: 62) for remarks on taxonomy. lobatus Peters, 1852 and angolensis Seabra, 1898 were removed by Taylor et al. (2018a: 22). COMMON NAMES: Afrikaans: Lander se saalneusvlermuis, Landersaalneusvlermuis. Castilian (Spain): Murciélago de Herradura. Chinese: 兰 德 菊 头 蝠 . Czech: vrápenec Landerův. English: Lander's Horseshoe Bat. French: Rhinolophe de Lander. German: Landers Hufeisennase, Kastanienrothe Kammnase, Rundlappigen Kammnase, Lander's Rundblattnase. Italian: Rinòlofo di Lànder. Kiluba (DRC): Kasusu. Portuguese: Morcego ferradura de Lander. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008an; IUCN, 2009; Monadjem et al., 2017cl). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cl). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008an; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004ap; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (Monadjem et al., 2016h). MAJOR THREATS: There appear to be no major threats to Rhinolophus landeri as a whole (Jacobs et al., 2008an; IUCN, 2009; Monadjem et al., 2017cl). CONSERVATION ACTIONS: Jacobs et al. (2008an) reported in IUCN (2009) and Monadjem et al. (2017cl) that in view of the wide range of Rhinolophus landeri it seems probable that it is present in some protected areas. No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Rhinolophus landeri has been widely reported from much of sub-Saharan Africa. It ranges from Senegal and The Gambia in the west, through most of West and Central Africa to Sudan and Ethiopia in the east. The species had been recorded as far south as eastern South Africa, but all southern and eastern African specimens are now referred to R. lobatus (see Taylor et al., 2018a: 24). On Mount Cameroon, they have been taken at an altitude of 1,200 m asl and at 900 328 ISSN 1990-6471 m asl on Mount Bintamane in Sierra Leone (Rosevear, 1965). Native: Benin (Capo-Chichi et al., 2004: 162); Burkina Faso (Kangoyé et al., 2015a: 610); Cameroon; Central African Republic; Congo; Côte d'Ivoire; Equatorial Guinea (including Bioko); Ethiopia; Gabon; Gambia (Emms and Barnett, 2005: 50); Ghana; Guinea (Fahr and Ebigbo, 2003: 128; Decher et al., 2016: 264); Liberia (Monadjem and Fahr, 2007: 47); Niger; Nigeria; Senegal; Sierra Leone; Somalia; Sudan; Togo. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Menzies (1973) [in Young (1975: 76)] indicates that, in some colonies in northern Nigeria, rufous bats get a grey fur during the molting season, and subsequently change back to rufous. SEXUAL DIMORPHISM: Krutzsch (2000: 117) mentions the presence of shoulder glands in the males. ECHOLOCATION: In Guinea, Fahr and Ebigbo (2003) reported the frequency of the CF-component as 103.3 kHz. In Kenya, O'Shea and Vaughan (1980) using a Holgate ultrasonic sensor reported a frequency of 55 kHz for individuals at Masalani, later Taylor et al. (2005) using a Pettersson D980 reported a maximum frequency of 110 (± 0) kHz for an individual from Bungule. In Burkina Faso, Kangoyé et al. (2015a: 613) recorded one male calling at 108.5 kHz, and a female calling at 105.6 kHz. In Sierra Leone, two males called at 102.0 (101.5 - 102.5) kHz (Weber et al., 2019: 21). Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 109 kHz and duration: 43.1 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - 2n = 58, FN = 60, BA = 4, and a submetacentric X chromosome (Rautenbach et al., 1993). Koubínová et al. (2010b) agree with Rautenbach et al. (1993), and reported for a female from Senegal 2n= 58, and a Fna= 64, which contained four pairs of biarmed autosomes three pairs of which were medium to small sized meta- or submetacentrics and one was a medium sized subtelocentric, the remaining 24 pairs were a gradated series of acrocentrics. Protein / allozyme - Unknown. HABITAT: Monadjem et al. (2016y: 366) recorded this species in the Mount Nimba area in forested habitats between 550 and 1,000 m. POPULATION: Structure and Density - This species is rather common locally, colonies can consist of hundreds of individuals (Jacobs et al., 2008an; IUCN, 2009; Monadjem et al., 2017cl). Trend:- 2016: Unknown (Monadjem et al., 2017cl). 2008:- Unknown (Jacobs et al., 2008an; IUCN, 2009). REPRODUCTION AND ONTOGENY: For Malawi, Happold and Happold (1990b: 566) reported that young are born early in the wet season. Orr and Zuk (2014: 902) refer to Racey (1982) [who in turn refers to Menzies (1973)] and indicated that in R. landeri it can take up to two months before the blastocyst gets implanted. PARASITES: HAEMOSPORIDA Adam and Landau (1973a: 5) report on the presence of a protozoan parasite of the genus Polychromophilus (Haemoproteidae) in R. landeri. About 50 % of the specimens from Congo they examined, were found to be infected. Gametocytes were found year-round. The bats were also infested by the nycteribiids Penicillidia fulvida Bigot, 1885 and Nycteribia schmidlii scotti Falcoz, 1923, of which the prior was a major carrier of Polychromophilus sporozoites. Schaer et al. (2013a: 17416) also found this parasite (possibly N. congolensis or N. gabonensis) in one out of two West African bats they examined. Polychromophilus ? murinus was reported by Garnham (1973: 237), based on a record by Adam and Landau. Espinosa-Álvarez et al. (2018: Suppl.) and Cai et al. (2019: Suppl.) reported Trypanosoma livingstonei from three bats from Mozambique. Schaer et al. (2015: 381) found Nycteria sp, in a bat from Guinea and Nycteria cf. congolensis in a R. landeri from Sierra Leone. Perkins and Schaer (2016: Suppl.) reported on Nycteria congolensis (Garnham, 1966) from Sierra Leone, and Nycteria sp. from Côte d'Ivoire or Guinea. Rosskopf et al. (2018: 31) also found Nycteria parasites in bats from Gabon. Justine (1989b: 537, 544) described two new stomach parasites: Capillaria landauae and African Chiroptera Report 2020 Capillaria brosseti (Nematoda) from R. landeri from Makokou (Gabon). Polyctenidae: Androctenes horvathi Jordan, 1912 recorded from Congo (Haeselbarth et al., 1966: 17). Streblidae: Ascodipteron lophotes Monticelli 1898 from Kenya (Haeselbarth et al., 1966: 106, but Maa (1965) suggests that these records need verification). Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Penicillidia pachymela Speiser, 1901, from Somalia, Sudan, Kenya, Tanzania, the Congo, Guinea and the Cameroon Mts (Haeselbarth et al., 1966: 114). Raymondia intermedia Jobling, 1936 in Sierra Leone (Haeselbarth et al., 1966: 102), Kenya (Haeselbarth et al., 1966: 102, host referred to as R. axillaris), see also Shapiro et al. (2016: 255). Raymondia simplex Jobling, 1955 at Kasongo, Congo (Haeselbarth et al., 1966: 104; Shapiro et al., 2016: 256). Raymondia waterstoni Jobling, 1931 (Shapiro et al., 2016: 256). 329 Ghana tested negative for Repirovirus, Henipavirus, Morbillivirus, Rubulavirus or Pneumovirus. In South Africa, however, Mortlock et al. (2015: 1841) reported that the only R. landeri specimen they examined, tested positive for Paramyxovirus sequences. Picornaviridae Hepatovirus - Zeghbib et al. (2019: Suppl) reported this virus from a Ghanaian bat. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Cameroon, Central African Republic, Côte d'Ivoire, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Kenya, Mali, Mauritania, Nigeria, Senegal, Sierra Leone, South Sudan, Sudan, The Gambia, Togo. VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Rhinolophus landeri: Astroviruses and Coronaviruses. Coronaviridae - Coronaviruses SARS-CoV - Seven out of 58 (12.1 %) Kenyan bats tested positive for CoV (Tao et al., 2017: Suppl.). Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that the single specimen they examined from Figure 99. Distribution of Rhinolophus landeri Rhinolophus lobatus Peters, 1852 1852. Rhinolophus lobatus Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 41, pl. 9; pl. 13, figs 16 - 17. Publication date: 1852. Type locality: Mozambique: S bank Zambesi River, Tette: Sena [17 28 S 35 01 E] [Goto Description]. Syntype: ZMB 24922: ♂, complete skeleton. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008). Syntype: ZMB 24927: ♀, complete skeleton. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008). Syntype: ZMB 2496: ♀, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008). Syntype: ZMB 375: ♀, skin only. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008). - Comments: Brown and Dunlop (1997: 1) mention the type locality as "Africa orientalis, Sena, Tette" (=Sena, Mozambique) restricted by Moreau et al. (1946: 399). For coordinates: see Ansell and Dowsett (1988: 33). Taylor et al. (2018a: 22) situate Sena at -15.679 S, 33.809 E [=15 40 44 S 33 48 32 E]. This is in the Tete province, but north of Tete and 57 km north of the Zambezi River, whereas they also mention the locality to be on the south bank of the Zambezi River. According to Google maps these coordinates point to Muchena, which is on the Revuboe River. We believe 330 ISSN 1990-6471 1898. ? the type locality is the same as "Vila de Sena", which is located at 17 26 30 S 35 01 38 E in the Sofala province, at the south bank of the Zambezi River. Rhinolophus angolensis Seabra, J. Sci. mat. phys. nat., ser. 2, 5: 250. Type locality: Angola: Hanha [12 15 S 13 45 E]. Rhinolophus landeri lobatus: (Name Combination) TAXONOMY: Used to be included in R. landeri Martin, 1838, but Taylor et al. (2018a: 22) found molecular differences which led them to recognize the taxon as a separate species. They also transferred angolensis Seabra, 1898, although this taxon might turn out to be a separate species. Taylor et al. (2019b: 535) indicated that the above analyses was based on a contaminated cyt-b sequence of R. landeri, but a new analyses with a correct sequence led to an identical result. CONSERVATION STATUS: Assessment History Regional South Africa (as R. landeri):- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016h). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). GENERAL DISTRIBUTION: For southern African "R. landeri", CooperBohannon et al. (2016: Table S2) calculated a potential distribution area of 1,008,924 km 2. Taylor et al. (2018a: 25) suggest that all savannah populations in southern and eastern Africa might belong to R. lobatus. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 560); Burundi; Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259Monadjem et al., 2010d: 560); Kenya; Malawi (Happold et al., 1988; Monadjem et al., 2010d: 560); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 560; Monadjem et al., 2010c: 379); Namibia; Rwanda; South Africa (Monadjem et al., 2010d: 560); Tanzania (including Unguja [=Zanzibar] [O'Brien, 2011: 286]); Uganda; Zambia (Ansell, 1969; Ansell, 1978; Monadjem et al., 2010d: 560); Zimbabwe (Cotterill, 2004a: 261; Monadjem et al., 2010d: 560) [All as R. landeri]. However, as most of the underlying specimens have not been re-examined, some records from the northeastern DRC and northern Uganda might actually represent R. landeri. ECHOLOCATION: In South Africa, Aldridge and Rautenbach (1987) reported (as R. landeri) a maximum frequency of 110 kHz for individuals from Pafuri, while Jacobs et al. (2007a) reported a peak frequency of 107.3 (± 2.1) kHz (detector type and locality not reported). Monadjem et al. (2010c: 379) reported from Mozambique that the peak echolocation frequencies for a single female was102.2 kHz (ANABAT), while a single male was recorded at 104 kHz (Pettersson S240x). Four specimens from Chihalatan and Malashane (Mozambique) had peak frequency of 106.8 ± 0.4 kHz (Taylor et al., 2018a: 24). PARASITES: HAEMOSPORIDA Texeira and Camargo [in Lima et al. (2013: 13)] described Trypanosoma livingstonei from a "R. landeri" specimen collected in Mozambique (see also Barbosa et al., 2016: 215). DIPTERA Streblidae: Raymondia aspera Maa, 1968 was reported from "R. landeri" in Mozambique (Shapiro et al., 2016: 254). VIRUSES: Astroviridae None of the nine Mozabican bats tested by Hoarau et al. (2018: 2) was positive for Astroviridae. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera of two "R. landeri" specimens from then Limpopo Province, South Africa, neither was positive (0/2). Joffrin et al. (2020: 4) reported on Alphacoronavirus in a bat from Mozambique. an DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Burundi, Congo (Democratic Republic of the), Kenya, Malawi, Mozambique, Rwanda, South Africa, Tanzania, Uganda, Zambia, Zimbabwe. African Chiroptera Report 2020 331 Figure 100. Distribution of Rhinolophus lobatus Rhinolophus mabuensis Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 *2012. Rhinolophus mabuensis Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, PLoS ONE, 7(9): 17. Publication date: 12 September 2012. Type locality: Mozambique: Mt. Mabu [161702S 362353E, 1043 m] [Goto Description]. Holotype: DNSM 10842: ad ♀, skull and alcoholic. Collected by: Michael Curran and Mirjam Kopp; collection date: 13 October 2008. Presented/Donated by: ?: Collector Unknown. Baculum prepared Taylor et al. (2012c: 17). Paratype: DNSM 11485: ad ♀, skull and alcoholic. Collected by: J. Bayliss; collection date: 5 September 2009. Presented/Donated by: ?: Collector Unknown. - Etymology: Taylor et al. (2012c: 17) selected the specific epithet to draw attention to the serious threats to the unique biodiversity isolated on the montane forest islands in northern Mozambique – notably Mts Mabu and Inago. None of these landforms lie within formally protected areas, and all are undergoing major habitat degradation and destruction from ever-increasing human activities - hunting, fires, timber harvesting and expanding agriculture. - ZooBank: C205928E-D72C-40D5-9647-FDC9391CB1A6. (Current Combination) TAXONOMY: Based on mitochondrial and nuclear DNA sequences and morphological and morphometric data, Taylor et al. (2012c) found that the Rhinolophus hildebrandtii group includes more species than previously recognized. ZOOBANK: C205928E-D72C-40D5-9647-FDC9391CB1A6 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Mozambique. COMMON NAMES: English: Mount Mabu Horseshoe Bat. German: Mt Mabu Hufeisennase. GENERAL DISTRIBUTION: Known only from two mountains in northern Mozambique but quite possibly extending to nearby Mts. Namuli, Chiperone, Mulanje and the Malawi Rift (Taylor et al., 2012c: 17). Accurate delimitation of this species’ range is subject to further collecting and reappraisal of existing museum material, previously refered to as R. hildebrandtii. Native: Mozambique (Taylor et al., 2012c: 17). Figure 101. Distribution of Rhinolophus mabuensis 332 ISSN 1990-6471 Rhinolophus maclaudi Pousargues, 1898 *1898. Rhinolophus Maclaudi Pousargues, Bull. Mus. natn. Hist. nat., Paris, sér 1, 3 (8): 358, f. 1, 2 (for 1897). Publication date: 22 January 1898. Type locality: Guinea: Conakry Island [09 31 N 13 43 W] [Goto Description]. Holotype: MNHN ZM-MO-1897-281: ♀, skull and alcoholic. Collected by: Dr. C. MacLaud. Number 177 in Rode (1941: 237). - Comments: Grubb et al. (1998: 78) mention 1898 as year of publication. - Etymology: Named in honour of dr. Maclaud, collector of the holotype (Pousargues, 1897: 361). (Current Combination) ? Rhinolophus maclaudi: (Current Spelling) TAXONOMY: R. maclaudi is an ecomorphological outlier having long-ears, which traditionally placed it in the R. philippinensis group (Andersen, 1905c). Csorba et al. (2003: xvii, 63 - 64) state that it is located well inside the Africa clade, in the maclaudi-group, and are followed by Simmons (2005: 358). Koopman (1993a: 167) included ruwenzorii and hilli, but this is revised by Fahr et al. (2002), who recognizes them as distinct species (followed by Csorba et al., 2003 and Simmons, 2005). COMMON NAMES: Czech: vrápenec Maclaudův. English: MacLaud's Horseshoe Bat. French: Rhinolophe de MacLaud. German: Maclauds Hufeisennase. CONSERVATION STATUS: Global Justification Listed as Endangered (EN B2ab(iii) ver 3.1 (2001)) because its area of occupancy (cave roosts) is probably less than 500 km², its distribution is severely fragmented through habitat degradation, and the extent of its habitat in the Fouta Djallon Highlands is declining (Fahr, 2008l; IUCN, 2009). Assessment History Global 2008: EN B2ab(iii) ver 3.1 (2001) (Fahr, 2008l; IUCN, 2009). 2004: EN B1ab(iii) ver 3.1 (2001) (Fahr, 2004e); IUCN, 2004. 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The relatively limited range of Rhinolophus maclaudi is quite densely populated, and it is threatened by habitat loss and degradation of remaining patches of forest, and by overharvesting for the bush meat trade (Fahr et al., 2002). Sagot and Chaverri (2015: 1670) mention habitat degradation or loss, and hunting as major threats for this species. CONSERVATION ACTIONS: Fahr (2008l) [in IUCN (2009)] reported that there appear to be no direct conservation measures in place. It is not known if Rhinolophus maclaudi is present in any protected areas. Protection of habitat, further research, and awareness education urgently required. GENERAL DISTRIBUTION: Rhinolophus maclaudi is known from a very few localities in eastern Guinea. Fahr et al. (2002) report that "apart from the type locality, supposedly Conakry Island, all records are located along the lower, southern slope of the Fouta Djallon region between Kindia and Mamou near the border with Sierra Leone". However, recent surveys have revealed additional localities in the Fouta Djallon Highlands, ranging as far north as the Gessorewoul River in Prefecture Mali (Weber and Fahr, 2007). Fahr (2007a: 104) re-identified specimens reported from Liberia by Simmons (2005: 358) as R. ziama. Specimens from Nigeria reported by Koopman et al. (1995: 6, 19), Happold (1987: 60 - 61), and Csorba et al. (2003: 64) were reidentified Fahr et al. (2002: 99) as R. hildebrandtii. Native: Guinea. Presence uncertain: Liberia (Simmons (2005: 358); Nigeria (Koopman et al., 1995: 6, 19; Happold, 1987: 60 - 61; Csorba et al., 2003: 64). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Csorba et al. (2003: 63) indicate that R. maclaudi is a very large species wiith enormous ears (40.0 44.0 mm) and a broad horseshoe, covering almost the whole muzzle. There is also a distinct ridge running on its surface nearly parallel to the outer ridge. There is no secondary horseshoe. The sella is very broad with a broadly rounded summit and parallel margined above. The lancet is long, narrow and cuneate. ROOST: For Guinea, Weber and Fahr (2006: 4) indicate that R. maclaudi largely or exclusively depends on the availability of caves as day roosts. African Chiroptera Report 2020 333 POPULATION: Structure and Density:- Little information is available on the population abundance or size of this species. The species roosts singly or in small groups (Fahr et al., 2002). Trend:- 2008: Decreasing (Fahr, 2008l; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Guinea. Figure 102. Distribution of Rhinolophus maclaudi Rhinolophus maendeleo Kock, Csorba and Howell, 2000 *2000. Rhinolophus maendeleo Kock, Csorba and Howell, Senckenb. biol., 80 (1/2): 233, 234, figs 1, 4, 7. Publication date: 22 December 2000. Type locality: Tanzania: Tanga district: 2.5 km W of Tanga, Mkulumuzi River Gorge: Amboni Cave Forest [05 05 S 39 02 E, 0 80 m] [Goto Description]. Holotype: SMF 79643: ad ♂, skull and alcoholic. Collected by: L. Stubblefield; collection date: 25 February 1992; original number: KMH 7673. Paratype: SMF 66960: ad ♀. Collected by: H. Grossmann; collection date: 29 August 1985; original number: KMH 3216. Presented/Donated by: ?: Collector Unknown. Etymology: From the Swaihili "maendeleo" for progress; a noun in apposition. Named in allusion to the increasing knowledge of the Tanzanian bat fauna (see Kock et al. (2000). (Current Combination) 2010. Rhinolophus cf. maendeleo: Monadjem, Schoeman, Reside, Pio, Stoffberg, Bayliss, Cotterill, Curran, Kopp and Taylor, Acta Chiropt., 12 (2): 379. TAXONOMY: adami species group (Csorba et al., 2003: 5 - 7; Simmons, 2005: 359). See Csorba et al. (2003: 6) for remarks on taxonomy. COMMON NAMES: Czech: vrápenec tanganjický. English: Maendeleo Horseshoe Bat. French: Rhinolophe de Tanga. German: Tanzania-Hufeisennase. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since it has only recently been discovered, and there is still very little information on its extent of occurrence, status and ecological requirements (Jacobs et al., 2008ae; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Jacobs et al., 2008ae; IUCN, 2009). 2004: DD ver 3.1 (2001) (Jacobs et al., 2004r; IUCN, 2004). Regional None known. MAJOR THREATS: The threats to Rhinolophus maendeleo are not known (Jacobs et al., 2008ae; IUCN, 2009). CONSERVATION ACTIONS: Jacobs et al. (2008ae) [in IUCN (2009)] reported that while Rhinolophus maendeleo is present in the Mazumbai Forest Reserve, further habitat protection is generally needed. Additional field surveys are needed to better determine the true extent of the species geographic range. GENERAL DISTRIBUTION: Rhinolophus maendeleo is a poorly known species that has only been recorded from two localities in Tanzania at elevations of up to 1,900 m asl, the type locality of 'Amboni Cave Forest, Mkulumuzi River Gorge, 2.5 km W of Tanga, Tanga District' (Anciaux de Faveaux, 1958) and Mazumbai Forest Reserve in the Usambara Mountains. It has recently been recorded from a single locality in 334 ISSN 1990-6471 Mozambique. Additional surveys are needed to better determine the range of this recently described species. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Mozambique, Tanzania. Native: Mozambique (Monadjem et al., 2010d: 560; Monadjem et al., 2010c: 379); Tanzania (Kock et al., 2000; Csorba et al., 2003: 6). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: Csorba et al. (2003: 6) indicate that the skull of R. maendeleo has an open foramen infraorbitale (without bony bar). DETAILED MORPHOLOGY: Baculum - See Csorba et al. (2003: 6). POPULATION: Structure and Density:- Rhinolophus maendeleo is currently known only from two specimens, and there is no information on population sizes or trends (Jacobs et al., 2008ae; IUCN, 2009). Figure 103. Distribution of Rhinolophus maendeleo Trend:- 2008: Unknown (Jacobs et al., 2008ae; IUCN, 2009). Rhinolophus mehelyi Matschie, 1901 *1901. Rhinolophus mehelyi Matschie, Sber. Ges. naturf. Freunde Berlin, 225. Type locality: Romania: Ilfov province: Bucharest [44 26 N 26 06 E]. - Etymology: In honour of Lajos Méhely (1862 - 1952) (see Kozhurina, 2002: 16). (Current Combination) 1904. E[uryalus] barbarus K. Andersen and Matschie, Sber. Ges. naturf. Freunde Berlin, 79. Type locality: Morocco: Tangiers [35 48 N 05 50 W]. Holotype: ZMB 12967: ♂, skull and alcoholic. Collected by: Kurt Floericke; collection date: 12 March 1898. See Turni and Kock (2008). Paratype: MHNG ♂, skull and alcoholic. Former ZMB 12968, exchanged in 1909; see Turni and Kock (2008: 28). Paratype: MHNG ♂, skull and alcoholic. Former ZMB 12972, exchanged in 1909; see Turni and Kock (2008: 28). Paratype: ZMB 12969: ♂, skull and alcoholic. Collected by: Kurt Floericke. See Turni and Kock (2008: 28). Paratype: ZMB 12970: ♂, skull and alcoholic. Collected by: Kurt Floericke. See Turni and Kock (2008: 28). Paratype: ZMB 12971: ♂, skull and alcoholic. Collected by: Kurt Floericke. See Turni and Kock (2008: 28). Paratype: ZMB 12972: Collected by: Kurt Floericke; collection date: 12 March 1898. Paratype: ZMB 12973: alcoholic (skull missing). Collected by: Dr. Lühe; collection date: 12 March 1898. From cave near Tebourba; see Turni and Kock (2008: 28). Paratype: ZMB 12974: ♂, alcoholic, skull and skeleton. Collected by: Dr. Lühe; collection date: 12 March 1898. From cave near Terbourba, see Turni and Kock (2008: 28) [lost]. Paratype: ZMB 12975: alcoholic (skull missing). Collected by: Dr. Lühe; collection date: 12 March 1898. From cave near Tebourba; see Turni and Kock (2008: 28). 1904. E[uryalus] meridionalis K. Andersen and Matschie, Sber. Ges. naturf. Freunde Berlin, 79. Type locality: Algeria: "Algeria". Holotype: ZMB 12976: ♀, skull and alcoholic. Collected by: Verreaux; collection date: undated. See Turni and Kock (2008: 26). Paratype: ZMB 3052: ♀. Collected by: Verreaux; collection date: undated. Number of paratype is 3052, not 12977 as mentioned by Andersen and Matschie (1904: 83): see Turni and Kock (2008: 29). 1955. Rhinolophus euryale tuneti Deleuil and Labbé, Bull. Soc. Sci. nat. Tunisie, 8: 52, 53. Type locality: Tunisia: Cap Bon: El Haouaria cave [37 03 N 11 01 E] [Goto Description]. Neotype: USNM 498999: ad ♂, skin and skull. Collected by: Tom Vaughan and Pamela Vaughan; collection date: 4 May 1975; original number: TPV 3244. Presented/Donated by: ?: Collector Unknown. Fisher and Ludwig (2015: 54) mention that a type specimen was not designated by Deleuil and Labbe (1955a), and that the type series has been lost. African Chiroptera Report 2020 1993. 1995. ? ? ? 335 They also indicate that Cockrum (1976: 868) incorrectly gave the wrong number when assigning the neotype (USNM 498998). - Comments: Horácek et al. (2000: 102) consider it a valid subspecies. Dalhoumi et al. (2011: 267) indicate that Deleuil and Labbe (1955a) based their description on a specimen belonging to R. mehelyi (type) and two belonging to R. blasii (paratypes), as was already suggested by Aellen and Strinati (1970: 231) and Cockrum (1976: 685). tunetae: Koopman, in: Wilson and Reeder, Mammal species of the World (2nd Edition): Chiroptera, 167. (Lapsus) tunetai: Pavlinov, Borissenko, Kruskop and Jahonton, Arch. Zool. Mus., Moscow State Univ., 133: 81. (Lapsus) Rhinolophus euryale barbarus: (Name Combination) Rhinolophus mehelyi mehelyi: (Name Combination) Rhinolophus mehelyi tuneti: (Name Combination) TAXONOMY: Revised by DeBlase (1972). euryale species group (Csorba et al., 2003: 17 - 18; Simmons, 2005: 359). See Csorba et al. (2003: 17 - 18) for remarks on taxonomy. Based on mitochondrial data, Najafi et al. (2019: 103, 106) estimated that R. mehelyi separated from R. euryale some 4.14 (3.17 - 5.09) mya. They also suggest that the African and southernEuropean haplotypes if R. mehelyi belong to one taxon, which is different from the Iranian haplotypes, and seem to support the division in subspecies (mehelyi and tuneti). However, further analyses of nuclear DNA is required to confirm this. Both groups became isolated some 1.18 (0.68 - 1.61) mya. Frisian: Mehely's hoefizernoas. Galician (Spain): Morcego de ferradura mediano. Georgian: მეჰელის ცხვირნალა. German: MehelyHufeisennase. Greek: Ρινόλοφος του Mehely. Hebrew: ‫חיוור פרסף‬, Parsaf Hiver. Hungarian: Méhely-patkósdenevér, Rhinolophos i mehelios. Irish Gaelic: Crú-ialtóg Mehely. Italian: Rinolofo di Méhely, Ferro di cavallo di Méhely. Latvian: Meheļa pakavdegunis. Lithuanian: Mehelio pasagnosis. Luxembourgish: MéhelÿHuffeisennues. Macedonian: Мехелов потковичар. Maltese: Rinolofu ta' Mehely. Montenegrin: Mehelijev potkovičar. Norwegian: Middelhavshesteskonese. Polish: Podkowiec średni. Portuguese: Morcego-de-ferraduramourisco. Rhaeto-Romance: Rinolof da Méhely. Romanian: Liliacul cu potcoavă a lui Mehely, Liliac-românesc. Russian: Подковонос Мегели (очковый). Serbian: Тамнооки потковичар [= Tamnooki potkovičar]. Scottish Gaelic: Crudhialtag Mehelyi. Slovak: Podkovár Mehélyho. Slovenian: Mehelyjev podkovnjak. Swedish: Kusthästskonäsa. Turkish: Mehely Nalburunlu Yarasası. Ukrainian: Підковик (Підковоніс) Мегеля. Welsh: Ystlum pedol Mehely. COMMON NAMES: Albanian: Lakuriq nate hundëpatkua i Mehely-it. Arabian: Khaffash. Armenian: Մեհելիի պայտաքիթ. Azerbaijani: Eynəkli nalburun. Basque: Mehelyi ferra-saguzar. Belarusian: Падкаванос Мегеля. Bosnian: Meheljev potkovasti šišmiš. Breton: Frigribell Méhely. Bulgarian: Подковонос на Мехели. Castilian (Spain): Murciélago mediano de herradura, Rinolofo mediano de herradura. Catalan (Spain): Ratpenat de ferradura mitjà, Rat penat mitjá de ferradura. Croatian: Meheljev potkovnjak. Czech: Vrápenec Mehelyův. Danish: Mehelys hestekonæse. Dutch: Mehely's hoefijzerneus, Mehely-hoefijzerneus. English: Mehely's Horseshoe Bat, Mediterranean Horseshoe Bat. Estonian: Meheli sagarnina. Finnish: Täpläherkko. French: Rhinolophe de Mehély, Rhinolophe des forêts, Rhinolophe de Roumanie. CONSERVATION STATUS: Global Justification The species is declining throughout its range, which is becoming increasingly fragmented. The rate of decline has been estimated as 10 % in the last 10 years in Andalucia (Spain). Elsewhere, declines have not been quantified but appear to have been considerable, with some colonies having been reduced to a fraction of their former size. Overall, declines throughout the global range are thought likely to exceed 30 % over 3 generations (27 years) over a time window including both the past and the future. The species is going extinct in France and is declining in Morocco due to disturbance in caves. The species only roosts in caves and does not use artificial roosts. Hence the species is listed as Vulnerable (VU A4c ver 3.1 (2001)) (Hutson et al., 2008n; IUCN, 2009; Alcaldé et al., 2016). Based on the measurements given by Andersen and Matschie (1904), Puechmaille et al. (2012: 216) suggest that R. barbarus Andersen et Matschie, 1904 and R. meridionalis Andersen et Matschie, 1904 are both synonyms of R. mehelyi Matschie, 1901 rather than of R. euryale Blasius, 1853. 336 ISSN 1990-6471 Assessment History Global 2016: VU A4c ver 3.1 (2001) (Alcaldé et al., 2016). 2008: VU A4c ver 3.1 (2001) (Hutson et al., 2008n; IUCN, 2009). 1996: VU A2c ver 2.3 (1994) (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Rhinolophus mehelyi is affected by disturbance and loss of underground habitats, changes in foraging habitats, and destruction of caves by tourism. Mortality due to collision with cars is a problem in some areas (e.g., Portugal). The reasons for the declines are not fully understood (Hutson et al., 2008n; IUCN, 2009; Alcaldé et al., 2016). According to Sherwin et al. (2012: 174), climatic change will have an inportant impact due to the following risk factors: its small range, roosting in caves or trees, its water stress and aerial hawking feeding style. Roost loss or disturbance, and habitat degradation or loss are considered the major threats for this species by Sagot and Chaverri (2015: 1670). CONSERVATION ACTIONS: Hutson et al. (2008n) [in IUCN (2009)] and Alcaldé et al. (2016) reported that Rhinolophus mehelyi is protected by national legislation in all European range states. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention where those apply. It is included in Annex II (and IV) of EU Habitats and Species Directive, and hence equires special measures for conservation including designation of Special Areas for Conservation. There is some habitat protection through Natura 2000 (some roosts are already protected by national legislation). An EU-LIFE funded project aims to ensure the long-term conservation of the large populations of cave and forest-dwelling bats, including this species, in Spain. There are no specific conservation measures in place in North Africa. Research is required on the causes of the declines across the range. GENERAL DISTRIBUTION: Rhinolophus mehelyi is largely restricted to the Mediterranean. It has a discontinuous distribution from north Africa (Morocco, Algeria, Tunisia and Egypt) and southern Europe (southern Portugal and Spain, possibly one occurrence in France, a few places in Italy and the Balkans) through Asia Minor, Anatolia, to Transcaucasia, Iran and Afghanistan (where its exact roost location is not known: Srinivasulu et al., in press). It is patchily distributed in some large and vulnerable colonies. It occurs up to 2,000 m in High and Saharan Atlas mountains, although it is typically found at lower altitudes in other parts of its range (e.g. in Spain it tends to occur below 700 m). Native: Afghanistan; Algeria (Horácek et al., 2000: 102); Armenia; Azerbaijan; Bosnia and Herzegovina; Bulgaria; Cyprus; Egypt; Georgia; Gibraltar; Greece; Iran, Islamic Republic of; Iraq; Israel; Italy (Sardegna, Sicilia); Jordan; Libyan Arab Jamahiriya; Macedonia, the former Yugoslav Republic of; Mediterrranean islands; Moldova; Montenegro; Morocco (El Ibrahimi and Rguibi Idrissi, 2015: 359); Palestinian Territory, Occupied; Portugal; Romania; Russian Federation; Serbia; Slovenia; Spain; Syrian Arab Republic; Tunisia (Dalhoumi et al., 2016b: 867; 2019b: 26); Turkey; Yugoslavia. Regionally extinct: Croatia. Presence uncertain: France. ECHOLOCATION: Jones and Siemers (2010: 450) indicate that juveniles emit lower frequencies than adults, and that the CF-frequency is positively related to the body condition (mass/forearm length). Puechmaille et al. (2014b: 4) demonstrated that the body length and weight for bats caught during the mating season showed a strong positive correlation (which was significant) with the peak frequency in both sexes: larger and heavier bats call at higher frequencies. Females clearly show a preference for higher calling males, and the latter have a larger offspring. Puechmaille et al. (2014b: 6) also found that the distribution of the echolocation frequencies was not normally distributed in adults, but left-skewed (meaning that there are more individuals calling at higher frequencies). Furthermore (p. 7), they indicate [referring to Jones and Ransome (1993)] that the peak frequency in horseshoe bats increases with about 0.45 kHz during the first year and an average reduction of 0.09 kHz per year occurs over their lifetime. For 16 Greek specimens, Papadatou et al. (2008b: 132) report a start frequency of 96.2 ± 7.06 kHz, a terminating frequency of 90.3 ± 2.16 kHz, a peak frequency of 109.5 ± 1.44 kHz, and a bandwidth of 5.1 ± 2.62 kHz. The duration of the call is 35.9 ± 12.04 msec, and the interpulse interval is 62.5 ± 23.92 msec. Walters et al. (2012: suppl.) presen the following figures for 41 calls from Bulgarian bats: duration: 27.73 ± 7.12 msec, Fmax: 108.59 ± 1.51 kHz, Fmin: African Chiroptera Report 2020 92.61 ± 4.73 kHz, bandwidth: 15.98 ± 4.44 kHz, Fpeak: 107.78 ± 2.74 kHz. Tunisian bats emitted sounds at 106.11 ± 1.31 kHz (111.5 - 117.5 kHz; Dalhoumi et al., 2016b: 866). Luo et al. (2019a: Supp.) reported the following data: Fpeak: 109.5 kHz, Fstart: 96.2 kHz, Fend: 90.3 kHz, and duration: 35.9 msec. Schuchmann et al. (2012: 161) found that both sexes were able to recognize the sex of conspecifics from their echolocation calls. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Ðulic and Soldatovic (1969) and Rushton (1970 463) reported 2n = 58, FN = 64, a submetacentric X chromosome, and an acrocentric Y chromosome. While Baker et al. (1974; 1975) reported FN = 60 and BA = 4. Volleth et al. (2002: 483) and Volleth and Eick (2012: 166) report 2n = 58 and a segment number [= autosomal segments + X] of 44. Arslan and Zima (2014: 8) indicate that different numbers of bi-armed auosomes have been reported, leading to FNa values between 60 and 64. Ðulic and Soldatovic (1969: 5) also reported the type of the autosomes as 4 metacentric, 2 smallsized and 24 acrocentric. Protein / allozyme - Unknown. HABITAT: In south-western Spain, Russo et al. (2005: 327) found that R. meheyli least preferred open habitats, but was frequently found in a seminatural oak savanna habitat. Salsamendi et al. (2012a: 122) pointed out that this was in an area where both R. mehelyi and R. euryale occurred, and they reported that R. mehelyi, in another area, always foraged in woodlands, preferably in cluttered spaces, but also in less-clutttered/moreopen ones, and was avoiding open spaces. They also indicated that the bats foraged generally less than 500 m from water bodies. The bats were using linear landscape elements (tree lines and riparian forests) to travel from their roosts to their feeding areas. Salsamendi et al. (2012a: 129) also indicated that habitat types used by males and females in south-western Spain were different: males foraged in riparian forest and broad-leaved woodlands, whereas females foraged in dehesa, scrubland and eucalypt plantations. HABITS: Kameneva (1976) [in Pervushina et al. (2012: 750)] found that among captive R. mehelyi individuals, conflicts took place during foraging 337 flights, although they were primarily friendly or neutral to one another when resting during the day. ROOST: Based on data from Sardinia, Russo et al. (2014: 9) rule out that R. euryale and R. mehelyi compete for roosting sites. DIET: In the Iberian peninsula, Salsamendi et al. (2009: 282) report that Lepidoptera form the most important prey category for this species (from 51.3 to 100.0 volume percent; 90.9 to 100.0 occurrence percentage), followed by Neuroptera, Diptera, Coleoptera, Hemiptera and undefined items. In Iran, Lepidoptera also form the dominant food item, followed by scarabaeid beetles and muscid dipterans (see Benda et al., 2012a: 247). In the southwestern Iberian peninsula, Arrizabalaga-Escudero et al. (2018: Suppl.) identified the following food items from faecal samples of R. mehelyi (food items marked with an asterisk were also found in faecal samples of R. euryale): Coleoptera (Cerambycidae: Arhopalus ferus (Mulsant, 1839)*); Diptera (Tachinidae: unidentified*; Anthomyiidae: Hylemya sp.; Muscidae: Neomyia cornicina (Fabricius, 1781); Calliphoridae: Pollenia vagabunda (Meigen, 1826)); Lepidoptera (Blastobasidae: Blastobasis phycidella (Zeller, 1839); Cosmopterigidae: Eteobalea intermediella (Riedl, 1966); Crambidae: Calamotropha paludella (Hübner, 1824)*; Ostrinia nubilalis (Hübner, 1796)*; Udea ferrugalis (Hübner, 1796); Erebidae: Catocala nymphagoga (Esper, 1787)*; Gelechiidae: Teleiopsis sp.*; Geometridae: Camptogramma bilineata (Linnaeus, 1758)*; Ennomos quercaria (Hübner, 1813)*; Pachycnemia hippocastanaria (Hübner, 1799)*; Rhoptria asperaria (Hübner, 1817)*; Lasiocampidae: Malacosoma neustria (Linnaeus, 1758); Lecithoceridae: Eurodachtha canigella (Caradja, 1920); Noctuidae: Agrotis segetum/trux*; Agrotis ipsilon (Hufnagel, 1766)*; Autographa gamma/pulchrina*; Calophasia platyptera (Esper, [1788])*; Heliothis incarnata Freyer, 1838; Mythimna albipuncta (Denis & Schiffermüller, 1775)*; Mythimna vitellina (Hübner, 1808); Mythimna loreyi/sicula*; Peridroma saucia (Hübner, 1808)*; Polyphaenis sericata (Esper, 1787)*; Sesamia nonagrioides (Lefèbvre, 1827)*; Praydidae: Prays fraxinella Bjerkander, 1784*; Pterophoridae: Crombrugghia laetus (Zeller, 1847); Emmelina monodactyla (Linnaeus, 1758); Stenoptilia zophodactyla Duponchel, 1840; Pyralidae: Pempelia palumbella (Denis & Schiffermüller, 1775)*; Tortricidae: Archips xylosteana (Linnaeus, 1758); Cnephasia sp. 1*; Cnephasia sp. 2*; Yponomeutidae: Zelleria oleastrella (Millière, 1864); Neuroptera 338 ISSN 1990-6471 (Hemerobiidae: Wesmaelius nervosus (Fabricius, 1793); Chrysopidae: Chrysoperla sp.*; Myrmeleontidae: Distoleon tetragrammicus (Fabricius, 1798)*). Of the 64 prey taxa, 32 (representing over 75 % of all food items) were consumed by both R. mehelyi and R. euryale. Benda et al. (2014c: 33) analysed 22 faecal pellets from Wadi Darnah (Libya) and found these to consist primarily of Blattoidea (99.1 % by volume) and 0.9 % of Lepidoptera. POPULATION: Structure and Density:- Hutson et al. (2008n) [in IUCN (2009)] and Alcaldé et al. (2016) indicate that Rhinolophus mehelyi is an infrequent species, which is reported to have declined in all parts of its range for which data are available. In Andalucia (Spain), the rate of decline has been estimated at 10 % over the last ten years. The species is close to extinction in France (Rodrigues and Palmeirim, 1999), Romania (Botnariuc and Tatole, 2005), and north-east Spain (J. Juste and T. Alcalde pers. Comm., 2006). In France, only one individual was recorded in 2004 (S. Aulagnier pers. Comm., 2006), and in Romania the population was estimated at 5,000 in the 1950s, but now numbers approximately 100 (Dumitrescu et al., 1962-3, Botnariuc and Tatole, 2005). It is also declining in southern Spain (Franco and Rodrigues de los Santos, 2001), Portugal (Rodrigues et al., 2003), the Russian Federation (K. Tsytsulina pers. Comm., 2005), Georgia, and Morocco (SW Asia Workshop 2005). In Iran mixed-species colonies including R. mehelyi, which in the 1970s were estimated to be over 10,000 individuals, now only number a few hundred individuals (M. Sharifi pers. obs., 2005). Summer nursery colonies typically number 30 - 500 individuals (although colonies of up to 3,000 individuals have been recorded, separated in smaller groups within the same cave). Winter clusters consist of up to 5,000 animals. Trend:- 2016: Decreasing (Alcaldé et al., 2016). 2008: Decreasing (Hutson et al., 2008n; IUCN, 2009). REPRODUCTION AND ONTOGENY: Benda et al. (2014c: 30) captured a lactating female on 15 May at Wadi Darnah (Libya). MATING: Puechmaille et al. (2011b: 40-41) found that during the mating season, females prefer males with higher frequency calls, indicating that this high frequency is probably a signal of good body condition. POSTNATAL DEVELOPMENT: In western Iran, Sharifi (2004: 155) determined that Fa and body mass followed a linear pattern of growth until day 14 (resp. 1.55 mm/day and 0.58 g/day) and subsequently decreased to reach a stable level. The length of the gap of the metacarpal-phalangeal joint showed an increase up to 10 days and decreased until it had closed at over 55 days. At about four weeks of age, the young started to fly. PARASITES: Beaucournu and Kock (1996) report the presence of the flea Rhinolophopsylla unipectinata arabs Jordan and Rothschild, 1921 on specimens from Algeria. Benda et al. (2014c: 33) recorded the following parasites from Libyan R. mehelyi: Ischnopsyllidae: Rhinolophopsylla unipectinata arabs Jordan and Rothschild, 1921, Nycteribiidae: Phthiridium biarticulatum (Hermann, 1864), Streblidae: Brachytarsina flavipennis Macquart, 1851 The following ectoparasites were found on Algerian bats by Bendjeddou et al. (2017: 15): Nycteribiidae: Nycteribia (Nycteribia) pedicularia Latreille, 1805 and Phthiridium biarticulatum Hermann, 1804; Streblidae: Brachytarsina (Brachytarsina) flavipennis Macquart, 1851; and Arachnida: Spinturnix myoti (Kolenati, 1856) and Eyndhovenia euryalis (G. Canestrini, 1885). From Iran, Sharifi et al. (2013: 181) mention the presence of the following ectoparasites: Eyndhovenia sp. (Acari: Spinturnicidae; on 89.65 % of the 87 bats examined, with 13.79 ind/bat, primarily on wing and tail membranes), Penicillidia sp. (Diptera: Nycteribiidae; 66.66 %, 2.31, mainly on fur) and one species from Streblidae (Diptera; 11.49 %, 1.80, on all parts of the body). In Azerbaijan, Duszynski (2002: 6) mentions Eimeria mehelyi Musaev and Gauzer, 1971 (Apicomplexa) as parasite. VIRUSES: Coronaviridae - Coronaviruses Luis et al. (2013: suppl.) mention the presence of a SARS coronavirus-related virus. Alphacoronavirus This virus was reported from Germany by Kohl and Kurth (2014: 3112). Betacoronavirus Kohl and Kurth (2014: 3112) report this virus from Bulgaria and Germany. African Chiroptera Report 2020 339 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Egypt, Morocco, Tunisia. Figure 104. Distribution of Rhinolophus mehelyi Rhinolophus mossambicus Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 *2012. Rhinolophus mossambicus Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, PLoS ONE, 7(9): 19. Publication date: 12 September 2012. Type locality: Mozambique: Nissa Game Reserve: Maputo Camp [121056S 373300E, 489 m] [Goto Description]. Holotype: DNSM 8578: ♂, skull and alcoholic. Collected by: Dr. Ara Monadjem; collection date: 1 July 2006. Presented/Donated by: ?: Collector Unknown. Baculum; skull and mandible in good condition (Taylor et al., 2012c: 19). Paratype: DNSM 11276: ad ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 8577: ad ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: DNSM 8579: ad ♂, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Baculum. Paratype: DNSM 8580: ad ♂, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Baculum. - Etymology: The name denotes the country of origin (Mozambique) of the type specimen of this species (Taylor et al., 2012c: 20). - ZooBank: 0D97DC39-F9CD-415EA7AE-4DCF70C807B4. (Current Combination) TAXONOMY: Based on mitochondrial and nuclear DNA sequences and morphological and morphometric data, Taylor et al. (2012c) found that the Rhinolophus hildebrandtii group includes more species than previously recognized. Carr et al. (2017) performed some additional analyses, which confirmed that R. mossambicus forms a well-defined clade based on mtDNA, but based on nuclear DNA, however, this taxon groups with R. fumigatus from Namibia. COMMON NAMES: English: Mozambican Horseshoe Bat. German: Mossambique-Hufeisennase. CONSERVATION STATUS: Global Justification Schoeman (2017) listed as Least Concern because it is relatively widespread in Mozambique and eastern Zimbabwe. Its extent of occurrence is >20,000 km2 and its area of occupancy (cave roosting sites) is probably >2,000 km 2. Although its population is probably decreasing through bushmeat hunting and guano mining, there is no evidence that the decrease has been rapid enough to warrant Near Threatened or threatened status. Assessment History Global 2016: LC ver. 3.1 (2001) (Schoeman, 2017). Regional None known. MAJOR THREATS: Schoeman (2017) report that this species is collected for bushmeat in Mozambique (particularly in the Inhassoro province). Additionally, the local communities disturb the roosts of this species with guano mining activities. 340 ISSN 1990-6471 CONSERVATION ACTIONS: R. mossambicus has been recorded from Niassa Game Reserve and Gorongosa National Park. Additional surveys are needed to learn more about the distribution, natural history and threats to this species. Existing cave roosts need to be protected from guano mining and hunting for bushmeat (S.M. Goodman and M.C. Schoeman unpublished data in Schoeman (2017). Other potential threats include habitat loss (due to charcoal collecting and logging) and fire (Schoeman, 2017). GENERAL DISTRIBUTION: Based on genetic data, the species is known from five localities in Mozambique from altitudes of 1,500 m - Chinizuia Forest, Gerhard’s Cave, Gorongosa Caves, Namapa, Niassa Game Reserve and one locality, Lutope-Ngolangola Confluence in NW Zimbabwe. These scattered records across the subregion suggest that R. mossambicus is widespread across the savanna biome of Zimbabwe and Mozambique (Taylor et al., 2012c: 20). PARASITES: DIPTERA: Streblidae: Raymondia aspera Maa, 1968 (Shapiro et al., 2016: 254). Raymondia waterstoni Jobling, 1931 (Shapiro et al., 2016: 256). VIRUSES: Astroviridae None of the 20 Mozambican bats tested by Hoarau et al. (2018: 2) was positive for this group of viruses. Coronaviridae Joffrin et al. (2020: 4) reported Alphacoronavirus from a Mozambican bat. an ZOOBANK: 0D97DC39-F9CD-415E-A7AE-4DCF70C807B4 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Malawi, Mozambique, Zimbabwe. POPULATION: Structure and Density:- There is no information on population size. Museum specimens have been collected from seven sites in Mozambique, and one locality in Zimbabwe (Monadjem et al., 2010c, 2010d; Taylor et al., 2012c; S.M. Goodman and M.C. Schoeman, unpublished data in Schoeman (2017)). Population sizes at roost sites were estimated between 50 – 400 (Monadjem et al., 2010d; S.M. Goodman and M.C. Schoeman unpublished data in Schoeman (2017)). Environmental niche models suggest additional roost sites may be present, but the localities are unknown (Taylor et al., 2012c). Trend:- 2016: Decreasing (Schoeman, 2017). Figure 105. Distribution of Rhinolophus mossambicus Rhinolophus rhodesiae Roberts, 1946 1946. 2018. Rhinolophus swinnyi rhodesiae Roberts, Ann. Transv. Mus., 20 (4): 304. Type locality: Zimbabwe: Limpopo valley: Bezwe river: a tributary of the Wanetsi (? Nuanetsi) River [ca. 21 22 S 30 45 E] [Goto Description]. Holotype: TM 1325: ad ♀. Collected by: Austin Roberts; collection date: 19 August 1913. Presented/Donated by: ?: Collector Unknown. - Etymology: The name refers to Southern Rhodesia (currently Zimbabwe), the country where the type specimen was collected. Rhinolophus rhodisiae: Hoarau, Le Minter, Joffrin, Schoeman, Lagadec, Ramasindrazana, Dos Santos, Goodman, Gudo, Mavingui, Lebarbenchon, Virology J., 15: Suppl.. Publication date: 30 June 2018. (Lapsus) TAXONOMY: Demos et al. (2019a: 11) resequenced the cyt-b of some of the specimens used by Taylor et al. (2018a) in the description of R. gorongosae and found that this diverged only between 1 and 1.4 % from that of R. simulator and R. rhodesiae. This "strongly suggests" that the genetic arguments for African Chiroptera Report 2020 341 raising R. swinnyi rhodesiae to full species were based on sequencing errors. ECHOLOCATION: Taylor et al. (2018a: 21) mention a peak frequency ranging from 99 to 102 kHz for eight bats from the Malashane cave (Mozambique). VIRUSES: Astroviridae None of the 31 Mozambican bats tested by Hoarau et al. (2018: 2) was positive for this group of viruses. Coronaviridae Joffrin et al. (2020: 4) reported Alphacoronavirus found in Mozambican bats. an DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Malawi, Mozambique, South Africa, Tanzania, Zambia, Zimbabwe. Figure 106. Distribution of Rhinolophus rhodesiae Rhinolophus ruwenzorii J. Eric Hill, 1942 *1942. Rhinolophus ruwenzorii J. Eric Hill, Am. Mus. Novit., 1180: 1, fig. 1. Publication date: 28 July 1942. Type locality: Congo (Democratic Republic of the): Kivu province: W slope of Mount Ruwenzori: Buhatu valley [00 22 N 29 44 E,, 7 500 ft] [Goto Description]. Holotype: AMNH 82394: ad ♀, skull and alcoholic. Collected by: James Paul Chapin, Dewitt L. Sage, Frank P. Mathews and The Ruwenzori-Kivu Expedition; collection date: 24 December 1926. - Comments: Fahr et al. (2002: 99) mention "S-side of Butahu Valley, 2286 m, W slope Rwenzori Mts., Congo (K.)." as type locality. - Etymology: Referring to the Ruwenzori mountains, where the type specimen was collected. (Current Combination) ? Rhinolophus maclaudi ruwenzorii: (Name Combination) TAXONOMY: Included in maclaudi by Koopman (1993a: 167), see Smith and Hood (1980: 170 ); but see Fahr et al. (2002), Csorba et al. (2003: 65 - 66) and Simmons (2005: 362). COMMON NAMES: Czech: vrápenec ruwenzorijský. English: Ruwenzori Horseshoe bat. French: Rhinolophe du Ruwenzori. German: Ruwenzori-Hufeisennase. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B1a+2b(ii,iii,iv,v) ver 3.1 (2001)) because its extent of occurrence is less than 20,000 km2, it is known from fewer than ten locations, and there is continuing decline in the extent and quality of its montane forest habitat, and probable loss of colonies through disturbance of roosting sites (Fahr, 2008m; IUCN, 2009). Assessment History Global 2008: VU B1a+2b(ii,iii,iv,v) ver 3.1 (2001) (Fahr, 2008m; IUCN, 2009). 2004: VU A4c; B1ab(iii) ver 3.1 (2001) (Fahr, 2004d; IUCN, 2004); and assessed as R. hilli by Fahr (2004f). Regional None known. MAJOR THREATS: Rhinolophus ruwenzorii is threatened by the deforestation of its montane habitat, generally through logging and mining activities, and the conversion of forest to farmland. It is dependent on cave habitats for roosting, and it is likely that disturbance of these sites could be an important threat to the species. It may be threatened by overharvesting for subsistence food, and while this needs to be fully confirmed populations are vulnerable to direct exploitation (Fahr, 2008m; IUCN, 2009). 342 ISSN 1990-6471 CONSERVATION ACTIONS: Fahr (2008m) [in IUCN (2009)] reported that Rhinolophus ruwenzorii has been recorded from some protected areas, such as the Mount Ruwenzori National Park. There is a need to reduce levels of deforestation within the Albertine Rift region as a whole. There is a need to initiate and develop public awareness programmes in the region, and to reduce the disturbance of roosting localities. Further field surveys are needed to locate additional population of this species. these total only eight distinct locations (Fahr, 2008m; IUCN, 2009). Trend:- 2008: Decreasing (Fahr, 2008m; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Rwanda, Uganda. GENERAL DISTRIBUTION: Rhinolophus ruwenzorii is restricted to the mountains of the Albertine Rift, where it has been recorded from the countries of the eastern Democratic Republic of the Congo, western Uganda, and Rwanda, at elevations of 1,666 to 2,666 m asl. The distribution includes the Ruwenzori Mountains, Kivu Region and KibaliIturi-Forest, Bwindi-Impenetrable Forest and Mutura in northwestern Rwanda (Fahr et al., 2002). Eastern Democratic Republic of the Congo, Western Uganda. Native: Congo (The Democratic Republic of the) Kityo and Kerbis (1996: 60); Rwanda (Fahr et al., 2002); Uganda Kityo and Kerbis (1996: 60). Figure 107. Distribution of Rhinolophus ruwenzorii POPULATION: Structure and Density:- This species is known only from 36 specimens collected from 13 localities, Rhinolophus sakejiensis Cotterill, 2002 *2002. Rhinolophus sakejiensis Cotterill, J. Zool., Lond., 256: 166, figs. 2a, 3, 4a, 5a, 6a, 7a. Type locality: Zambia: Mwinilunga district; Ikelenge Pedicle: btw Sakeji/Zambezi rivs,c.11 km NNE source Zambezi: Kavunda [11 17 S 24 21 E, 1 388 m] [Goto Description]. Holotype: NHMZ 29153: ad ♂, skull and alcoholic. Collected by: F.P.D. ("Woody") Cotterill; collection date: 11 October 1990; original number: FWC 1090. Baculum extracted. Paratype: HZM 1.32236: ad ♂, skull and alcoholic. Collected by: F.P.D. ("Woody") Cotterill; collection date: 11 October 1990; original number: FWC 1092. Baculum extracted. Paratype: NHMZ 29154: sad ♂, skull and alcoholic. Collected by: F.P.D. ("Woody") Cotterill; collection date: 11 October 1990; original number: FWC 1091. Baculum extracted. (Current Combination) TAXONOMY: Cotterill (2002b: 5) considers it a member of the ferrumequinum species group, and is followed by Simmons (2005: 362). See Csorba et al. (2003: 132 - 133) for remarks on taxonomy. COMMON NAMES: Czech: vrápenec sakežianský. English: Sakeji Horseshoe bat. French: Rhinolophe de Sakeji. German: Sakeji-Hufeisennase. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since it has only recently been discovered, and there is still very little information on its extent of occurrence, ecological requirements and threats (Cotterill, 2008a; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Cotterill, 2008a; IUCN, 2009). 2004: DD ver 3.1 (2001) (IUCN, 2004; Cotterill, 2004c). African Chiroptera Report 2020 Regional None known. MAJOR THREATS: The threats to Rhinolophus sakejiensis are not well known, but it is presumably threatened by the loss of roosting sites resulting from logging and the conversion of suitable habitat to agricultural use (Cotterill, 2008a; IUCN, 2009). CONSERVATION ACTIONS: Cotterill (2008a) [in IUCN (2009)] reported that it is not known if Rhinolophus sakejiensis is present in any protected areas. Further surveys are needed to better determine the extent of the distribution of this species, to gather more information on the species natural history, and to review the threats to this species. 343 POPULATION: Structure and Density:- Rhinolophus sakejiensis is currently known only from the type collection of three bats, taken from a cluster of six roosting bats (Cotterill, 2008a; IUCN, 2009). Trend:- 2008: Unknown (Cotterill, 2008a; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Zambia. GENERAL DISTRIBUTION: Rhinolophus sakejiensis is known only from the type locality of Kavunda, between the Sakeji and Zambezi rivers: c. 11 km north-north-east of the source of the Zambezi River in the Ikelenge Pedicle, Mwinilunga District of north-west Zambia (Cotterill, 2002a). It is likely that the species is also present within similar suitable habitat in Angola and the Congo basin (Cotterill, 2002a). Native: Zambia (Monadjem et al., 2010d: 560). Figure 108. Distribution of Rhinolophus sakejiensis Rhinolophus silvestris Aellen, 1959 *1959. Rhinolophus silvestris Aellen, Archs Sci. Genève, 12 (2): 228. Type locality: Gabon: Lastourville: N'Dumbu cave [ca. 00 50 S 12 03 E] [Goto Description]. Holotype: MHNG 965.040: ad ♂, skull and alcoholic. Collected by: Pierre Strinati and Villy Aellen; collection date: 4 August 1957. - Etymology: From the Latin for forest dweller (see Flannery, 1995a: 321). (Current Combination) 2016. Rhinolophus sylvestris: Karadjian, Hassanin, Saintpierre, Gembu Tungaluna, Ariey, Ayala, Landau and Duval, PNAS, 113 (35): 9835. Publication date: 30 August 2016. (Lapsus) ? Rhinolophus clivosus silvestris: (Name Combination) TAXONOMY: Considered a subspecies of clivosus by Hayman and Hill (1971: 23), but Barratt et al. (1995: 386), Cotterill (2002a), Csorba et al. (2003: 46 - 48) and Simmons (2005: 363), which indicate the relationships of Rhinolophus deckeni and R. silvestris are unclear, with the forms possibly being conspecific. Considered part of the ferrumequinum species group (Cotterill, 2002a; Csorba et al., 2003: 46 - 48; Simmons, 2005: 363). COMMON NAMES: English: Forest Horseshoe Bat. French: Rhinolophe sylvestre. German: WaldHufeisennase. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Cotterill, 2008b; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Cotterill, 2008b; IUCN, 2009). 2004: VU B2ab(iii) ver 3.1 (2001) (Cotterill, 2004g; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. 344 ISSN 1990-6471 MAJOR THREATS: Rhinolophus silvestris is considered to be threatened by habitat loss through conversion of land to agricultural use and logging operations. Additional threats include general disturbance of cave roost sites and overharvesting of the species for the bushmeat trade (Cotterill, 2008b; IUCN, 2009). CONSERVATION ACTIONS: Cotterill (2008b) [in IUCN (2009)] reported that Rhinolophus silvestris does not appear to have been recorded from any protected areas. An additional examination of the taxonomic relationship between this species and Rhinolophus deckeni is needed. Further field surveys are needed to better determine the distribution, natural history and the extent of threats to this little-known bat. GENERAL DISTRIBUTION: Rhinolophus silvestris a poorly known species has only been recorded from three localities in Gabon (Benga; Belinga; and the type locality of Dumbu Cave, Latoursville), and a single site in Congo (Meya-Nzouari). clivosus sylvestris. About 50 % of the specimens from Congo they examined, were found to be infected. Gametocytes were found year-round. The bats were also infested by the nycteribiids Penicillidia fulvida Bigot, 1885 and Nycteribia schmidlii scotti Falcoz, 1923, of which the prior was a major carrier of Polychromophilus sporozoites. The Haemosporidan Nycteria gabonensis Rosin, Landay & Hugot, 1978 was reported by Landau et al. (2012: 142), Perkins and Schaer (2016: Suppl.) and Karadjian et al. (2016: 9835) from the Republic of Congo and Gabon. Justine (1989a: 334; 1989b: 553) described two new stomach parasites: Capillaria chabaudi and Capillaria gabonensis (Nematoda) from R. silvestris from Belinga (Gabon). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo, Gabon. Native: Congo (Bates et al., 2013: 335); Gabon. POPULATION: Structure and Density:- Rhinolophus silvestris is usually found in small colonies of fewer than 20 bats. It is believed to be in overall decline (Cotterill, 2008b; IUCN, 2009). Trend:- 2008: Decreasing (Cotterill, 2008b; IUCN, 2009). PARASITES: Adam and Landau (1973a: 5) report on the presence of a protozoan parasite of the genus Polychromophilus (Haemoproteidae) in R. Figure 109. Distribution of Rhinolophus silvestris Rhinolophus simulator K. Andersen, 1904 *1904. Rhinolophus simulator K. Andersen, Ann. Mag. nat. Hist., ser. 7, 14 (83): 384. Publication date: 1 November 1904. Type locality: Zimbabwe: Mashonaland: Mazoe [17 30 S 31 03 E, 4 000 ft] [Goto Description]. Holotype: BMNH 1902.2.7.10: ad ♂, skull and alcoholic. Collected by: J.ff. Darling Esq. - Etymology: From the masculine Latin substantive simulàtor, meaning "simulator", as R. simulator, according its descriptor, "[... ] by quite superficial inspection, could be taken for a curiously small and long-tailed form of [R. capensis]." (see Lanza et al., 2015: 115). (Current Combination) 1914. Rhinolophus bembanicus Senna, Annuario Mus. Zool. R. Univ. Napoli, (2) 4: no 9, p. 1, 2, text-fig 1, 2. Publication date: 27 March 1914. Type locality: Zimbabwe: Lake Bengueolo [=Lake Bangweulu]. - Comments: Ansell and Dowsett (1988: 34) indicate that bembanicus might represent a separable species, but the type specimen no longer exists. 1936. Rhinolophus alcyone alticolus Sanborn, Field Mus. Nat. Hist., Zool. Ser., (362) 20 (14): 108. Publication date: 15 August 1936. Type locality: Cameroon: Mount Cameroon [04 13 N 09 11 E, 5 800 ft] [Goto Description]. Holotype: FMNH 42596: ad ♀. Collected African Chiroptera Report 2020 ? ? ? 345 by: Rudyerd Boulton and Laura Boulton; collection date: 2 July 1934; original number: 53. - Comments: Fahr (2007a: 103) refers to Csorba et al. (2003), who indicate that alticolus might possibly represent a separate species. Rhinolophus alticolus: Rhinolophus simulator alticolus: (Name Combination) Rhinolophus simulator simulator: (Name Combination) TAXONOMY: Figure 110. Rhinolophus simulator from Madimatle Cave, South Africa. Includes alticolus and bembanicus (see Koopman, 1975: 387; Hayman and Hill, 1971: 25; Koopman, 1993a: 169; Simmons, 2005: 363). Considered part of the capensis species group (Csorba et al., 2003: 10 - 12; Simmons, 2005: 363). Monadjem et al. (2016y: 366) refer to Csorba et al. (2003), who indicated that alticolus might represent a separate species, as their specimens from the Mount Nimba area are about 2,000 km from the nearest records in Nigeria and Cameroon, which, in turn, are again thousands of kilometers from the main distribution area. However, they still retain alticolus as a subspecies of simulator. COMMON NAMES: Afrikaans: Bosveldsaalneusvlermuis, Bosveldvlermuis. Chinese: 布 什 维尔德菊头蝠. English: Bushveld Horseshoe Bat. French: Rhinolophe de brousse, Rhinolophe du Veld, Rhinolophe du Bushveld. German: BuschveldHufeisennase, Buschland-Rundblattnase. Italian: Rinòlofo sirnulatóre, Fèrro di cavàllo simulatóre. Portuguese: Morcego ferradura das savanas. ETYMOLOGY OF COMMON NAME: The colloquial name refers to its occurrence in savanna woodland in the subregion (see Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: R. simulator may be present in the late Holocene at Nkupe, South Africa (Avery, 1991: 6). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008ao; IUCN, 2009; Monadjem et al., 2017cm). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cm). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008ao; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004al; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016l). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Rhinolophus simulator is threatened in parts of its range by cave disturbance and the loss of habitat resulting from the conversion of land to agricultural use and mining operations (Jacobs et al., 2008ao; IUCN, 2009; Monadjem et al., 2017cm). CONSERVATION ACTIONS: Jacobs et al. (2008ao) [in IUCN (2009)] and Monadjem et al. (2017cm) reported that there appear to be no direct conservation measures in place. In view of the species East African and southern African distribution, it seems likely that it is present in some protected areas. GENERAL DISTRIBUTION: Rhinolophus simulator is widely distributed in subSaharan Africa. There are records in West Africa from the Wonegizi Mountains and Mount Nimba (Guinea and Liberia), and from central Nigeria. In Central Africa it has been reported from Cameroon. In East Africa there are many more records, with the species being reported from as far north as Ethiopia and southern Sudan, ranging southwards through Uganda, Kenya and Tanzania to Zambia, Malawi, Mozambique, Zimbabwe, eastern South Africa, Swaziland and southern Botswana. In South Africa, its distribution is strongly associated with the amount of 346 ISSN 1990-6471 precipitation in the warmest quarter of the year (Babiker Salata, 2012: 49). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 855,537 km2. Native: Botswana (Skinner and Smithers, 1990; Csorba et al., 2003: 12; Monadjem et al., 2010d: 560); Cameroon (Rosevear, 1965; Csorba et al., 2003: 12); Côte d'Ivoire; Ethiopia (Hill and Morris, 1971; Largen et al., 1974; Csorba et al., 2003: 12); Guinea (Brosset, 1985; Csorba et al., 2003: 12: Denys et al., 2013: 282); Kenya (Koopman, 1975; Aggundey and Schlitter, 1984; Csorba et al., 2003: 12); Liberia; Malawi (Ansell and Dowsett, 1988; Happold et al., 1988; Happold and Happold, 1997b: 818; Csorba et al., 2003: 12; Monadjem et al., 2010d: 560); Mozambique (Smithers and Lobão Tello, 1976: 561; Csorba et al., 2003: 12; Monadjem et al., 2010c: 380); Nigeria (Rosevear, 1965 Happold, 1987; Csorba et al., 2003: 12); South Africa (Ellerman et al., 1953; Taylor, 1998; Rautenbach et al., 1984; Maree and Grant, 1997; Csorba et al., 2003: 11; Monadjem et al., 2010d: 561); Sudan (Koopman, 1975; Csorba et al., 2003: 12); Swaziland (Monadjem, 2005: 6; Monadjem et al., 2010d: 561); Tanzania (Aellen, 1957; Csorba et al., 2003: 12; Stanley and Goodman, 2011: 42; Stanley and Goodman, 2011: 42); Uganda; Zambia (Ansell, 1967; Ansell, 1969; Ansell, 1973; Ansell, 1978; Csorba et al., 2003: 12; Cotterill, 2004a: 261; Monadjem et al., 2010d: 561) ; Zimbabwe (Smithers and Wilson, 1979; Csorba et al., 2003: 12; Monadjem et al., 2010d: 561). ECHOLOCATION: Jacobs (1996) and Fenton and Bell (1981) reported a maximum frequency of 78 kHz for an individual from Sengwa in Zimbabwe. Monadjem et al. (2007a), using an Anabat detector, reported a maximum frequency of 84.1 (± 0.36) kHz for individuals from various localities in Swaziland. Additional parameters from the same country were given by Monadjem et al. (2017c: 179): Fmin: 72.1 ± 2.97 (68.6 - 76.0) kHz, Fknee: 83.8 ± 0.18 (83.6 84.1) kHz, Fc: 83.4 ± 0.20 (83.7 - 84.2) kHz, and duration: 16.8 ± 4.70 (12.0 - 25.9) msec. For individuals in South Africa, Taylor (1999b), using a Pettersson D980, reported a maximum frequency at Shongweni of 82.7 (± 0.4) kHz, while Jacobs et al. (2007a) reported a peak frequency of 80.6 (± 1.0) kHz (detector type and locality not indicated). Taylor (2000) reported a constant frequency component at 83 kHz. Mutumi and Jacobs (2013: 108) investigated the peak frequency over the entire distribution range and found that this frequency was not correlated to body size or vegetation index, but in one location (where the mean annual rainfall and temperatures where the highest), they found that the peak frequency was significantly lower. They suggest that this lower frequency was linked with an increased detection range due to the more humid conditions. The resting call frequency was studied by Mutumi et al. (2016), who concluded that the acoustic divergence was best predicted by latitude, geography and climate-induced differences in atmospheric attenuation. Taylor et al. (2013b: 19) reported 9 calls from Waterberg, South Africa with the following characteristics: Fmax: 82.0 ± 0.58 (81.1 - 82.9) kHz, Fmin: 79.3 ± 1.42 (75.9 - 80.6) kHz, Fknee: 80.5 ± 0.46 (79.8 - 81.2) kHz, Fchar: 81.5 ± 0.47 (80.9 82.1) kHz, duration: 12.4 ± 1.71 (9.6 - 14.4) msec. An additional 16 calls from Swaziland had the following parameters: Fmax: 84.0 ± 0.34 (83.2 84.4) kHz, Fmin: 78.6 ± 3.77 (71.5 - 82.3) kHz, Fknee: 83.1 ± 0.83 (80.3 - 83.7) kHz, Fchar: 83.4 ± 1.20 (79.2 - 84.1) kHz, duration: 13.1 ± 3.66 (7.0 - 18.9) msec. Linden et al. (2014: 40) reported the following parameters from the Soutpansberg range (RSA): Fmin: 76 - 81 kHz, Fpeak: 79 - 84 kHz, Fknee: 80 - 84 kHz, Slope: 70 - 368 OPS, duration: 7 - 19 msec. 20 calls from Mapungubwe National Park (RSA) recorded by Parker and Bernard (2018: 57) had the following characteristics: Fchar: 79.70 ± 1.02 kHz, Fmax: 79.86 ± 0.99 kHz, Fmin: 75.43 ± 4.93 kHz, Fknee: 79.71 ± 1.01 kHz, duration: 37.79 ± 9.97 msec, with 7.46 ± 3.46 calls/sec. Jacobs (2016: 119) indicates that the second harmonic of this bat coincides with the third harmonic of R. capensis. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach (1986) reported 2n = 58, FN = 60, BA = 4, and submetacentric X and Y chromosomes. Protein / allozyme - Unknown. ROOST: Taylor et al. (1999: 70) reported a colony of 50 - 60 bats roosting in the Shongweni Dam tunnel (RSA). DIET: From Sudwala (RSA), Jacobs (2000: 201) reported the following proportions of prey: Lepidoptera (57.6 ± 34.9), Coleoptera (18.3 ± 23.0), Isoptera (19.4 ± 29.8), Hymenoptera (1.5 ± 4.1), Diptera (2.2 ± 4.2), Hemiptera (0.2 ± 1.1), Mandotea (0.2 ± 0.7), and unknown (0.6 ± 2.5). Weier et al. (2019b: 6) examined droppings from this species in the Levubu region (Limpopo, RSA) African Chiroptera Report 2020 and found DNA sequences of Nezara viridula (Linnaeus, 1758) (Hemiptera). PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- Rhinolophus simulator can be locally common. Colonies may consist of a couple of hundred individuals, although these are often quite fragmented because of the species association with caves for breeding (Jacobs et al., 2008ao; IUCN, 2009; Monadjem et al., 2017cm). Trend:- 2016: Decreasing (Monadjem et al., 2017cm). 2008: Decreasing (Jacobs et al., 2008ao; IUCN, 2009). REPRODUCTION AND ONTOGENY: Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] reported that births take place around mid November. In Malawi, the births took place early in the wet season (Happold and Happold, 1990b: 566). 347 VIRUSES: Polyomaviridae Carr et al. (2017) reported two new viruses from R. blasii from Zambia: Rhinolophus simulator polyomavirus 4 and Rhinolophus simulator polyomavirus1, besides the two already known: Rhinolophus simulator polyomavirus 2 and Rhinolophus simulator polyomavirus 3 Reoviridae Sasaki et al. (2016: 2489) retrieved a VP7 type Rotavirus from a Zambian R. simulator: RVA/Batwt/ZMB/ LUS12-14/2012/G3P[3] (LUS12-14). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Botswana, Cameroon, Congo (Democratic Republic of the), Eswatini, Ethiopia, Guinea, Kenya, Liberia, Malawi, Mozambique, Nigeria, South Africa, South Sudan, Tanzania, Zambia, Zimbabwe. PARASITES: BACTERIA: Dietrich et al. (2016a: 4) found Rickettsia in the faeces of bats from South Africa. Clément et al. (2019: 5) reported on 27 R. simulator specimens of which 2 were infected by Trypanosoma livingstonei_A and T. livingstoneii_C. Streblidae: Brachytarsina africana (Walker, 1849) has a wide distribution in sub Saharan Africa (Haeselbarth et al., 1966: 100). Raymondia hardyi Fiedler, 1954 from South Africa (Shapiro et al., 2016: 254). Raymondia waterstoni Jobling, 1931 (Haeselbarth et al., 1966: 104; Shapiro et al., 2016: 256). Figure 111. Distribution of Rhinolophus simulator Rhinolophus smithersi Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, 2012 *2012. Rhinolophus smithersi Taylor, Stoffberg, Monadjem, Schoeman, Bayliss and Cotterill, PLoS ONE, 7(9): 19. Publication date: 12 September 2012. Type locality: Zimbabwe: Sebungwe: Gokwe Communal Land: Ngolangola Gorge at confluence with Lutope River [181705S 280500E, 1000 m] [Goto Description]. Holotype: NHMZ 33647: ♀, skull and alcoholic. Collected by: F.P.D. ("Woody") Cotterill and Peter John Taylor; collection date: 26 October 2000; original number: FWC 4764. Presented/Donated by: ?: Collector Unknown. Skull and mandible in good condition ( Taylor et al., 2012c: 19). Paratype: NHMZ 33652: ♂, skull and alcoholic. Collected by: F.P.D. ("Woody") Cotterill and Peter John Taylor; collection date: 26 October 2000; original number: FWC 4769. Presented/Donated by: ?: Collector Unknown. - Etymology: Taylor et al. (2012c: 19) selected the specific epithet in recognition of the late Reay Henry Noble Smithers (1907– 1987), former Director of the National Museums of Zimbabwe, prodigious collector and 348 ISSN 1990-6471 2015. researcher of mammals including bats, and author of important regional texts on the mammalogy of Botswana, Zimbabwe and Mozambique, notably his definitive monograph – The Mammals of the Southern African Subregion subsequently updated and revised. The species name combines the surname Smithers and genitive singular case-ending ‘‘i’’ indicative of masculine gender. - ZooBank: E78B0F44-7534-4991-975A-F176CB668DDE. (Current Combination) [Rhinolophus] smthersi: Kaipf, Rudolphi and Meinig, NABU - Biodiversity Assessment in Kafa, Ethiopia, 13. TAXONOMY: Based on mitochondrial and nuclear DNA sequences and morphological and morphometric data, Taylor et al. (2012c) found that the Rhinolophus hildebrandtii group includes more species than previously recognized. COMMON NAMES: English: Smithers's Horseshoe Bat. Smithers Hufeisennase. German: CONSERVATION STATUS: Global Justification Taylor (2017) reports that the species does not trigger any thresholds for any threatened categories under any of the criteria. Recent collecting has shown that, although essentially limited to one province of South Africa (with an isolated small Zimbabwean population), the species is quite widespread so areas of occupancy and extent of occurrence exceed the thresholds for Vulnerable under criterion B. There is no evidence for declines in the species or its habitat. However, colony sizes are very small and the species appears to have a scattered occurrence in the landscape (e.g., it is rarely collected with harp traps or recorded with bat detectors during surveys of the Soutpansberg). Since it is limited by availability of roosting sites, and possibly suitable water sources, it is hard to believe that the entire population of the species could be much more than 1,000 individuals, so it could potentially be Vulnerable under criterion D1. Until the popualtion size can be confirmed to be below 1,000, however, the species is assessed as Near Threatened (nearly meets VU D1). Assessment History Global 2016: NT D1 ver 3.1 (2001) (Taylor, 2017). Regional South Africa:- 2016: NT D1 ver 3.1 (2001) (Taylor et al., 2016b). MAJOR THREATS: Taylor (2017) report that there are no known major threats to this species at present as much of its range occurs throughout the Soutpansberg, Blouberg and Waterberg ranges of Limpopo where human impacts and habitat transformation are not yet severe. The threat of extensive planned coal, platinum, natural gas (fracking) and other mining developments over much of the Limpopo Valley and the foothills of the Soutpansberg and Waterberg mountains and the Mahabeng Plateau, could impact heavily on populations, e.g. through roosting and foraging habitat loss, noise, air and water pollution and water abstraction leading to degradation of riparian habitats. Suppression of fire together with over-grazing of cattle and game and climate change has resulted in serious bush encroachment of savannas across much of its range (e.g. the western Soutpansberg) which has been shown to have a negative effect on biodiversity generally. In the eastern Soutpansberg, afforestation and alien plant invasions have considerably altered natural habitats. Limpopo is extremely drought-prone and water-stressed and with projected climate change, since the species seems to be dependent on water sources for drinking, extreme droughts in the area have had potentially devastating effects on wildlife generally. CONSERVATION ACTIONS: Taylor (2017) report that not counting the Zimbabwean population, the entire South African range of the species is included within two UNESCO Biosphere Reserves, the Vhembe and Waterberg Biosphere Reserves. Defining and refining core and buffer areas is a critical part of the ongoing management of these Reserves. Roosting sites of bats should be included as a layer in determining the location of such zones within biosphere reserves. This means that important bat underground (natural and manmade) roosts (including those of R. smithersi) should be included wherever possible in core or buffer areas where developments (including mining) would have to be regulated. Such conservation zonation plans (including Strategic Environmental Frameworks, EMFs) would inform planning by Provincial Nature Conservation (including the protected areas expansion strategy) and municipal Integrated Development Plans (IDPs), thereby affording protection to roosting sites. African Chiroptera Report 2020 GENERAL DISTRIBUTION: Known from Lutope-Ngolangola Gorge south of the Zambezi Escarpment in NW Zimbabwe and also from Pafuri in the Limpopo Valley in the foothills of the Soutpansberg Mountains of northern Limpopo Province, South Africa but likely more widespread across savanna woodlands of the Limpopo and Zambezi valleys, and their escarpments (the Gwembe horst, and the Soutpansberg and Waterberg Mountains, respectively) (Taylor et al., 2012c: 19). Accurate delimitation of this species’ range is subject to further collecting and reappraisal of existing museum material, previously refered to as R. hildebrandtii. Native: South Africa (Taylor et al., 2012c: 19); Zimbabwe (Taylor et al., 2012c: 19); Malawi (tentavively, all of the material previously assigned to R. hildebrandtii, but Meredith Happold (pers. comm., 24/02/2019) indicated that two species might be involved). Kaipf et al. (2015: 22) report "Rhinolophus smithersi or a new species" from God's Bridge in Ethiopia. ECHOLOCATION: For 7 calls from specimens from Blouberg, South Africa, Taylor et al. (2013b: 18) recorded the following parameters: Fmax: 45.2 ± 0.17 (44.9 45.4) kHz, Fmin: 40.6 ± 1.28 (38.1 - 41.7) kHz, Fknee: 44.7 ± 0.13 (44.5 - 44.8) kHz, Fchar: 44.6 ± 0.13 (44.3 - 44.7) kHz, duration: 34.9 ± 6.69 (26.0 - 46.9) msec. 13 calls from Waterberg resulted in the following parameters: Fmax: 47.8 ± 0.31 (47.0 - 48.4) kHz, Fmin: 45.7 ± 0.91 (44.1 - 44.7) kHz, Fknee: 47.2 ± 0.31 (46.3 - 47.7) kHz, Fchar: 47.3 ± 0.29 (46.4 - 47.7) kHz, duration: 18.2 ± 4.52 (10.9 - 24.8) msec, and 10 calls from Soutpansberg: Fmax: 46.0 ± 0.12 (45.9 - 46.2) kHz, Fmin: 44.8 ± 0.20 (44.4 - 45.0) kHz, Fknee: 45.2 ± 0.07 (45.2 45.4) kHz, Fchar: 45.5 ± 0.05 (45.5 - 45.6) kHz, duration: 12.0 ± 2.22 (9.6 - 16.5) msec. 19 calls recorded at the Mapungubwe National Park (RSA) by Parker and Bernard (2018: 57) had the following characteristics: F char: 47.23 ± 1.62 kHz, Fmax: 47.47 ± 1.50 kHz, F min: 43.27 ± 2.65 kHz, Fknee: 47.26 ± 1.54 kHz, duration: 44.74 ± 22.83 msec, with 8.02 ± 3.70 calls/sec. POPULATION: Structure and Density:- Rhinolophus smithersi has colony sizes very small but more precise information about population size is missing (Taylor, 2017). Trend:- 2016: Stable (Taylor, 2017). 349 PARASITES: DIPTERA: Streblidae: Ascodipteron brevior Maa, 1965, female cysts found at the base of the ears from Namibia and South Africa (Haeselbarth et al., 1966: 106 host referred to as R. hildebrandtii). Raymondia hardyi Fiedler, 1954 from South Africa (Shapiro et al., 2016: 254; based on a record by Theodor (1968), who mentioned "R. hildebrandtii" as host). Nycteribiidae: Nycteribia scissa rhodesiensis Theodor, 1957 from Zimbabwe and Malawi (Haeselbarth et al., 1966: 108, host as R. hildebrandtii). Nycteribia rotundata Theodor, 1957 from Mazoe, Zimbabwe (Haeselbarth et al., 1966: 110, host referred to as R. hildebrandtii). Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114, host referred to as R. hildebrandtii). ZOOBANK: E78B0F44-7534-4991-975A-F176CB668DDE DISTRIBUTION BASED ON VOUCHER SPECIMENS: Botswana, Malawi, Mozambique, South Africa, Zambia, Zimbabwe. Figure 112. Distribution of Rhinolophus smithersi 350 ISSN 1990-6471 Rhinolophus swinnyi Gough, 1908 *1908. Rhinolophus swinnyi Gough, Ann. Transv. Mus., 1: 71, 2 text-f. Publication date: April 1908. Type locality: South Africa: Cape province: Ngqeleni district: W Pondoland: "Ngqeleni district" [30 40 S 29 02 E] [Goto Description]. Holotype: TM 1021: ad ♂, skin and skull. Collected by: H.H. Swinny; collection date: 22 February 1908; original number: 153. Paratype: TM 1022: ad ♂, skin and skull. Collected by: H.H. Swinny; collection date: 23 February 1908; original number: 154. Notes: skull found lying loose- may not belong to this specimen [signed A. Roberts]). - Comments: Meester et al. (1986: 41) mentioned the type locality as "Ngqeleni district, west of Port St. Johns, Transkei". (Current Combination) 1913. Rhinolophus swinnyi piriensis Hewitt, Records Albany Mus., 2: 402. Publication date: 6 February 1913. Type locality: South Africa: E Cape province: near King William's Town: Pirie [32 43 S 27 18 E]. TAXONOMY: Koopman (1966, 1982) suggests that this species may be conspecific with denti. capensis species group (Csorba et al., 2003: 12 - 14; Simmons, 2005: 364). See Csorba et al. (2003: 11) for remarks on taxonomy. COMMON NAMES: Afrikaans: Swinny se saalneusvlermuis, Swinnysaalneusvlermuis, Swinnyse Vlermuis. Czech: vrápenec mosambický. English: Swinny's Horseshoe Bat. French: Rhinolophe de Swinny. German: Swinnys Hufeisennase. Portuguese: Morcego ferradura de Swinny. ETYMOLOGY OF COMMON NAME: Named after H.H. Swinny, who collected the original specimens from the Nqqueleni district, Eastern Cape (Taylor, 2005). CONSERVATION STATUS: Global Justification Although Rhinolophus swinnyi is known mainly from sparse records from a large area, it is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large overall population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Cotterill, 2008c; IUCN, 2009; Monadjem and Cotterill, 2017b). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Cotterill, 2017b). 2008: LC ver 3.1 (2001) (Cotterill, 2008c; IUCN, 2009). 2004: NT ver 3.1 (2001) (Cotterill, 2004h; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: VU C2a(i) ver 3.1 (2001) (Jacobs et al., 2016h). 2004: EN C2a(i) ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: Populations may be locally threatened by deforestation, largely resulting from logging operations, local use of timber and firewood, and general conversion of land to agricultural use (Cotterill, 2008c; IUCN, 2009; Monadjem and Cotterill, 2017b). CONSERVATION ACTIONS: Cotterill (2008c) [in IUCN (2009)] and Monadjem and Cotterill (2017b) reported that Rhinolophus swinnyi is present in some protected areas (e.g. Kruger National Park, South Africa). Further studies are needed into the taxonomic status of this species, the distribution and possible threats. GENERAL DISTRIBUTION: Rhinolophus swinnyi has been recorded from eastern parts of South Africa, much of Zimbabwe, northwestern Mozambique, with additional scattered records further north in Malawi, Zambia, Democratic Republic of the Congo and Tanzania (including the island of Unguja [=Zanzibar]) (Skinner and Chimimba, 2005; O'Brien, 2011: 286). It may be present in Angola, but this needs confirmation. In South Africa, its distribution is mainly affected by temperature seasonality (Babiker Salata, 2012: 49). Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,054,284 km2. Native: Congo (The Democratic Republic of the) (Hayman et al., 1966; Happold et al., 1988; Kock African Chiroptera Report 2020 and Howell, 1988; Csorba et al., 2003: 14; Monadjem et al., 2010d: 561); Malawi (Happold and Happold, 1997b: 818; Csorba et al., 2003: 14); Mozambique (Smithers and Lobão Tello, 1976; Skinner and Smithers, 1990; Csorba et al., 2003: 14; Monadjem et al., 2010d: 561; Monadjem et al., 2010c: 380); South Africa (Maree and Grant, 1997; Rautenbach and Espie, 1982; Kock and Howell, 1988; Taylor, 1998; Csorba et al., 2003: 13; Monadjem et al., 2010d: 561); Tanzania (Skinner and Smithers, 1990; Kock and Howell, 1988; Csorba et al., 2003: 14); Zambia (Ansell, 1969; Ansell, 1978; Csorba et al., 2003: 14; Monadjem et al., 2010d: 561); Zimbabwe (Cotterill, 1996a; Cotterill, 2004a: 261; Csorba et al., 2003: 14; Monadjem et al., 2010d: 561). Presence uncertain: Angola 351 Trend:- 2016: Unknown (Monadjem and Cotterill, 2017b). 2008: Unknown (Cotterill, 2008c; IUCN, 2009). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: ECHOLOCATION: Aldridge and Rautenbach (1987) reported a maximum frequency of 115 kHz for individuals from Pafuri in South Africa. Jacobs et al. (2007a) reported a slightly different peak frequency of 107.0 (± 0.8) kHz for individuals from South Africa. Monadjem et al. (2010c: 380) reported from Mozambique that the peak echolocation frequencies ranged between 99 - 103 kHz (Petterson D240x). Taylor et al. (2018a: 25) mention a mean frequency of 106.6 ± 0.4 kHz for 10 specimens from the Eastern Cape Province and 105.6 ± 0.76 kHz from KwaZulu-Natal. Mutumi et al. (2016) found that acoustic divergence in RF was best predicted by latitude, geography and climate-induced differences in atmospheric attenuation. Jacobs (2016: 119) indicates that the second harmonic of this bat coincides with the third harmonic of R. capensis, which possibly might illustrate "harmonic hopping". Figure 113. Clockwise from top lateral, ventral and dorsal skull images of TM 1021, holotype of Rhinolophus swinnyi Gough, 1908. PARASITES: DIPTERA: Streblidae: Raymondia waterstoni Jobling, 1931 (Shapiro et al., 2016: 256). DISTRIBUTION BASED ON VOUCHER SPECIMENS: South Africa. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach (1986) reported 2n = 58, FN = 62, BA = 6, a subtelocentric X chromosome and a metacentric Y chromosome. Protein / allozyme - Unknown. POPULATION: Structure and Density:- In parts of its range Rhinolophus swinnyi is considered to be uncommon, however, Taylor (2000) records that it is fairly common in Zimbabwe. It generally forms small colonies of fewer than ten animals (Cotterill, 2008c; IUCN, 2009; Monadjem and Cotterill, 2017b). Figure 114. Distribution of Rhinolophus swinnyi 352 ISSN 1990-6471 Rhinolophus willardi Kerbis Peterhans and Fahr, 2013 *2013. Rhinolophus willardi Kerbis Peterhans and Fahr, in: Kerbis Peterhans, Fahr, Huhndorf, Kaleme, Plumptre, Marks and Kizungu, Bonn. Zool. Bull., 62 (2): 190, figs 3, 4, 6. Publication date: November 2013. Type locality: Congo (Democratic Republic of the): Misotschi-Kabogo highlands [05 06 19 S 29 03 56 E, 1 880 m] [Goto Description]. Holotype: FMNH 195182: ad ♂, skull and alcoholic. Collected by: A.J. Plumptre and E.A. Mulungu; collection date: 17 February 2007; original number: DCM 1680. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 195082: ad ♀, skin and skeleton and skull. Collected by: B.D. Marks; collection date: 14 February 2007; original number: MHH 837. Presented/Donated by: ?: Collector Unknown. From 4 km SW of the village Talama, "Camp 2", 4°59’29’’S, 29°04’49’’E, 1950 m. Paratype: FMNH 195083: ad ♀, skin and skeleton and skull. Collected by: B.D. Marks; collection date: 14 February 2007; original number: MHH 838. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 195084: ad ♀, skin and skeleton and skull. Collected by: Prince Kiswele Kaleme; collection date: 16 February 2007; original number: PK 754. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 198083: ad ♀, complete skeleton. Collected by: B.D. Marks; collection date: 14 February 2007; original number: MHH 838. Presented/Donated by: ?: Collector Unknown. From 4 km SW of the village Talama, "Camp 2", 4°59’29’’S, 29°04’49’’E, 1950 m. Paratype: FMNH 198084: ad ♀, complete skeleton. Collected by: B.D. Marks; collection date: 16 February 2007; original number: PK 754. Presented/Donated by: ?: Collector Unknown. From 4 km SW of the village Talama, "Camp 2", 4°59’29’’S, 29°04’49’’E, 1950 m. - Etymology: In honour of Dr. David Willard (Collection Manager, Division of Birds, FMNH) in recognition of his unparalleled 35+ years of service to the Field Museum of Natural History (see Kerbis Peterhans et al., 2013: 191). (Current Combination) COMMON NAMES: English: Willards horseshoe bat. Willards Hufeisennase. German: DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the). Figure 115. Distribution of Rhinolophus willardi Rhinolophus ziama Fahr, Vierhaus, Hutterer and Kock, 2002 *2002. Rhinolophus ziama Fahr, Vierhaus, Hutterer and Kock, Myotis, 40: 102, 109, figs 1 - 7. Type locality: Guinea: Guinée Forestière: border "Réserve Biosphère de la Massif du Ziama": Sérédou, W edge of: near Park Station [ca. 08 23 N 09 17 W] [Goto Description]. Holotype: ZFMK MAM-1999.0934: ad ♂, alcoholic, skull and skeleton. Collected by: Henning Vierhaus; collection date: 12 August 1992; original number: HV 2590. Paratype: AMNH 265708: ad ♂, skull and alcoholic. Collected by: Dr. Robert W. Dickerman; collection date: 4 March 1990; original number: 2144. 7 mi ?, 1 mi E Ziggida, Lofa county, Wonegizi Mts., Liberia: see Fahr et al. (2002: 109). - Etymology: In reference to the African Chiroptera Report 2020 353 protected area near the type locality, the "Réserve de la Biosphère du Massif de Ziama", and as a noun in apposition (Fahr et al., 2002: 110). (Current Combination) TAXONOMY: maclaudi species group (Fahr et al., 2002; Simmons, 2005: 365). COMMON NAMES: Czech: vrápenec liberijský. English: Ziama Horseshoe Bat (first use Simmons, 2005: 365). French: Rhinolophe du Ziama. German: ZiamaHufeisennase. CONSERVATION STATUS: Global Justification Rhinolophus ziama is listed as Endangered (EN B1ab(iii) ver 3.1 (2001)) because its extent of occurrence is less than 5,000 km 2, all individuals appear likely to be in fewer than five locations, and there is continuing decline in the extent and quality of its habitat (Fahr, 2008n; IUCN, 2009). Assessment History Global 2008: EN B1ab(iii) ver 3.1 (2001) (Fahr, 2008n; IUCN, 2009). 2004: EN B1ab(iii) ver 3.1 (2001) (Fahr, 2004c; IUCN, 2004). abundance, natural history, and threats to this species. GENERAL DISTRIBUTION: Rhinolophus ziama is a West African bat, known from two localities in the Guinea Highlands (c. 600 m asl) of southeast Guinea (Ziama Forest) and north-west Liberia (Wonegizi Mountains) (Fahr et al., 2002). Surrounding areas have been surveyed but the species has not been found elsewhere. Native: Guinea; Liberia; Sierra Leone (Decher et al., 2010; Norris et al., 2015: 83).. POPULATION: Structure and Density:- Rhinolophus ziama is known only from four specimens (Fahr, 2008n; IUCN, 2009). Trend:- 2008: Decreasing (Fahr, 2008n; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Guinea, Liberia. Regional None known. MAJOR THREATS: Rhinolophus ziama is threatened by deforestation of its habitat, largely through logging and mining operations, and conversion of land to agricultural use. It is also considered possible that the species could be threatened by overharvesting for subsistence food in the future (Fahr, 2008n; IUCN, 2009). CONSERVATION ACTIONS: Fahr (2008n) [in IUCN (2009)] reported that it is not known if the species is present in any protected areas. There is a need to protect suitable areas of forest habitat for this species, and to initiate appropriate bat conservation awareness programmes among local people. In addition, further studies are needed into the distribution, Figure 116. Distribution of Rhinolophus ziama Family RHINONYCTERIDAE J. E. Gray, 1866 *1866. Rhinonycteridae Gray, Proc. zool. Soc. Lond., 81. 1866. Rhinonycterina Gray, Proc. zool. Soc. Lond., 1866, I (vi): 81. Publication date: May 1866. - Comments: Proposed as tribe (?). Type genus: Rhinonicteris J. Gray, 1866, and originally included the genus Rhinonicteris J. Gray, 1847. Retained as subtribe by Koopman (1994: 68) (see Jackson and Groves, 2015: 251). *2009. Triaenopini Benda and Vallo, Folia Zool., 58 (Mon. 1): 2, 33 [Goto Description]. Comments: Type genus: Traenops Dobson, 1871. Proposed as tribe and originally 354 ISSN 1990-6471 ? included the genera Triaenops Dobson, 1871 and Paratriaenops Benda and Vallo, 2009. Synonymized with Rhinonycteridae by Foley et al. (2014: 319) (see Jackson and Groves, 2015: 251). Rhinonicteridae (Alternate Spelling) TAXONOMY: Based on the phylogram, the timetree, and a unique retrotransposon insertion, Foley et al. (2014: 313) elevated the subtribe Rhinonycterina Gray, 1866 to family level: Rhinonycteridae. They also found (p. 318) that the monophyletic grouping of this family is supported by a unique shared indel in the THY gene fragment. The validity of this family was also confirmed by Amador et al. (2016: 3, 6). Single known subfamily - Rhinonycterinae Currently (Simmons and Cirranello, 2020) recognized extant genera of Rhinonycteridae: Cloeotis Thomas, 1901; Paratriaenops Benda and Vallo, 2009; Rhinonicteris Gray, 1847, Triaenops Dobson, 1871. Additionally, there are extict genera: †Archerops Hand and Kirsch, 2003; †Brachipposideros Sigé, 1968; †Brevipalatus Hand and Archer, 2005, COMMON NAMES: English: Trident bats (Armstrong et al., 2016: 116). SIMILAR SPECIES: Key characters for the genera: Cloeotis – Head and body length is 33 – 50 mm, tail length is 22 – 33 mm (tail completely enclosed by interfermoral membrane and is longer than the femur), and forearm length is 30 – 39 mm. Posterior component of noseleaf with three tall tapering projections. Two upper premolars on each side, M3 not reduced (four ridges) (Nowak, 1994: 118, Happold, 2013al: 357). Paratriaenops – Head and body length is 35 – 62 mm, tail length is 20 – 34 mm (tail completely enclosed by interfemoral membrane), and the forearm length is about 49 – 61 mm. Posterior component of noseleaf with three tall tapering projections. Two upper premolars on each side, M3 not greatly reduced (three ridges). Rhinonicteris – Head and body length is 45 – 53 mm, tail length is 24 – 28 mm, forearm length is 47 – 50 mm. The complex upper portion of the noseleaf surmounts the nostrils like a crown. The crown-like portion of the leaf is extensively honeycombed and sculptured (Nowak, 1994: 116). Triaenops – Head and body length is 35 – 62 mm, tail length is 20 – 34 mm (tail completely enclosed by interfemoral membrane), and forearm length is 45 – 61 mm. Posterior component of noseleaf with three tall tapering projections as in Cloeotis. Cranially, Triaenops differs from genera in its raised and inflated rostrum and relatively much larger cochleae but resembles Rhinonycters in the structure of the anterorbital region and zygoma and in the curiously thickened premaxillae, two upper premolars on each side, M3 not greatly reduced (three ridges) (Nowak, 1994: 115, Happold, 2013al: 357). MOLECULAR BIOLOGY: The family's karyotype 2n value varies between 36 and 40 (Sotero-Caio et al., 2017: 5). †Genus Brachipposideros Sigé, 1968 *1968. Brachipposideros Sigé, Paleovert., 1 (3): 83. †Genus Brevipalatus Hand and Archer, 2005 *2005. Brevipalatus Hand and Archer, Palaeontology, 48 (2): 372. Genus Cloeotis Thomas, 1901 *1901. Clœotis Thomas, Ann. Mag. nat. Hist., ser. 7, 8 (43): 28. Publication date: 1 July 1901 [Goto Description]. - Comments: Type species: Clœotis percivali Thomas, 1901. Etymology: From the Greek "κλοτός", meaning collar and "ούς" or "ώτός", meaning ear, African Chiroptera Report 2020 2016. ? 355 referring to Thomas' statement that "the whole ear is very like a man's 'stand-up' collar with angles in front rounded off." (see Palmer, 1904: 191). Alternatively "cloeotis" might simply mean short-eared. Cleotis: Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 42. Publication date: 9 May 2016. Cloeotis: (Current Spelling) TAXONOMY: Reviewed by Hill (1982b). Currently (Simmons and Cirranello, 2020) recognized species of the genus Cloeotis: percivali Thomas 1901. COMMON NAMES: Czech: jihoafričtí cíponosi. English: Short-eared Trident Bats, Short-eared Bats, African Trident Bats, African Trident-nosed Bats, Trident-nosed Bats. French: Cléothes. German: KleinohrDreizackblattnase. Italian: Cloeòtidi. Cloeotis percivali Thomas, 1901 *1901. Clœotis Percivali Thomas, Ann. Mag. nat. Hist., ser. 7, 8 (43): 28. Publication date: 1 July 1901. Type locality: Kenya: Coast province: N of Mombassa: Takaungu [03 42 S 39 51 E, 20 m] [Goto Description]. Holotype: BMNH 1901.5.1.11: ♂. Collection date: 15 February 1901. Presented/Donated by: Arthur Blayney Percival. - Etymology: In honour of Arthur Blayney Percival, game ranger in the former British East Africa, and collector of the type specimen (see Lanza et al., 2015: 133). (Current Combination) 1917. Clœotis percivali australis Roberts, Ann. Transv. Mus., 5: 264-5. Publication date: 16 May 1917. Type locality: South Africa: W Transvaal province: Rustenburg district: Mooimeisiesfontein [25 01 S 27 37 E] [Goto Description]. Holotype: TM 1670: ad ♀, skin and skull. Collected by: W. Powell; collection date: 28 May 1915. Paratype: TM 1669: ad ♀, skin and skull. Collected by: W. Powell; collection date: 26 May 1915. Roberts (1917b: 265) called this a "Metatype". Paratype: TM 1671: ad ♀, skin and skull. Collected by: W. Powell; collection date: 27 May 1915. Roberts (1917b: 265) called this a "Metatype". Paratype: TM 1672: ad ♀, skin and skull. Collected by: W. Powell; collection date: 28 May 1915. Roberts (1917b: 265) called this a "Metatype". Paratype: TM 1673: ad ♀, skin and skull. Collected by: W. Powell; collection date: 28 May 1915. Roberts (1917b: 265) called this a "Metatype". Paratype: TM 1674: ad ♂, skin and skull. Collected by: W. Powell; collection date: 26 May 1915. Roberts (1917b: 265) called this a "Metatype". Paratype: TM 1675: ad ♂, skin and skull. Collected by: W. Powell; collection date: 26 May 1915. Roberts (1917b: 265) called this a "Metatype". Paratype: TM 1676: ad ♂, skin and skull. Collected by: W. Powell; collection date: 26 May 1915. Roberts (1917b: 265) called this a "Metatype". - Comments: Considered a valid subspecies by Taylor (1998: 41). Monadjem et al. (2010d: 147) mentioned TM 1690 as type specimen, but Roberts (1917b: 265) indicated TM 1670, which is part of the eight percivali specimens collected by W. Powell at Mooimeisjesfontein and mentioned by the author (TM 1669 - 1676). 2019. Cloeotis percivalli: Waghiiwimbom, Bakwo Fils, Atagana, Tsague Kenfack and Tamesse, Afr. J. Ecol., "6". Publication date: 23 October 2019. (Lapsus) ? Cloeotis percivali australis: ? Cloeotis percivali percivali: (Name Combination) ? Cloeotis percivali: (Current Spelling) 356 ISSN 1990-6471 TAXONOMY: MAJOR THREATS: There have been seemingly stochastic extinctions of colonies, though it is not known if the animals simply move to new locations. Roost disturbance seems to be an important factor, but there might be others, and there has been speculation that DDT might be a cause of population disappearances (Mickleburgh et al., 2008do; IUCN, 2009; Monadjem et al., 2017bc). Figure 117. Cloeotis percivali (TM 46644) from Miggies gat, Mpumalanga, South Africa. Meester et al. (1986: 43 - 44) recognized two subspecies: percivali and australis. COMMON NAMES: Afrikaans: Drietand-bladneusvlermuis, Drietandblaarneusvlermuis. Chinese: 珀 氏 三 叉 蝠 . Czech: cíponos jihoafrický. English: Percival's Short-eared Trident Bat, Percival's Trident Bat, Short-eared Trident Bat, African Trident Bat, East African Trident Bat. French: Cléothe de Percival, Asellia d'Afrique orientale, Phyllorhine à petites oreilles, Phyllorhine de Percival. German: Percivals Kleinohr-Dreizackblattnase. Italian: Cloeòtide di Pèrcival. Kiluba (DRC): Kasusu. Portuguese: Morcego tridentado. SiSwati: Lilulwane. CONSERVATION STATUS: Global Justification Although poorly known and with some recorded population declines, it is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008do; IUCN, 2009; Monadjem et al., 2017bc). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bc). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008do; IUCN, 2009). 2004: VU A2bc+A3bc, C1 ver 3.1 (2001) (Mickleburgh et al., 2004db; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional South Africa:- 2016: E C2a(i) ver 3.1 (2001) (Balona et al., 2016). 2004: CR A2a ver 3.1 (2001) (Friedmann and Daly, 2004). 1996: Indeterminate (Smithers, 1986). Swaziland:- 2003: DD ver 3.1 (2001) (Monadjem et al., 2003). CONSERVATION ACTIONS: Mickleburgh et al. (2008do) [in IUCN (2009)] and Monadjem et al. (2017bc) report that in South Africa, KwaZulu-Natal province they are protected from human disturbance. It might occur in some protected areas. Protection of maternity/breeding and roosting sites from disturbance is important. GENERAL DISTRIBUTION: Largely confined to southern Africa, with records from South Africa (KwaZulu-Natal, Limpopo, and Mpumalanga), Swaziland, south-east Botswana, southern Zambia, Zimbabwe (the core of the distribution), and extralimital records from southern Democratic Republic of Congo, Malawi, northwestern Mozambique, and coastal Kenya (Taylor, 2000). Reported from Mafia Island Tanzania, but no vouchers were collected (Kock and Stanley, 2009; O'Brien, 2011: 287). For southern Africa (south of 8°S), CooperBohannon et al. (2016: Table S2) calculated a potential distribution area of 955,464 km 2. In the RSA, its distribution is best predicted by geology (Babiker Salata, 2012: 50). Native: Botswana (Smithers, 1971, Tiunov, 1986: 536); Cameroon (Waghiiwimbom et al., 2019b: 6), Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 536); Kenya; Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 536); South Africa (Seamark, 2005c: 3; Monadjem et al., 2010d: 536; Balona, 2016: 5); Swaziland (Monadjem et al., 2005; Monadjem et al., 2010d: 536); Tanzania; Zambia (Ansell, 1969; Ansell, 1986; Monadjem et al., 2010d: 536); Zimbabwe (Monadjem et al., 2010d: 536). FUNCTIONAL MORPHOLOGY: Davies et al. (2013b: 6) found that this species' cochlea contains a highly modified basal turn compared to the other species they studied, and these modifications are consistent with the tonotopic organisation of the cochlea in which the basal area corresponds to the highest frequencies used by this bat. African Chiroptera Report 2020 ECHOLOCATION: For an individual from Sengwa in Zimbabwe, Jacobs (1996) and Fenton and Bell (1981) reported a maximum frequency of 212 kHz, the highest frequency pure tone documented from the natural world (Bell and Fenton, 1984 in Thiagavel et al., 2017: 1). Monadjem et al. (2007a), using an Anabat detector reported a maximum frequency of 103.4 (± 0.74) kHz for individuals from various localities in Swaziland. While Monadjem et al. (2010d: 57) report a maximum frequency of 103.2 (± 0.7) using an Anabat detector. Taylor (1999b) using a Pettersson D980 detector reported a maximum frequency of 104.2 (± 0.4) kHz for C. percivali from Jozini in South Africa. Taylor (2000) reported a constant frequency of 104 kHz (fundamental) and 208 kHz (first harmonic). Fenton (2004b: 3) indicates that the calls are dominated by sounds higher than 200 kHz. Monadjem et al. (2010d: 144) report HD-CF calls with high peak frequency (207.8 ± 3 kHz, n = 6) and intermediate duration (4.6 ± 1.2 msec, n = 6), with a second harmonic, where the fundamental harmonic is at around 104 kHz. Monadjem et al. (2007a) also reported sexual dimorphism in the call with males calling at a higher frequency than females (males 103.7 ± 0.71 kHz, females 102.9 ± 0.52 kHz). In Swaziland, Monadjem et al. (2017c: 179) recorded the following parameters: Fmin: 99.2 ± 1.02 (98.1 - 100.1) kHz, Fknee: 103.1 ± 0.65 (102.6 - 103.8) kHz, Fc: 102.6 ± 0.51 (102.2 - 103.2) kHz, duration: 2.3 ± 0.04 (2.2 - 2.3) msec. Three bats (one unknown sex, one female and one male) from the Fikirini caves in Kenya called with Fmax respectively: 217.1, 225.3, and 219.6 kHz and a duration of 3.9, 4.0 and 4.0 msec (Webala et al., 2019b: 15). MOLECULAR BIOLOGY: DNA - Eick et al. (2005) and Matthee et al. (2006). Karyotype - 2n = 40 (Rautenbach et al., 1993). Protein / allozyme - Unknown. HABITAT: Its elevational range is from sea level to 1,000 m (Mickleburgh et al., 2008do). ROOST: Caves and mine tunnels for day roosting (Taylor, 2000). DIET: It feeds exclusively on moths (Taylor, 2000). From Zambia, Black (1979: 54) reported the occurrence of the following prey taxa in the 357 stomachs of 55 bats: Lepidoptera (moths): 100, Coleoptera (beetles): 7.3, Diptera (flies): 5.5, Unidentified insects: 3.6, Hemiptera (bugs): 1.8, and Isoptera (termites): 1.8. Only 7 of the 55 bats had eaten prey other than moths. For South Africa, Jacobs (2000: 201) mentioned the following prey proportions: Lepidoptera (98.7 ± 3.2), Coleoptera (0.3 ± 1.2), Diptera (0.2 ± 0.7), and unknown (0.8 ± 3.1). PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- Large fluctuations in population numbers are known, and it is prone to local extinctions. It is never found in very large colonies, with the largest known colony (of about 300 individuals) being reported from KwaZuluNatal, South Africa, on the southern section of its range. However, the current estimated population size of the KwaZulu-Natal colony is only 50 animals. There are no population estimates from elsewhere in its range, although the largest population might be in Zimbabwe (Mickleburgh et al., 2008do; IUCN, 2009; Monadjem et al., 2017bc). Trend:- 2016: Unknown (Monadjem et al., 2017bc). 2008: Unknown (Mickleburgh et al., 2008do; IUCN, 2009). 2004: Unknown (Mickleburgh et al., 2004db; IUCN, 2004). PARASITES: Streblidae: Raymondia alulata Speiser, 1908 recorded at Kanye, Botswana (Haeselbarth et al., 1966: 102; Shapiro et al., 2016: 253). Raymondia hardyi Fiedler, 1954 from South Africa (Shapiro et al., 2016: 254). Raymondia huberi huberi Frauenfeld, 1855 (Shapiro et al., 2016: 254). Raymondia seminuda Jobling,1954 from southern Botswana (Haeselbarth et al., 1966: 103; Shapiro et al., 2016: 255). Raymondia waterstoni Jobling, 1931 (Haeselbarth et al., 1966: 104). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Botswana, Congo (Democratic Republic of the), Eswatini, Kenya, South Africa, Zambia, Zimbabwe. 358 ISSN 1990-6471 Figure 118. Distribution of Cloeotis percivali Genus Paratriaenops Benda and Vallo, 2009 *2009. Paratriaenops Benda and Vallo, Folia Zool., 58 (1): 1, 31 [Goto Description]. - Comments: Type species: Triaenops furcula Trouessart, 1906. - Etymology: The name refers to close similarity of Paratriaenops with the genus Triaenops; Greek prefix para - means besides, next to. Masculinum. TAXONOMY: Differs primarily from Triaenops Dobson, 1871 and Cloeotis Thomas, 1901 in the shape and morphology of the noseleaf, and lacking of lateral supplementary leaflets (Benda and Vallo, 2009). Based on a considerable genetic distance (21.6 26.2 % in 731 bp sequence of cyt b) and substantial morphological differences from the continental forms of Triaenops as well as from Malagasy T. menamena, Benda and Vallo (2009: 1) propose generic status for the group of Malagasy species, T. furculus, T. auritus, and T. pauliani. They also separate Paratriaenops and Triaenops from the other hipposiderid bats in the tribus Triaenopini. Ramasindrazana et al. (2013: 436) support the recognition of Paratriaenops as a separate genus based on an additional character: the inversed sexual dimorphism, where females are larger than males. Currently (Simmons and Cirranello, 2020) recognized species of the genus Paratriaenops: auritus (G. Grandidier, 1912); furculus (Trouessart, 1907); pauliani (Goodman and Ranivo, 2008). COMMON NAMES: Czech: malí cíponosi. Dreizack-Blattnasen. German: Madagaskar- Paratriaenops auritus (G. Grandidier, 1912) *1912. Triænops aurita G. Grandidier, Bull. Mus. natn. Hist. nat., Paris, sér. 1, 18 (1): 8, fext-fig. Publication date: 25 January 1912. Type locality: Madagascar: extreme N Madagascar: near Diégo-Suarez [=Antsiranana] [ca. 12 16 S 49 17 E] [Goto Description]. Holotype: MCZ 45080: ♂, skin and skull. Collected by: Dr. Mazières. Body mummified; see Peterson (1987: 81). on label: "envoi du Dr. Mazières, Diégo Suarez - Madag. 1910": see Ranivo and Goodman (2006: 974), Helgen and McFadden (2001: 143). 2009. Paratriaenops auritus: Benda and Vallo, Folia Zool., 58: 33. (Name Combination, Current Combination) ? Triaenops auritus: (Current Spelling) TAXONOMY: Until recently, only known from the holotype (see Peterson et al., 1995: 81), but additional specimens were reported by Ranivo and Goodman (2006: 977). Included in furculus (Hayman and Hill, 1971; Koopman, 1993a: 175, 1994), but Peterson et al., 1995: 81), Russ et al. African Chiroptera Report 2020 (2001), Hutson et al. (2001: 18), Ranivo and Goodman (2006: 974) consider it a valid species. Benda and Vallo (2009) placed it in the genus Paratriaenops. COMMON NAMES: Czech: cíponos zlatý. English: Grandidier's trident bat. French: Triaenops à grandes oreilles. German: Grandidiers DreizahnBlattnase. CONSERVATION STATUS: Global Justification This species is listed as Vulnerable (VU B1ab(iii) ver 3.1 (2001)) as it has what appears to be restricted geographic range of less than 20,000 km2 in northwestern Madagascar. The habitat is severely fragmented, and there is a continuing decline in the area and quality of habitat mainly due to agricultural activities. Further survey work is needed to help better ascertain the limits of the range of this species (Andriafidison et al., 2008n; IUCN, 2009; Monadjem et al., 2017a). Assessment History Global 2016: VU B1ab(iii) ver 3.1 (2001) (Monadjem et al., 2017a). 2008: VU B1ab(iii) ver 3.1 (2001) [assessed as Triaenops auritus] (Andriafidison et al., 2008n; IUCN, 2009). 2000: DD ver 2.3 (1994) [assessed as Triaenops auritus] (IUCN, 2000). Regional None known. MAJOR THREATS: As a forest dependent species any decline in extent or quality of the remaining forest poses a threat to this species. In northern Madagascar, forest is lost because of expanding agriculture. There is also the threat of disturbance to cave roosting sites (Andriafidison et al., 2008n; IUCN, 2009; Monadjem et al., 2017a). CONSERVATION ACTIONS: Andriafidison et al. (2008n) [in IUCN (2009)] and Monadjem et al. (2017a) report that this species is known to occur in three protected areas: Réserve Spéciale d’Ankarana, Réserve Spéciale d’Analamerana and Daraina forest (Ranivo and Goodman, 2006; Russell et al., 2007). Conservation should be focused at the known cave roosts to secure the existing populations and closely monitor seasonal and annual population changes. GENERAL DISTRIBUTION: Paratriaenops auritus is endemic to the island of Madagascar where it has a restricted range in the north-western tip of the island, ranging between 50 359 m and 160 m above sea-level (Ranivo and Goodman, 2006; Robinson et al., 2006; Russell et al., 2007). The Andrafiamena Mountains likely form the southern limit of this species (S. M. Goodman pers. comm.). Native: Madagascar (Ranivo and Goodman, 2006: 963; Robinson et al., 2006; Russell et al., 2007). BIOGEOGRAPHY: See Russell et al. (2007). DETAILED MORPHOLOGY: Baculum: The only specimen examined by Rakotondramanana and Goodman (2017: 56) had an elongated baculum with a pointed tip and expanded rounded basal lobe; length: 1.78 mm, width: 0.49 mm. ECHOLOCATION: Kofoky et al. (2009: 380) reported the calls from 10 individuals, which consisted of a distinctive narrowband CF component that ended with short FM sweeps with a maximum energy of 100.3 kHz. Ramasindrazana et al. (2013: 434) mention the following values for 5 ♂♂ specimens: Resting frequency: 107.8 ± 1.51 (106.9 - 109.6) kHz, Fmax: 108.8 ± 1.48 (106.5 - 110.2) kHz, Fmin: 106.5 ± 1.59 (104.8 - 108.2) kHz, duration: 11.7 ± 3.65 (7.6 17.5) msec, interpulse interval: 28.9 ± 8.02 (19.5 38.6) msec, and for 2 ♀♀: Resting frequency: 95.6 (95.1, 96.1) kHz, Fmax: 96.4 (95.6, 97.2) kHz, Fmin: 93.9 (93.8, 94.1) kHz, duration: 13.7 (13.5, 14.0) msec, interpulse interval: 25.5 (25.4, 25.7) msec. MOLECULAR BIOLOGY: DNA - Russell et al. (2007). Karyotype - Unknown Protein / allozyme - Unknown HABITAT: Dammhahn and Goodman (2013: 108) mention that the bat's foraging habitat is the lower portion of forest. POPULATION: Structure and Density:- This species is thought to be relatively abundant within a localized area in the northern part of Madagascar. The largest roost recorded was 2,000 individuals in Réserve Spéciale d’Ankarana (S. G. Cardiff pers. obs. in Andriafidison et al. (2008n) and Monadjem et al. (2017a)) and up to 1,000 individuals were recorded from a mine tunnel approximately 5 km from Andavokoera (J. Ranivo pers. comm. in in Andriafidison et al. (2008n) and Monadjem et al. (2017a). 360 ISSN 1990-6471 Trend:- 2016: Unknown (Monadjem et al., 2017a). 2008: Unknown (Andriafidison et al., 2008n; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Figure 119. Distribution of Paratriaenops auritus Paratriaenops furculus (Trouessart, 1907) *1907. Triaenops furcula Trouessart, Bull. Mus. natn. Hist. nat., Paris, sér. 1, 12 (7): 446 (for 1906). Publication date: 26 February 1907. Type locality: Madagascar: Tuléar province: 20 km S Tuléar, St. Augustine Bay: Sarodrana cave [23 33 S 43 45 E]. Holotype: MNHN ZM-MO-1912-40a (186): ♂, alcoholic (skull missing). Collected by: Guillaume Grandidier; collection date: 19 May 1898. - Comments: Issue 12 (7) was for a meeting on 27 November 1906, but was only distributed around 26 February 1907 (see Bull. Vol 13 (2). Trouessart (1907: 446) mentions Triaenops in name for the new taxon, but refers to the other taxa as Triænops). 2009. Paratriaenops furculus: Benda and Vallo, Folia Zool., 58: 33. (Name Combination, Current Combination) ? Triaenops furculus: (Current Spelling) TAXONOMY: Hayman and Hill (1971: 30) and Koopman (1993a: 175) included aurita, but see Peterson et al. (1995), Russ et al. (2001), Hutson et al. (2001: 18), Simmons (2005: 378), Ranivo and Goodman (2006: 963) who consider it a valid species. Benda and Vallo (2009) placed it into the genus Paratriaenops. COMMON NAMES: Czech: cíponos malý. English: Trouessart's Trident Bat, Trouessart's Triden Bat, Madagascar Leaf-nosed Bat. French: Triaenops malgache, Triaenops de Madagascar, Triaenops de Trouessart. German: Trouessarts DreizahnBlattnase. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Sabatier and Legendre (1985: 23) and Gunnell et al. (2014: 3) refer to fossils from the Anjohibe, Tsimanampetsotsa, and Andrahomana caves on Madagascar. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution. There is no evidence that this species is declining fast enough to place it in a category of higher threat although its roosting colonies should be monitored especially in areas undergoing habitat change (Andriafidison et al., 2008o; IUCN, 2009; Monadjem et al., 2017by). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017by). 2008: LC ver 3.1 (2001) [assessed as Triaenops furculus] (Andriafidison et al., 2008o; IUCN, 2009). 1996: VU A2c ver 2.3 (1994) (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There may be possible threats from disturbance at the cave sites. This species is tolerant to some African Chiroptera Report 2020 degree of forest degradation, but appears to require forested areas to survive (Goodman et al., 2005a) and may therefore be one of the few Malagasy bat species really susceptible to deforestation and forest fragmentation (Andriafidison et al., 2008o; IUCN, 2009; Monadjem et al., 2017by). CONSERVATION ACTIONS: Andriafidison et al. (2008o) [in IUCN (2009)] and Monadjem et al. (2017by) report that this species is present in Parc National du Tsingy de Bemaraha, Parc National de Namoroka, Parc National Kirindy-Mite and Parc National Tsimanampetsotsa (Goodman et al., 2005a). Roosting colonies need to be the focus of further conservation and research. GENERAL DISTRIBUTION: Paratriaenops furculus is found on Madagascar where it is restricted to lowland (an elevation span of 30 m to 200 m above sea level) areas in the west and south-west (Goodman et al., 2005a; Ranivo and Goodman, 2006: 963). It is also found on Cosmoledo and Aldabra Atolls in the outer Seychelles (von Brandis, 2004: 135; Hutson, 2004a). Native: Madagascar (Goodman et al., 2005a; Ranivo and Goodman, 2006: 963); Seychelles (Hutson, 2004a). BIOGEOGRAPHY: See Russell et al. (2007). DETAILED MORPHOLOGY: Baculum: The penis bone is elongated with a slightly pointed tip and triangular basal lobe; length: 1.66 ± 0.154 (1.48 - 1.93) mm, width: 0.37 ± 0.105 (0.22 - 0.53) mm. ECHOLOCATION: Kofoky et al. (2009: 380) reported the calls from 18 individuals, the pulses were of longer duration than Triaenops menamena and consisted of a narrowband CF component that T. menamena, began and ended with short FM sweeps with a high duty cycle (52.5%) and with up to two harmonics, with the second harmonic containing the most energy at about 103.4 kHz. The frequency of maximum amplitude (FMA) was reported as 100.2 ± 0.2 kHz (n = 18) by Bambini et al. (2011: 84). Ramasindrazana et al. (2013: 434) mention for 6 ♂♂ specimens the following values: Resting frequency: 113.1 ± 1.80 (110.6 - 115.1) kHz, Fmax: 114.8 ± 1.90 (111.9 - 116.8) kHz, Fmin: 110.5 ± 2.16 (106.8 - 113.0) kHz, duration: 12.4 ± 2.71 (8.9 - 361 16.9) msec, interpulse interval: 23.4 ± 8.50 (14.4 36.9) msec, and for 10 ♀♀: Resting frequency: 98.2 ± 1.41 (96.8 - 101.2) kHz, Fmax: 99.5 ± 1.21 (98.1 -102.3) kHz, Fmin: 95.7 ± 2.20 (91.4 - 99.4) kHz, duration: 16.0 ± 4.55 (8.1 - 22.5) msec, interpulse interval: 37.7 ± 18.00 (13.9 - 68.5) msec. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003) and Russell et al. (2007). Karyotype - Unknown Protein / allozyme - Unknown HABITAT: Hutson (2004a: 129) reports, that on Madagscar, this bat is essentially a cave inhabitant (often sea caves), but on Aldabra (Seychelles) - where there are no true caves - it might inhabit some vertical solution holes, which are at most about 5m deep, and other small hollows which might provide a limited cave-like environment. ROOST: Recorded in a variety of roosts including caves, shallow sea caves, rock overhangs, under bridges, and large road drainage pipes (Goodman and Ranivo, 2008). DIET: On Madagascar, Rakotoarivelo et al. (2007) found that Paratriaenops furculus mainly consumed Lepidoptera. This was confirmed by Bambini et al. (2011: 81), who also indicated that Coleoptera were also part of the diet. These two order also made up the major volume percentage of preys found in feacal pellets collected by Ramasindrazana et al. (2012: 120-121) in the Tsimanampetsotsa NP: Lepidoptera (65.7 ± 4.8), Coleoptera (27.6 ± 4.6), Psocoptera (1.9 ± 0.5), Diptera (1.9 ± 0.7), Hymenoptera (1.5 ± 0.7), Homoptera/Hemiptera (1.0 ± 0.7), and Neuroptera (0.4 ± 0.3). The percentages of Lepidoptera and Coleoptera were about equal during the dry season (46.0 ± 8.5 versus 47.7 ± 8.3), but differed considerably during the wet season (74.7 ± 3.9 versus 18.3 ± 3.4). PREDATORS: Goodman et al. (2015c: 78) found the remains of one individual in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. POPULATION: Structure and Density:- There are few data available on Paratriaenops furculus, but it appears to be a relatively rare member of the chiropteran assemblage in western Madagascar as determined by mist netting and acoustic sampling (Kofoky et al., 2007; Rakotoarivelo and 362 ISSN 1990-6471 Randrianandrianina, 2007). However, it can occur in large colonies and a roost of over 10,000 individuals was reported from the Sept Lacs in southern Madagascar (Olsson et al., 2006). There is no information on population size of P. furculus from Aldabra, but it appears to be rather rare with only a few specimen records (Hutson, 2004a). Trend:- 2016: Unknown (Monadjem et al., 2017by). 2008: Unknown (Andriafidison et al., 2008o; IUCN, 2009). Paramyxoviridae Wilkinson et al. (2012: 160) tested 1 individuals from the Madagascar using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Mélade et al. (2016b: 4) tested 14 Madagascan specimens, of which one was positive for paramyxoviruses. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. PARASITES: Lagadec et al. (2012: 1696) and Gomard et al. (2016: 5) report the presence of Leptospira sp bacteria in this species. Perkins and Schaer (2016: Suppl.) report the presence of an unidentified haemosporidan. Ramasindrazana et al. (2018: Suppl.) identified the haemosporidian Polychromophilus melanipherus Dionisi, 1899 from this bat species. Ramasindrazana et al. (2016: 7) report one male specimen (out of five examined) to be infected by an unnamed filarial Nematode. VIRUSES: Astroviridae Lebarbenchon et al. (2017: Suppl.) found 11 out of 31 examined bats to test positive for Astroviridae. Figure 120. Distribution of Paratriaenops furculus Paratriaenops pauliani (Goodman and Ranivo, 2008) 1941. [Triaenops] furinea Tate, Am. Mus. Novit., 1140: 3. Type locality: Seychelles: Aldabra. Comments: Goodman and Ranivo (2008: 685) mention that Tate (1941: 3) used the name "Triaenops furinea Trouessart" for the species on Aldabra. To their knowledge Trouessart never named the form occurring on Aldabra; the use of "furinea" is presumably a misreading of the name furcula and is considered a nomen nudum (Hayman and Hill, 1971: 30; Hill, 1982b). *2008. Triaenops pauliani Goodman and Ranivo, Zoosystema, 30 (3): 681, 685, figs 2, 3. Type locality: Seychelles: Aldabra Atoll: Picard Island [c 09 24 S 46 12 E] [Goto Description]. Holotype: BMNH 1913.2.18.1: ad ♂, skull and alcoholic. Collected by: J.C.F. Fryer. Based on the itinerary of the Percy Sladen Trust Expedition to the Seychelles, Fryer was in the western Seychelles (Astove, Cosmoledo, Assumption, and Aldabra) from 13 September1908 to 24 January 1909 (Gardiner et al.,1910), see Goodman and Ranivo (2008). Paratype: BMNH 1913.2.18.2: ad ♀, skull and alcoholic. Collected by: J.C.F. Fryer. Paratype: BMNH 1978.185: ad ♀, alcoholic (skull not removed). Collected by: J.J. Whitelaw; collection date: 04 May 1977. Picard Island, Aldabra Station [09 24 04.4 S 46 12 21.4 E, 5 m]. Paratype: CUMZ E5609.A: ad ♀. Collected by: J.C.F. Fryer. Picard Island, between 13 September 1908 and 24 January 1909. Paratype: CUMZ E5609.B: ad ♂. Collected by: J.C.F. Fryer. Picard Island, between 13 September 1908 and 24 January 1909. Paratype: FMNH 185795: sad ♂, skull and alcoholic. Collected by: U. Samedi; collection date: 09 April 2004. Picard Island, Aldabra Station [09 24 04.4 S 46 12 21.4 E, 5 m]. - Etymology: The name pauliani is a patronym after the late Prof. Renaud Paulian, who conducted extensive research in the western Indian Ocean islands and was African Chiroptera Report 2020 2009. 363 responsible for a major synthesis of the bio-geography of Madagascar and neighbouring islands (Paulian 1961, see Goodman and Ranivo (2008: 685). Paratriaenops pauliani: Benda and Vallo, Folia Zool., 58: 33. (Name Combination, Current Combination) TAXONOMY: Benda and Vallo (2009) placed it into the genus Paratriaenops. COMMON NAMES: Czech: cíponos aldabránský. English: Paulian's Trident Bat. French: Triaenops de Paulian. German: Aldabra- Dreizahnblattnase. CONSERVATION STATUS: Global Justification This species is known from two different islands in the extreme western Seychelles, which include the Aldabra and Cosmoledo Atolls. These two atolls are separate by over 100 km of seas and the occurrence of this species on the latter atoll has been questioned (Goodman and Ranivo, 2008). As nothing is known about this species' ecology and most records (observations and specimens) are of animals in human structures on Aldabra Atoll, further field data is needed to properly assess this taxon (Goodman, 2017b). Assessment History Global 2016: DD ver. 3.1 (2001) (Goodman, 2017b). Regional None known. are distinctly arc-shaped, while in members of the P. auritus/furculus group the external lancets are distinctly spear-shaped and the difference in length of the external and internal lancets is less marked (Goodman and Ranivo, 2008). In overall external measurements, P. pauliani is smaller than P. auritus/furculus group. GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: The cranial measurements are similar to those of P. auritus and P. furculus and fall within the range of size variation of these latter two taxa. There is an elongation of the anterior portion of the upper toothrow, with the I1-M3 measurements falling outside the range for P. furculus and P. auritus (see Goodman and Ranivo, 2008). POPULATION: Structure and Density:- Population size and trends are not known, but based on very irregular records on Aldabra Atoll (von Brandis, 2004; Hutson, 2004a), presumably it is not very common (Goodman, 2017b). Trend:- 2016: Unknown (Goodman, 2017b). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Seychelles. MAJOR THREATS: The main threats to this species are not known (Goodman, 2017b). CONSERVATION ACTIONS: Goodman (2017b) report that the Aldabra Atoll has the status of a UNESCO World Heritage site since late 1982 and as such, its terrestrial habitats are well protected. GENERAL DISTRIBUTION: Is endemic to the Aldabra Atoll, specifically Picard Island (Goodman and Ranivo, 2008). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: The noseleafs of P. pauliani are shorter than those in individuals of P. auritus or P. furculus, and the external lancets are shorter than the internal lancet, and outer margins of the external lancets Figure 121. Distribution of Paratriaenops pauliani 364 ISSN 1990-6471 Genus Triaenops Dobson, 1871 *1871. Triaenops Dobson, J. Asiatic Soc. Bengal, 40 (2): 455, pl. 28. Publication date: 28 December 1871. - Comments: Type species: Triænops persicus Dobson, 1871. Etymology: From the Greek "τρίανα" (trìaina), meaning trident and "ώφ", meaning face (or "ὄφις" (ópsis), meaning "aspect, appearance"), referring to the posterior part of the noseleaf, which terminates in three pointed projections (see Palmer, 1904: 687); Lanza et al., 2015: 155). (Current Combination) 1995. Triaenopus: Yoshiyuki, Bull. Nat. Sci. Mus., Tokyo, Ser. A (Zool.), 21 (2): 119. Publication date: 22 June 1995. (Lapsus) 2002. Trienops: Paniutina, Plecotus et al., pars spec.: 36. (Lapsus) ? Triaenops sp.: TAXONOMY: Hill (1982b) and Garbutt (1999) suggested that T. rufus and T. persicus be synonymized. This is not supported by Russell et al. (2007). The Madagascan species are reviewed by Ranivo and Goodman (2006). Currently (Simmons and Cirranello, 2020) recognized species of the genus Triaenops: afer Peters, 1877; menamena Goodman and Ranivo, 2009, parvus Benda and Vallo, 2009 - Yemen, persicus Dobson, 1871 - Yemen, Oman, United Arab Emirates, Iran, Pakistan; rufus A. MilneEdwards. And the extinct †goodmani Samonds, 2007. COMMON NAMES: Czech: velcí cíponosi. English: Greater Trident Bats, Trident Bats, Trident Leaf-nosed Bats, Triple nose-leaf bats. French: Triaenops. German: Dreizahn-Blattnasen. Italian: Triènopi. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Samonds (2007: 55 - 57) found Triaenops sp. Within the deposits (NCC-1 locality) dating estimated as 69,600 to 86,800 years old within the Anjohibe cave, Madagascar. GENERAL DISTRIBUTION: Is broadly distributed in Africa, portions of the Middle East, and islands in the western Indian Ocean. BIOGEOGRAPHY: Russell et al. (2007)'s data reject a single-dispersal hypothesis (single origin hypothesis), which predicts that linages endemic to Madagascar form an exclusive clade. Russell et al. (2007) found that Malagasy Triaenops are paraphyletic with respect to the mainland African species T. persicus, indicating that the distribution of Malagasy and African Triaenops involved at least two independent dispersal events. Russell et al. (2008a) estimate that at 2.25 Mya Triaenops still formed a single ancestral population between T. rufus/P. auritus, while the timing of the later T. rufus dispersal to Madagascar is less clear (about 148 kya and 660 kya). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia, Kenya, Madagascar, Somalia, Tanzania. †Triaenops goodmani Samonds, 2007 *2007. Triaenops goodmani Samonds, Acta Chiropt., 9 (1): 46, fig 3 A-B. Type locality: Madagascar: Anjohibe cave [Goto Description]. Holotype: UADBA 9010:. Partial left dentary: see Samonds (2007: 46). - Etymology: Specific name for Dr. Steven M. Goodman, in recognition of his significant contributions to the field of modern Malagasy bat research (see Samonds, 2007). (Current Combination) COMMON NAMES: English: Goodman's Trident Bat (Armstrong et al., 2016: 116). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Samonds (2007: 46 - 49) found samples in deposits (OLD SE) dated at 10,000 yrs old or younger within the Anjohibe cave, Madagascar. Time frame: Pleistocene (Brown et al., 2019: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. African Chiroptera Report 2020 365 Triaenops afer Peters, 1877 *1877. Triænops afer Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 913 (for 1876). Publication date: 1877. Type locality: Kenya: Mombassa [04 03 S 39 40 E]. Holotype: ZMB 5074: ♂, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt. See Turni and Kock (2008: 40). (Current Combination) 1968. Triaenops persicus majusculus Aellen and Brosset, Rev. suisse Zool., 75 (14): 450. Type locality: Congo: Loudima: Grotte de Doumboula [04 15 S 13 00 E] [Goto Description]. Holotype: MNHN ZM-MO-1968-412: ad ♂, alcoholic (skull not removed). Collected by: Jean-Paul Adam; collection date: 19 June 1964. 18 paratypes. 2017. Tr[iaenops] affer: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 4. Publication date: 6 April 2017. (Lapsus) 2017. Triaenops affer: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 5. Publication date: 6 April 2017. (Lapsus) ? Triaenops afer: (Current Spelling) ? Triaenops persicus afer: (Name Combination) ? Triaenops persicus: TAXONOMY: Benda and Vallo (2009: 26) found differences in the structure of the skull, baculum and Cyt. b of Triaenops persicus occurring in the Middle East and those in East Africa. COMMON NAMES: Chinese: 非 洲 三 叉 蝠 . Czech: cíponos africký, vrápenec cíponosý. English: African Trident Bat, Peter's Trident Bat. German: Afrikanische Dreizahn-Blattnase. Italian: Triènope àfro. Ukrainian: Трилистоніс африканський [= Trylystonis afrykans'kyy] CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Monadjem and Shapiro, 2017a). Assessment History Global 2016: LC ver. 3.1 (2001) (Monadjem and Shapiro, 2017a). Regional None known. MAJOR THREATS: Monadjem and Shapiro (2017a) reports that there appear to be no major threats to this widespread species as a whole. The species is locally threatened in parts of its range by disturbance of roost sites and mining activities. Populations have been recorded roosting in caves and mines (Taylor, 2005). CONSERVATION ACTIONS: Monadjem and Shapiro (2017a) reports that there appear to be no direct conservation measures in place. It occurs in several protected areas in Mozambique (Monadjem et al., 2010c) and Tanzania. Monadjem and Shapiro (2017a recommends further taxonomic studies are needed for specimens of Triaenops from the Albertine Rift. GENERAL DISTRIBUTION: Native: Ethiopia (Kruskop et al., 2016: 58); Mozambique (Monadjem et al., 2010c: 381). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Lucati and López-Baucells (2016: Suppl.) refers to Howell (1980), who mentioned two piebald (allwhite fur/skin patches, eyes always normally coloured) T. persicus specimens from a mine in Tanzania. DETAILED MORPHOLOGY: Brain: Adult hippocampal neurogenesis was studied by Chawana R. et al. (2016: 1551) [as Triaenops persicus]. ECHOLOCATION: From Mozambique, Monadjem et al. (2010c: 381) reported that the echolocation calls are sexually dimorphic, peak frequencies of males ranged between 71 - 75 kHz (ANABAT) and those of females, between 82 - 85 kHz (ANABAT). The calls of nine Kenyan bats were reported by Webala et al. (2019b: 15) with Fmax: 88.2 - 91.2 kHz (5 ??), 82.5 - 83.9 kHz (2 FF) and 75.2 - 75.7 kHz (2 MM). Three calls of a third male had a Fmax of 30.1 kHz and a duration of 8.4 - 12.2 msec. 366 ISSN 1990-6471 POPULATION: Structure and Density:- Monadjem and Shapiro (2017a) report that some colonies of this bat can be quite large, with up to half a million animals recorded in the Koalin Mines of Tanzania. In peripheral parts of the range, such as Zimbabwe, colonies are often much smaller. Trend:- 2016: Unknown (Monadjem and Shapiro, 2017a). REPRODUCTION AND ONTOGENY: Martin and Bernard (2000: 32) indicate that the corpus luteum is only present for the first part of the pregnancy after which complete regression occurs. PARASITES: BACTERIA Bartonellae Bartonella - Kosoy et al. (2010: 1877) and Bai and Kosoy (2012: 58) reported a prelevence of Bartonella spp. cultered from blood samples of 7/8 (87.5 %) T. persicus specimens from Kenya. Seven Bartonella spp. gltA sequences were identified, and formed a monophyletic geogroup (Kosoy et al., 2010: 1878). See also Kosoy (2010: 719) for further information. HAEMOSPORIDA Garnham (1973: 237) refers to Adam and Landau for a record of Polychromophilus ? murinus in a "Triaenops persicus" from Africa. VIRUSES: Astroviridae These viruses were found by Waruhiu et al. (2017) in their country-wide survey of Kenyan bats. Hoarau et al. (2018: 2) found them in 35 of the 51 Mozambican bats they tested. Coronaviridae Alphacoronavirus: Of the 30 T. afer specimens Tao et al. (2017: 3) tested in Kenya, eight (26.7 %) were positive for CoV (subgenus Setracovirus; see Markotter et al., 2020: 6). Joffrin et al. (2020: 5) reported Alphacoronaviruses from bats from Mozambique and Kenya. Paramyxoviridae Mortlock et al. (2015: 1841) reported that 12 out of 16 examined T. afer specimens from Kenya tested positive for Paramyxovirus sequences. Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 12 Kenyan "T. persicus" specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Central African Republic, Congo, Ethiopia, Kenya, Malawi, Mozambique, Somalia, Tanzania, Uganda, Zambia, Zimbabwe. SIPHONAPTERA Ischnopsyllidae: Araeopsylla scitula Rothschild, 1909 was reported by Beaucournu and Kock (1996) from "Triaenops persicus" bat from Kenya. DIPTERA: Streblidae: Raymondia aspera Maa, 1968 from Mozambique (Shapiro et al., 2016: 254). Raymondia huberi huberi Frauenfeld, 1855 (Shapiro et al., 2016: 254). Raymondia seminuda Jobling, 1954 (Shapiro et al., 2016: 255). Raymondia waterstoni Jobling, 1931 (Shapiro et al., 2016: 256). ACARI: Trombiculidae: Stekolnikov (2018a: 50, 173) found Whartonia oweni Vercammen-Grandjean and Brennan, 1957 and Microtrombicula tanzaniae Goff, 1982 on "Triaenops persicus afer". Figure 122. Distribution of Triaenops afer Triaenops menamena Goodman and Ranivo, 2009 1881. Triænops Humbloti A. Milne-Edwards. Type locality: : "Africa": Aden (Yemen) or Somalia.. Holotype: MNHN ZM-MO-1985-836 (184): ♂, alcoholic (skull missing). Collected by: Léon Humblot; collection date: 1880; original number: 184 or 184A. See Peterson et al. (1995: 82). The type series (184 and 184A) contains 3 MM + 3 FF) African Chiroptera Report 2020 367 (Peterson et al., 1995). Rode (1941: 239) refers to the holotype (184) and six paratypes (184a). Paratype: MNHN 184A [a]: ♀. Collected by: Léon Humblot; collection date: 1880; original number: 184 or 184A. See Peterson et al. (1995: 82). Paratype: MNHN 184A [b]: ♀. Collected by: Léon Humblot; collection date: 1880; original number: 184 or 184A. See Peterson et al. (1995: 82). Paratype: MNHN 184A [c]: ♀. Collected by: Léon Humblot; collection date: 1880; original number: 184 or 184A. See Peterson et al. (1995: 82). Paratype: MNHN ZM-MO-1985-838: ♂. Collected by: Léon Humblot; collection date: 1880; original number: 184 or 184A. See Peterson et al. (1995: 82). Paratype: MNHN ZM-MO-1985-839: ♂. Collected by: Léon Humblot; collection date: 1880; original number: 184 or 184A. See Peterson et al. (1995: 82). - Comments: See Goodman and Ranivo (2009) for futher discussion on the type locality. According to Ranivo and Goodman (2006: 974) the skull of the holotype has been mixed up with the skulls of the paratypes (MNHN 1985-838 and 1985-839). T. humbloti is retained as a synonym for both T. persicus and T. menamena due to the fact that purely taxonomically it is a synonym of T. persicus, but most of the specimens from Madagascar were erroneously assigned this name. In general, the authorship for the Madagascan use would be assigned to a subsequent author or to the same author in a later publication, but in this case, the author already made the error in the original publication. 1881. Triænops rufus A. Milne-Edwards, C. R. séances Acad. Sci., Paris, 91: 1035. Publication date: 1881. Type locality: "Africa": Aden (Yemen) or Somalia [Goto Description]. Holotype: MNHN ZM-MO-1997-1854 (185): ♀, skull and alcoholic. Collected by: Léon Humblot; collection date: 1880. Presented/Donated by: ?: Collector Unknown. See Peterson et al. (1995: 82). Ranivo and Goodman (2006: 974) indicate that the skull is still present, but in very bad shape. Specimen labled "Holotype no. 185" (see Goodman and Ranivo, 2009). Paratype: MNHN ZM-MO-1997-1855 (185A): ♀, alcoholic (skull not removed). Collected by: Léon Humblot; collection date: 1880. Presented/Donated by: ?: Collector Unknown. See Peterson et al. (1995). Paratype: MNHN ZM-MO-1997-1856 (185A): ♀, alcoholic (skull missing). Collected by: Léon Humblot; collection date: 1880. Presented/Donated by: ?: Collector Unknown. See Peterson et al. (1995). Paratype: MNHN ZM-MO-1997-1857 (185A): ♀, alcoholic (skull missing). Collected by: Léon Humblot; collection date: 1880. Presented/Donated by: ?: Collector Unknown. See Peterson et al. (1995). - Comments: (see Goodman and Ranivo, 2009). *2009. Triaenops menamena Goodman and Ranivo, Mammalia, 73 (1): 48, 54, fig 1. Publication date: 31 March 2009. Type locality: Madagascar: Mahajanga province: Réserve Naturelle Intégrale de Namoroka, 4 km SSW Namoroka village: Grotte d'Ampidiranamhaja [16 25.957 S 45 17.180 E, 120 m] [Goto Description]. Holotype: FMNH 178563: ♂, skull and alcoholic. Collected by: Julie Ranivo and Fania H. Ratrimomanarivo; collection date: 25 September 2003; original number: S.M. Goodman 13825. - Etymology: From the Malagasy word "mena" meaning "red", which, in common usage, when repeated twice, means "reddish" (see Goodman and Ranivo, 2009: 54). (Current Combination) ? Triaenops humbloti: ? Triaenops rufus: TAXONOMY: These Madagascan specimens were previously included in persicus by Hayman and Hill (1971: 30), Hill (1982b), Koopman (1993a: 175), probably based on the type specimens, which are now believed to be collected either in Yemen or Somalia (see Goodman and Ranivo, 2009: 52). Based on actual Madagascan material, Peterson et al. (1995: 82), Russ et al. (2001), Hutson et al. (2001: 18), Goodman et al. (2005a: 158), and Simmons (2005: 379) recognized the specimens as representing a separate species. See also the phylogentic analysis by Russell et al. (2007: 847). Goodman and Ranivo (2009) extensively discuss the origin of the type specimens of Triænops Humbloti Milne-Edwards, 1881 and Triænops rufus Milne-Edwards, 1881. Their conclusion is that both forms were not from Madagascar but from Yemen or Somalia. Therefore, from a strict taxonomical view, these two names are synonyms of Triaenops persicus Dobson, 1871. However, we are also retaining these names in the synonymy of Triaenops menamena Goodman and Ranivo, 2009 since these names were always used to identify Madagascan forms, which are currently assigned to T. menamena. 368 ISSN 1990-6471 COMMON NAMES: Czech: cíponos madagaskarský. English: Rufous Trident Bat, Red Madagascan Trident Bat. French: Triaenops roux. German: Rötliche Dreizahn-Blattnase. 2007; Rakotoarivelo et al., 2007; Goodman and Ranivo, 2009). CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution across Madagascar. It can tolerate some degree of habitat transformation, is relatively common, and is not thought to be declining fast enough to place it in a category of higher threat (Andriafidison et al., 2008p; IUCN, 2009). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 57) indicate that the baculum is ‘stick-like’ with a bifid distal tip and straight mid-shaft. Its basal part is triangular with two distinct lateral projections; length: 1.67 ± 0.12 (1.50 - 1.80) mm, width: 0.71 ± 0.07 (0.61 - 0.79) mm. Assessment History Global 2008: LC ver 3.1 (2001) [assessed as Triaenops rufus] (Andriafidison et al., 2008p; IUCN, 2009). 2000: DD ver 2.3 (1994) [assessed as Triaenops rufus] (IUCN, 2000). Regional None known. MAJOR THREATS: There are no significant threats to this species even though it is occasionally hunted for food (Goodman, 2006) and is often associated with forests that are subject to ongoing degradation (Russell et al., 2007). CONSERVATION ACTIONS: Andriafidison et al. (2008p) [in IUCN (2009)] report that this species is known to occur in many protected areas: Réserve Spéciale d’Ankarana, Réserve Spéciale d’Analamerana, Parc National du Tsingy de Bemaraha, Parc National de Namoroka, Parc National de Tsimanampetsotsa, Parc National d’Ankarafantsika, Parc National d’Isalo, Réserve Spéciale d’Ambohitantely, Parc National de Masoala and Réserve Naturelle Intégrale de Tsaratanana (Ranivo and Goodman, 2006; Russ et al., 2003; Russell et al., 2007). GENERAL DISTRIBUTION: Triaenops menamena is endemic to the island of Madagascar. It is restricted to the western dry habitats and extreme northeastern and southeastern Madagascar in humid habitats (Goodman and Ranivo, 2009). BIOGEOGRAPHY: See Russell et al. (2007). ECHOLOCATION: Kofoky et al. (2009: 379) reported the calls from 15 individuals, which produced relatively short CF component followed by brief FM components, the duty cycle was high (25.5 %), with the second harmonic generally contained most energy at about 93.2 kHz, a maximum of three harmonics were also observed. Bambini et al. (2011: 84) reported the frequency of maximum amplitude (FMA) to be 95.8 ± 0.1 kHz (n = 21). Ramasindrazana et al. (2013: 434) reported the following data for 11 ♂♂: Resting frequency: 82.3 ± 1.43 (79.6 - 84.0) kHz, Fmax: 83.6 ± 1.26 (81.4 85.2) kHz, Fmin: 79.5 ± 3.21 (72.2 - 82.3) kHz, duration: 7.5 ± 0.97 (5.5 - 9.4) msec, interpulse interval: 38.3 ± 30.15 (9.8 - 107.4) msec, and for 20 ♀♀: Resting frequency: 93.5 ± 1.47 (90.0 96.8) kHz, Fmax: 94.7 ± 1.49 (91.8 - 98.3) kHz, Fmin: 91.3 ± 2.00 (87.6 - 95.4) kHz; duration: 7.8 ± 1.54 (5.1 - 10.6) msec, interpulse interval: 35.6 ± 24.75 (14.9 - 127.4) msec. MOLECULAR BIOLOGY: DNA - Russell et al. (2007). Karyotype - Unknown Protein / allozyme - Unknown HABITAT: There are no significant threats to this species even though it is occasionally hunted for food (Goodman, 2006) and is often associated with forests that are subject to ongoing degradation (Russell et al., 2007). Papers prior to 2009 refer to T. rufus; see taxonomy section above for more details. Dammhahn and Goodman (2013: 108) mention that this species' foraging habitat is the lower portion of forest. Ravelomanantsoa et al. (2019: 111) captured this bat near a marshy area near Ambereny. Native: Madagascar (Peterson et al., 1995; Russ et al., 2003; Goodman et al., 2005a; Simmons, 2005; Ranivo and Goodman, 2006; Russell et al., ROOST: Wilkinson et al. (2012: 160) indicate that caves are the typical roosts for this species on Madagascar. African Chiroptera Report 2020 369 DIET: On Madagascar Rakotoarivelo et al. (2007) found T. menamena (reported as T. rufus) mainly consumed Lepidoptera. Bambini et al. (2011: 87) reported that T. menamena showed a high preference for Hemiptera, and less for Lepidoptera. a third form having sequences closely related to L. borgpetersenii. The numbers reported by Ramasindrazana et al. (2012: 120-121) from the Tsimanampetsotsa NP confirmed that Lepidoptera formed the major food source of this bat with 67.2 ± 4.5 volume percentage. The Coleoptera formed the second food source with 26.9 ± 4.1. The other insect orders contributed only small amounts: Hymenoptera (3.3 ± 1.0), Psocoptera (0.9 ± 0.2), Neuroptera (0.6 ± 0.2), Diptera (0.4 ± 0.2), Homoptera/Hemiptera (0.4 ± 1.9), Blattaria (0.1 ± 0.0); and Aranea (0.1 ± 0.0). During the dry season, however, Coleoptera contributed the most to the bats diet: 69.0 ± 5.1, whereas Lepidoptera counted for 18.7 ± 4.1 volume percentage. During the wet seaon, the Lepidoptera dominated the diet with 90.5 ± 2.1 volume percent versus 6.7 ± 2.0 for the Coleoptera. Paramyxoviridae Wilkinson et al. (2012: 160) tested 10 individuals from the Madagascar using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 2 positive results for viral nucleic acids. Wilkinson et al. (2014) tested 30 individuals from Madagascar with a Respiro-, Morbilli- and Henipavirus specific RTPCR assay, 19 samples tested positive for paramyxovirus RNA. Rakotondramanana et al. (2015: 77) present the following volume percentages of prey types from the faeces of 16 bats: Coleoptera: 11.6 ± 19.46; Hymenoptera: 11.7 ± 24.20; Lepidoptera: 64.1 ± 32.64; Trichoptera: 0.6 ± 2.25; Isoptera: 11.5 ± 26.32; Homoptera: 0.5 ± 1.55; Arachnida: 0.06 ± 0.25. In frequency percentages, the values are: Coleoptera: 35.0 ± 43.51; Hymenoptera: 42.5 ± 40.58; Lepidoptera: 88.8 ± 30.96; Trichoptera: 3.8 ± 15.00; Isoptera: 17.5 ± 37.86; Homoptera: 3.75 ± 10.88; Arachnida: 1.2 ± 5.00. VIRUSES: Astroviridae Eight out of 13 specimens tested by Lebarbenchon et al. (2017: Suppl.) were positive for Astroviridae. In their extensive study, Mélade et al. (2016b: 3) found this species to be the most infected by paramyxoviruses of all Madagascan bats (21 out of 42 specimens = 50 %). Rhabdoviridae Two out of 11 bats tested by Mélade et al. (2016a: 6) showed a seroreaction against Lagos bat lyssavirus. UTILISATION: Goodman (2006) describes the hunting method used by local inhabitants. See also Jenkins et al. (2007b), Goodman et al. (2008d) and Jenkins and Racey (2008), who refer to T. rufus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. POPULATION: Structure and Density:- The species is thought to be locally common in many parts of western Madagascar. A single roost in deciduous forest near the Onilahy River in southern Madagascar contained over 40,000 individuals (Olsson et al., 2006). Trend:- 2008: Unknown (Andriafidison et al., 2008p; IUCN, 2009). PARASITES: BACTERIA Lagadec et al. (2012: 1696) reported that 8 out of 10 bats they examined, tested positive for Leptospira sp. bacteria, and Gomard et al. (2016: 5) found 27 out of 42 bats (64.3 %) to be positive for this bacterium. Dietrich et al. (2018a: 3) identified these further as L. interrogans, L. sp. and Figure 123. Distribution of Triaenops menamena 370 ISSN 1990-6471 Superfamily RHINOPOMATOIDEA Dobson, 1872 *1872. Rhinopomatoidea Dobson. - Comments: The name of the Superfamily is based on Rhinopomatidae Dobson, 1872. (Current Combination) TAXONOMY: Based on morphological data, the family Rhinopomatidae was placed in the superfamily Rhinopomatoidea by Simmons (1998) and Simmons and Geisler (1998), but more recent molecular studies have contradicted many groupings based on morphological data and, pending resolution of the controversies, for this reason Simmons (2005) did not recognize any chiropteran superfamilies. Family RHINOPOMATIDAE Dobson, 1872 1838. Rhinopomina Bonaparte, Syn. Vert. Syst., in Nuovi Ann. Sci. Nat., Bologna, 2: 111 (sep. p. 7). - Comments: . *1872. Rhinopomatidae Dobson, J. Asiatic Soc. Bengal, 41 (3): 221. (Current Combination) 1907. Rhinopomidae Miller, Bull. U.S. natl. Mus., 57: 80. Publication date: 29 June 1907. Comments: Type genus Rhinopoma Geoffroy St.-Hilaire, 1818. TAXONOMY: This is a monotypic family, with all extant species belonging to the genus Rhinopoma. Koopman (1993a) and Simmons (2005) assigned the name Rhinopomatidae to Bonaparte, 1838 Syn. Vert. Syst. In Nuovi Ann. Sci. Nat., Bologna, 2: 111; but see Corbet and Hill (1992: 81) and Van Cakenberghe and De Vree (1994). Miller (1907) proposed the name Rhinopomidae and this was adopted by several authors: however, the original name is most commonly used (Aulagnier, 2013d:409). The family was reviewed by Hill (1977a) and more recently by Van Cakenberghe and De Vree (1994) and Hulva et al. (2007b). Currently recognized genera of the family Rhinopomatidae: †Qarunycteris Gunnell, Simmons and Seiffet, 2008; Rhinopoma E. Geoffroy, 1818. COMMON NAMES: Czech: víkonosovití. Dutch: Langstaartvleermuizen, Muisstaartvleermuizen. English: Mouse-tailed bats. French: Rhinopomes, Chauves-souris à queue de souris, Rhinopomatidés. German: Mausschwanzfledermäuse, MausschwanzFledermäuse. Italian: Rinopormàtidi, Pipistrèlli a códa di tòpo. Norwegian: Langhalete flaggermus, klaffneser. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The oldest remains of extant rhinopomatids were collected from Egyptian tombs, which have been inhabited by these bats for ca. 3000 years (Aulagnier, 2013d: 410). The family-level stem and crown ages were calculated by Shi and Rabosky (2015: 1537) to be respectively 51.9 and 26.9 mya. GENERAL DISTRIBUTION: The family is distributed from Africa (north of the Equator) to southern Asia through Arabia, the Middle East and India, mainly in semi-arid and arid habitats (Aulagnier, 2013d: 409). The three species of rhinopomatids occuring in Africa occur in the tropics; two also extend northwards into temperate habitats. In Africa, one species is found in semi-desert vegetation zones, and the remaining two are found in both desert and semi-desert vegetation zones. Two species have also been found in montane habitats (but not above ca. 1200 m) (Aulagnier, 2013d:409-410). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: These bats of primitive structure are characterized by a tail that is exceptionally long and has at least half of its length projecting beyond the narrow interfemoral membrane, slit-like valvular nostrils that can be closed, and a small triangular nose leaf surmounting a hog-shaped muzzle (that gave the name to the genus) (Aulagnier, 2013d:409). Rhinopomatids are small to medium-sized with moderately long pelage that is generally more or less sepia grey (but varying in colour and tone across the geographic range), and usually paler ventrally. The face, rump and posterior portion of the abdomen are naked and medium brown. A thickened narial pad is present on the end of the muzzle, surmounted by a distinct ridge-like dermal outgrowth (a rudimentary noseleaf). The ears are more or less triangular and joined across the African Chiroptera Report 2020 forehead by a connecting band of skin; each ear has a simple but moderately long and erect tragus. Eyes are relatively large (Aulagnier, 2013d:409). Wings rather narrow; second finger with two bony phalanges (like all other African families); all of the fingers are shorter, relative to the length of the forearm, than in any other bats. The tip of the wing can be shortened by folding to facilitate cursorial locomotion, but not to the same extent as in the Emballonuridae and Molossidae. This is achieved by the terminal phalanx of the third finger bending round, but not fully, upon the ventral surface of the wing (Rosevear, 1965). The hindlimbs are moderately long; the toes (except the hallux) have three phalanges. The tail has sensitive hairs at its distal end (Aulagnier, 2013d: 409). DENTAL FORMULA: 1113/ 2123 = 28. The upper incisor (I2) is very small, barely emerging from the gum. The canines are conical and simple. The first and second upper molars are lacking distinct hypocones. GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: The skull is relatively short and broad; there are no postorbital processes. Separate nasal swellings are present on each side of the rostrum. The sagittal crest is low and sharp. The auditory bullae are rather large. The premaxillae are separate from each other and from the adjacent part of the skull. 371 SEXUAL DIMORPHISM: Sexes similar in colour, and males are often slightly larger than females (Aulagnier, 2013d: 409). ECHOLOCATION: When echolocating, ultrasound pulses are emitted through the mouth; the echolocation calls are loud with little modulation of frequency when the bats are foraging, and become more frequencymodulated when the bats are approaching prey or when they are emerging from their day-roosts in groups (Schmidt and Joermann, 1986, Aulagnier, 2013d:410). MOLECULAR BIOLOGY: Within the family, Sotero-Caio et al. (2017: 5) indicated that the 2n value varies from 36 to 42. ROOST: Rhinopomatids roost in dry caves, crevices, various underground structures and buildings including tombs and pyramids; their day- roosts are impregnated with a characteristic smell (Aulagnier, 2013d: 409). They sometimes share day-roosts with other species of bats, including other species of rhinopomatids, but the different species do not roost in exactly the same places (Aulagnier, 2013d: 409). Rhinopomatids sometimes roost in clusters numbering up to many thousands of individuals, but most of the groups are smaller (10 - 100 bats) (Aulagnier, 2013d: 410). DETAILED MORPHOLOGY: Rhinopomatids can accumulate fat in the lower abdominal region; this is metabolized when food is scarce (Aulagnier, 2013d: 410). Their water metabolism reflects extreme adaptations to desert habitats; the concentration of urea in the blood is high, and the skin is poorly vascularized so that loss of water through evaporation is minimal, even at high temperatures (Aulagnier, 2013d: 410). MIGRATION: Some populations are suspected to migrate (Aulagnier, 2013d: 410). FUNCTIONAL MORPHOLOGY: Their wings are adapted for fast flight in open areas, gliding or rapid wing-flapping, but are poor fliers and tire quickly (Aulagnier, 2013d: 410). MATING: Little is known of their breeding habits in Africa (Aulagnier, 2013d: 410). ACTIVITY AND BEHAVIOUR: The sexes segregate for at least part of the year (Aulagnier, 2013d: 410). These bats do not hibernate, but they can remain torpid for several days (Aulagnier, 2013d: 410). †Genus Qarunycteris Gunnell, Simons and Seiffert, 2008 *2008. Qarunycteris Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 6. - Comments: Type species - Qarunycteris moerisae. - Etymology: Combination of Qarun for Birket Qarun, a large lake near BQ-2 Quarry with Nykteris, Greek for bat (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). Curently recognized species of the genus Qarunycteris: †moerisae Gunnell, Simons and Seiffert, 2008. 372 ISSN 1990-6471 SIMILAR SPECIES: See Gunnell et al. (2008). GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). †Qarunycteris moerisae Gunnell, Simons and Seiffert, 2008 *2008. Qarunycteris moerisae Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 3, 6, fig. 4F. Type locality: Egypt: Fayum Depression: Quarry BQ-2: Umm Rigl Member of the Birket Qarun Formation. Holotype: CGM 83671:. M2. - Etymology: For Lake Moeris, the ancient name of Birket Qarun (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). Earliest late Eocene (Priabonian - 37.97 - 33.23 mya Brown et al., 2019: Suppl.). SIMILAR SPECIES: See Gunnell et al. (2008). GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). Timeframe: Genus Rhinopoma E. Geoffroy St.-Hilaire, 1818 *1818. Rhinopoma E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 113. Publication date: 1818. - Comments: Type: Vespertilio microphyllus Brünnich, 1782. - Etymology: From the Greek "ρις" (rhís) or "ρινός" (rhinós), meaning "nose or snout", and "πώμα" (pomatòs), meaning "lid, cover", referring to the valvular nostrils, which open through a narrow, transverse slit (see Palmer, 1904: 607; Lanza et al., 2015: 180). (Current Combination) 1821. Rhynopoma Bowdich, Anal. Nat. Class. Mamm, : 30. - Comments: Type: Vespertilio microphyllus Brünnich, 1782. 1854. Rhinopomus: Gervais, Histoire Naturelle des Mammifères, 202. - Comments: Lapsus (Allen, 1939a: 64). (Lapsus) 2017. Rhinopona: Loumassine, Allegrini, Bounaceur, Peyre and Aulagnier, Mammalia, xx (x): xxx. Publication date: 23 February 2017. (Lapsus) TAXONOMY: Revised by Hill (1977a) and Van Cakenberghe and De Vree (1994). Allen (1939a: 64) assigned Rhinopoma to Oken, 1816, Lehrbuch d. Naturgesch., pt. 3, zool., Sect. 2, p. 926, but this name was rendered "Not Available" by Opinion 417 (International Commission on Zoological Nomenclature, 1956). A key of the genus is given in Van Cakenberghe and De Vree (1994: 21). Ethiopian records of Rhinopoma muscatellum Thomas, 1903, mentioned by Koopman (1993a) represent Rhinopoma macinnesi Hayman, 1937 (see Van Cakenberghe and De Vree, 1994: 20), therefore R. muscatellum is no longer included in the list of African species. Currently (Simmons and Cirranello, 2020) recognized species of the genus Rhinopoma: cystops Thomas, 1903; hadramauticum Benda, 2009 – Yemen; hardwickii Gray, 1831; macinnesi Hayman, 1937; microphyllum (Bünnich, 1782); muscatellum Thomas, 1903 – United Arab Emirates, Oman, Yemen, southwestern Iran, southern Afghanistan, western Pakistan, southwestern India (Simmons, 2005: 381). COMMON NAMES: Czech: víkonosi, nosalecové, klaponosové. English: Mouse-tailed bats. French: Rhinopomes. African Chiroptera Report 2020 German: Mausschwanz-Fledermäuse. Rhinopòma. 373 Italian: DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". PARASITES: Streblidae: Brachytarsina alluaudi (Falcoz, 1923) (Haeselbarth et al., 1966: 101). Rhinopoma cystops Thomas, 1903 1816. Rhinopoma brevicaudatum Oken, Lehrbuch Naturgeschichte, Jena, 3 (2): 926. Type locality: Egypt: "Egypt". - Comments: Not available, ICZN Opinion 417, 1956 (Van Cakenberghe and De Vree, 1994: 14). 1866. Rhinopoma longicaudatum Fitzinger, in: Heuglin and Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 54 (1) 10: 547. Type locality: Sudan: Sennaar [15 35 N 33 38 E, 425 m]. - Comments: Nomen nudum: Kock (1969a: 35), Van Cakenberghe and De Vree (1994: 14). 1866. Rhinopoma sennaariense Fitzinger, in: Heuglin and Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 54 (1) 10: 547. Type locality: Sudan: Anglo-Egyptian Sudan, near Roseires: Sennaar and Fazoglo [=Fazughli] [ca. 11 20 N 34 45 E]. - Comments: Type locality restricted to Fazughli by Kock (1969a: 35). Nomen nudum: see Harrison and Bates (1991: 29), Van Cakenberghe and De Vree (1994: 14). Validated by Kock (1969a) according to Qumsiyeh and Jones (1986: 1). 1868. Rhinopoma sennarense Hartmann, Z. Gesel. f. Erdk., 3 (28/29): 44. - Comments: Nomen nudum: Van Cakenberghe and De Vree (1994: 14). 1877. Rhinopoma senaarense, potius senarense: Heuglin, Reise in Nordost Afrika, 2: 24. Publication date: 1877. - Comments: Emendation of sennariense Fitzinger 1866 (see Kock, 1969a: 35); Lapsus: Van Cakenberghe and De Vree (1994: 14). (Lapsus, Emendation) *1903. Rhinopoma cystops Thomas, Ann. Mag. nat. Hist., ser. 7, 11 (65): 496. Publication date: 1 May 1903. Type locality: Egypt: Qena province: Luxor [25 41 N 32 39 E] [Goto Description]. Holotype: BMNH 1902.1.17.2: ad ♀, skull and alcoholic. Collected by: Nathanial Charles Rothschild. Presented/Donated by: Nathanial Charles Rothschild. See Thomas (1903b: 497); Kock (1969a: 35); Qumsiyeh (1985: 20). Paratype: BMNH 1902.1.17.1: ad ♂. Collected by: Nathanial Charles Rothschild. Presented/Donated by: ?: Collector Unknown. - Etymology: From the Greek substantive "κύστις" (kystis) meaning "bladder" and "ὄφις" (ópsis) meaning "eye-ball", referring to the large eyes (see Lanza et al., 2015: 182, Lanza et al., 2015: 182). (Current Combination) 1917. Rhinopoma longicaudum: Geyr von Scheppenburg, J. Ornithol., 65 (3): 283. Publication date: July 1917. (Lapsus) 1954. Vespertilio brevicauda Stresemann, Abh. d. Akad. Wiss., Berlin, 1: 172. Type locality: Egypt: Giza province: Saqqara [29 51 N 31 13 E]. Holotype: ZMB 46211: ♂, skull and alcoholic. Collected by: Friedrich Wilhelm Hemprich and Christian Gottfried Ehrenberg; collection date: April 1821; original number: A.1.6.69. Formerly ZMB 564.3 "Sendung Nr 69-71"; see Kock (1969a: 35); Turni and Kock (2008). Holotype: ZMB 564.c: ♂. Collected by: Friedrich Wilhelm Hemprich and Christian Gottfried Ehrenberg; collection date: April 1821. Presented/Donated by: ?: Collector Unknown. - Comments: Label name from Hemprich, without nomenclatural status, see Kock (1969a: 35), Van Cakenberghe and De Vree (1994: 14). 1954. Vespertilio ferox Stresemann, Abh. d. Akad. Wiss., Berlin, 1: 172. Type locality: Egypt: Giza province: Saqqara [29 51 N 31 13 E]. Holotype: ZMB 46210: ♂, skull and alcoholic. Collected by: Friedrich Wilhelm Hemprich and Christian Gottfried Ehrenberg; collection date: April 1821; original number: A.1.6.68. Formerly ZMB 564.2 "Sendung nr 67-68"; see Kock (1969a: 35), Turni and Kock (2008). Holotype: ZMB 564.b: ♂. Collected by: Friedrich Wilhelm Hemprich and Christian Gottfried Ehrenberg; collection date: April 1821. Presented/Donated by: ?: Collector Unknown. - Comments: Label name from Hemprich, without nomenclatural status, see Kock (1969a: 35), Van Cakenberghe and De Vree (1994: 14). 1956. Rhinopoma hardwichei cystops: Möhres and Kulzer, Verh. D. zool. Ges., 19: 60. Comments: Lapsus (see Van Cakenberghe and De Vree, 1994: 15). (Lapsus) 374 ISSN 1990-6471 1961. 1969. 1971. 1975. 1985. 2015. 2015. ? ? ? ? ? ? ? Rhinopoma lardwickei cystops: Madkour, Bull. Zool. Soc. Egypt, 16: 50. - Comments: Lapsus (see Kock, 1969a: 35; Van Cakenberghe and De Vree, 1994: 15). (Lapsus) Rhinopoma hardwickei sennaariense Kock, Abh. Senckenberg. naturforsch. Ges., 521: 40, 51. Type locality: Sudan: Blue Nile province: Fazogli [=Fazughli] [11 20 N 34 45 E] [Goto Description]. - Comments: Validation of sennaariense Fitzinger, 1866, nomen nudum. Hill (1977a) pointed out that sennaariense Kock, 1969 is a synonym of arabium (see Harrison and Bates, 1991: 30). Rhinopoma hardwickei sennaarinese: Hill and Morris, Bull. Br. Mus. (nat. Hist.) Zool., 21 (2): 30. - Comments: Lapsus (see Van Cakenberghe and De Vree, 1994: 15). (Lapsus) Rhinopoma senaariense: Koopman, Bull. Am. Mus. Nat. Hist., 154 (4): 367. - Comments: Lapsus (see Van Cakenberghe and De Vree, 1994: 15). (Lapsus) R[hinopoma] h[ardwickei] sinaariense: Qumsiyeh, Spec. Publ. Mus. Texas Tech. Univ., 23: 22. - Comments: Lapsus: attributed to Kock (1969a) by Qumsiyeh (1985: 22). (Lapsus) Rhinopoma cytops: Kangoyé, Ouéda, Granjon, Thiombiano, Guenda and Fahr, Biod. J., 6 (2): 613. Publication date: 30 June 2015. (Lapsus) Rhinopoma hardweickei: Mohammed, Endocrinol. Metab. Synd., 4: 1. Publication date: 5 January 2015. - Comments: Rhinopoma hardweickei for R. harwickei [= R. cystops]. (Lapsus) Rhinopoma hardwickei arabium: (Alternate Spelling) Rhinopoma hardwickei cystops: (Name Combination) Rhinopoma hardwickei hardwickei: - Comments: Only for the African specimens. Rhinopoma hardwickei:, . - Comments: Emendation of hardwickii, since the species was named after Maj. Gen. Hardwicke, but according to Hill (pers. comm.) this was unjustified since Gray latinized the name. Only for the African specimens. (Emendation) Rhinopoma hardwickii arabium: (Lapsus) Rhinopoma hardwickii cystops: (Name Combination) Rhinopoma Hardwickii:, . - Comments: Not of Gray (1831), which is restricted to Asia (see Hulva et al., 2007b), but when used for the African specimens. TAXONOMY: See Schlitter and Qumsiyeh (1996, Mammalian Species, 263), Van Cakenberghe and De Vree (1994: 20) as Rhinopoma hardwickii. Hulva et al. (2007b) restrict R. hardwickii to Asia (Iran to Indonesia) and reinstate cystops as the species name for the African and Western Asian representatives. Ahmim and Tahri (2017: 71) indicate that in Algeria, Niger, and Egypt, two forms of "R. Hardwickii" differing in size are known, which casts doubt on the recent taxonomic discrimination of the genus Rhinopoma, implying that besides R. cystops another small sized species might need to be recognized in northern Africa. Benda et al. (2020: 2586) found that Ethiopian bats are very similar in size to the animals from the southern part of the Arabian peninsula, and smaller than those from Somalia, and they suggest that possibly all specimens from the Horn of Africa should be assigned to Rhinopoma cystops arabium rather than to R. c. cystops, although they can not exclude that there might be a mosaic-like border between the two lineages. COMMON NAMES: Arabian: Khafash abu danab saghir, Khaffash. Czech: víkonos africký. Dutch: Kleine klapneusvleermuis. English: Hardwicke's Mouse-tailed Bat, Lesser Mouse-tailed Bat, Small mouse-tailed bat, Lesser Rat-tailed Bat, Egyptian Mouse-tailed bat. French: Petit Rhinopome, Rhinopoma moyen, Rhinopome de Hardwicke, Chauve-souris à queue de souris. German: Hardwickes Mausschwanz-Fledermaus, Kleine Mausschwanzfledermaus. Hebrew: Yaznuv Katan. Italian: Rhinopòma àfro-ovestasiàtico. CONSERVATION STATUS: Global Justification Assessed as Least Concern (LC ver 3.1 (2001)) as it is a widespread and common species with no major threats (Benda, 2017a). Assessment History Global 2016: LC ver 3.1 (2001) (Benda, 2017a). 2008: LC ver 3.1 (2001) [assessed as part of Rhinopoma hardwickii] (Benda et al., 2008e; IUCN, 2009). 2004: LC ver 3.1 (2001) [assessed as part of Rhinopoma hardwickii] (Benda, 2004b; IUCN, 2004). 1996: VU [assessed as R. hadithaensis] (Baillie and Groombridge, 1996); LR/lc (Baillie and Groombridge, 1996). African Chiroptera Report 2020 Regional None known. MAJOR THREATS: Benda (2017a) reports that no specific measures are known or are in place, but occurs in protected areas across the range. A study on the impacts of pesticides is required, especially ways in which the impact might be minimised. Aerial hawking and water stress are considered to be the major risk factors related to climatic change (Sherwin et al., 2012: 174 as Rhinopoma hardwickei). Mansour et al. (2016: 66) examined the occurrence of heavy metals in bats from two caves in the Saqqara region of Egypt and found that cobalt was not detected in any sample analyzed. Cd, Mo, Ni and Pb were found in low concentrations (generally less than 1.0 µg/g tissue). In the livers, moderate concentrations of Ba, Cr, Cu and Mn and very high concentrations of Ca, Fe and Mg were recorded. Liver tissue from males contained much higher levels of Al and Ba than those from females. CONSERVATION ACTIONS: Benda et al. (2008e) [in IUCN (2009)] and Benda (2017a) reported that no specific measures are known or are in place, but presumably occurs in protected areas across the range. A study on the impacts of pesticides is required, especially ways in which the impact might be minimised. GENERAL DISTRIBUTION: Rhinopoma cystops occurs across central and northern Africa; from Morocco to Israel, Palestine, Jordan Iraq and Afghanistan and south to Kenya. Occurs up 1,100 m asl in Morocco and Algeria. The Kenyan material mentioned by Koopman (1993a) belongs to R. macinnesi, see Van Cakenberghe and De Vree (1994: 17 - 18). Native: Afghanistan; Algeria (Horácek et al., 2000: 97); Burkina Faso (Kangoyé et al., 2015a: 613); Cameroon; Chad; Djibouti (Pearch et al., 2001: 392); Egypt (Sinai; first record; see Benda et al., 2008e: 1; Carpenter et al., 2014: 180); Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 147, Benda et al., 2020: 2585); India; Iran, Islamic Republic of; Iraq; Israel; Jordan; Kenya (but see note above); Kuwait; Libyan Arab Jamahiriya; Mali; Mauritania (Brito et al., 2010: 452, as R. hardwickei, Allegrini et al., 2011: 3); Morocco (Benda et al., 2004d; Benda et al., 2010a: 154; El Ibrahimi and Rguibi Idrissi, 2015: 358, as R. hardwickei); Niger; Nigeria; Oman; Saudi Arabia; Somalia; Sudan; 375 Syrian Arab Republic; Tunisia (Horácek et al., 2000: 97; Dalhoumi et al., 2016b: 866); Western Sahara; Yemen (Socotra). DETAILED MORPHOLOGY: Mohammed (2015) extensively discuss pituitary gland of Egyptian "R. hardwickei". the SEXUAL DIMORPHISM: Kangoyé et al. (2015a: 613) indicate that most of the average body measurements are larger in females than in males, but that the average cranial measurements are larger in males. ECHOLOCATION: Frequency of CF-component (second harmonic) during pursuit of insect (Egypt): 36 - 40 kHz; callduration 6 - 10 ms; call-repetition rate ca. 10 call/s. Darting flight is accompanied by shortening the signals to 1 ms or less and increasing the callrepetition rate to ca. 100 calls/s; laboratory tests indicated that Doppler shift compensation is probably not used by this species (Simmons et al., 1984). When three individuals flew together in the laboratory, each used a different CF-frequency suggesting that they regulate the CF-frequency to avoid jamming each other (Habersetzer, 1981). After landing, CF calls of mean duration 74 ms are emitted and these have maximum energy in fundamental harmonic; the purpose of these calls (echolocation and/or communication) is not known (ibid.). The first harmonic frequencies (18 - 20 kHz; Simmons et al., 1984) of calls made in a roost in Morocco were audible to humans (Aulagnier and Destre, 1985). In Jordan, Benda et al. (2010b: 204) reported the following parameters for 59 calls: Start frequency: 36.4 ± 1.5 (33.2 - 39.6) kHz, End frequency: 32.5 ± 1.4 (29.5 - 35.8) kHz, Peak frequency: 34.3 ± 1.2 (32.0 - 37.1) kHz, Call duration: 8.0 ± 1.6 (4.5 12.0) msec, Inter-pulse interval: 116.0 - 29.9 (60.0 - 210.0) msec. From Tunisia, Dalhoumi et al. (2016b: 866) reported an almost constant frequency with three or four harmonics and maximum energy in the second harmonic (at 32 kHz). Moores and Brown (2017: 612) recorded the following parameters in southern Morocco: Fstart: 34.6 ± 0.5 (33.6 - 35.6) kHz, Fmax: 33.6 ± 0.4 (32.8 - 34.4) kHz, Fend: 33.0 ± 0.5 (32.5 - 34.1) kHz, duration: 9.4 ± 0.8 (8.6 - 11.1) msec, interpulse interval: 260 ± 122 (69 - 520) msec. MOLECULAR BIOLOGY: DNA - Giannini and Simmons (2005); Ali (2011). Karyotype - Ray-Chaudhuri et al. (1968) and Rushton (1970: 463) reported 2n = 36, FN = 68, 376 ISSN 1990-6471 BA = 34, a submetacentric X chromosome and an acrocentric Y chromosome for specimens from India, this data now relates to Rhinopoma hardwickii. Qumsiyeh and Baker (1985) reported an identical karyotype for specimens from Palestine, which are now referred to Rhinopoma cystops see Hulva et al. (2007a). Sayed (2011: 659) also reported 2n = 36 and FN = 68 for R. hardwickei specimens from Egypt, with a mediumsized submetacentric X chromosome and an acrocentric Y chromosome. Pairs 13 and 15 show a broad G-negative region around their centromere, and complete heterochromatin in the short and long arm. Pair 5 also shows a broad Gnegative region around its centromere, and the Y chromosome has complete heterochromatin. REPRODUCTION AND ONTOGENY: Gharaibeh (1997: 47) reported two pregnant females captured in Tunisia on 26 May 1996. MATING: Khajuria (1972: 307) report on the mating behaviour of the closely related Indian R. h. hardwickei, but whether this is also applicable for R. cystops is uncertain. PARASITES: TREMATODA Morsy et al. (2018: 319) reported the presence of Urotrema scabridum (Braun, 1900) and Renschetrema indicum Kifune, 1984 in "R. hardwickii" specimens collected in Cairo. Protein / allozyme - Unknown. ROOST: In Israel, Levin et al. (2015: 1, 3) found that R. cystops hibernates during five months in winter in caves that were geothermally heated to a stable temperature of 20 °C, with almost 100 % relative humidity. They also found that the bat's skin temperature was 1 - 3 °C above the average cave temperature, and that during the same period, the outside temperatures fluctuated between 4 and 22 °C. During the hibernation, no period arousal or periods of normothermy were observed. Benda et al. (2014c: 10) observed a colony of about 40 specimens in two ground floor rooms of a ruin of an Italian fortress in the Al Jaghbub oasis (Libya). DIET: Benda et al. (2014c: 14) examined 40 faecal pellets of R. cystops from the Al Jaghbub oasis (Libya). The diet consisted primarily of ants (Hymenoptera: Formicoidea; 81 % of volume) and furthermore of beetles (Coleoptera, mostly of the family Tenebrionidae). ACARI Argasidae: Argas vespertilionis was reported from Libyan bats by Benda et al. (2014c: 15), and two larvae of Argas confusus (Argasidae) on R. cystops specimens in Jordan (Benda et al., 2010b: 212). Trombiculidae: Grandjeana mauritanica Kalúz and Ševcík, 2014 on R. cystops specimens from Mauritania (Kalúz and Ševcík, 2014: 31; Stekolnikov, 2018a: 136). Bendjeddou et al. (2017: 15) reported the following ectoparasites from Algeria: Nycteribia (Listropoda) schmidlii schmidlii Schiner, 1853 (Nycteribiidae) and Argas vespertilionis (Latreille, 1802) (Arachnida). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Burkina Faso, Chad, Djibouti, Egypt, Ethiopia, Kenya, Libya, Mali, Mauritania, Morocco, Niger, Senegal, Somalia, South Sudan, Sudan, Tunisia. POPULATION: Structure and Density:- Appears to be particularly abundant near oases. However, both distribution and abundance are undoubtedly insufficiently investigated because the roosts and suitable habitats are often unreachable. Colonies range in size from a few individuals up to several hundred maximum is about 1000 individuals (Egypt) (Benda, 2017a). This species is assumed stable throughout its distribution range. Population information remains unknown for most of its African distribution. Loumassine et al. (2017c:28) reported on a colony of about 100 animals in an old mine in Algeria. Figure 124. Distribution of Rhinopoma cystops Trend:- 2016: Stable (Benda, 2017a). African Chiroptera Report 2020 377 Rhinopoma macinnesi Hayman, 1937 *1937. R[hinopoma] cystops macinnesi Hayman, in: St. Leger, Ann. Mag. nat. Hist., ser. 10, 19 (113): 530. Publication date: 1 May 1937. Type locality: Kenya: Lake Rudolf, near Central Island: Bat Island [03 27 N 36 04 E, 700 m] [Goto Description]. Holotype: BMNH 1936.11.4.45: ad ♂, skull and alcoholic. Collected by: D.G. MacInnes; collection date: 24 April 1934. - Etymology: In honour of Mr. Donald Gordon MacInnes, eminent English palaeontologist, who worked in eastern Africa and collected of the type specimen (see Lanza et al., 2015: 189). ? Rhinopoma macinnesi: (Name Combination, Current Combination) TAXONOMY: Considered a synonym of hardwickei by Koopman (1993a), but see Van Cakenberghe and De Vree (1994: 20). COMMON NAMES: Chinese: 麦 氏 鼠 尾 蝠 . Czech: víkonos východoafrický. English: MacInnes's Mousetailed Bat. French: Rhinopome de MacInnes. German: MacInnes' Mausschwanz-Fledermaus. Italian: Rinopòma di MacÌnnes. CONSERVATION STATUS: Global Justification Rhinopoma macinnesi is listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Aulagnier, 2008; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Aulagnier, 2008; IUCN, 2009). 2004: VU D1 ver 3.1 (2001) (Aulagnier, 2004a; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). GENERAL DISTRIBUTION: Rhinopoma macinnesi is endemic to East Africa, where it has mostly been recorded from Kenya with additional records from the eastern edge of Uganda, Somalia, Ethiopia and possibly Eritrea (Aulagnier, 2008; IUCN, 2009). Native: Ethiopia; Kenya; Somalia; Uganda Presence uncertain: Eritrea POPULATION: Structure and Density:- Little information is available on the population abundance or size of this species. It is believed to have a small population as several expeditions to the species range have not collected specimens (Aulagnier, 2008; IUCN, 2009). Trend:- 2008: Unknown (Aulagnier, 2008; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Eritrea, Kenya, Somalia. Regional None known. MAJOR THREATS: Rhinopoma macinnesi may be threatened by habitat loss, although this requires confirmation (Aulagnier, 2008; IUCN, 2009). CONSERVATION ACTIONS: Aulagnier (2008) [in IUCN (2009)] reported that there appear to be no direct conservation measures in place. It is possible that the species is present in some protected areas. Further research is needed into the distribution, natural history and possible threats to this species. Figure 125. Distribution of Rhinopoma macinnesi 378 ISSN 1990-6471 Rhinopoma microphyllum (Brünnich, 1782) *1782. Vespertilio Microphyllus Brünnich, Dyrenes Historie, 1: 50, pl. 6; figs. 1 - 4. Type locality: Egypt: Giza province: Giza [30 01 N 31 13 E]. Holotype: MNHN A.121/T196: ♀, alcoholic (skull not removed). Number 196 in Rode (1941: 242), who calls this the holotype. Holotype: ZMUC. Collected by: Hasselquist. See Kock (1969a). Paratype: MNHN A.123/196B: ad, alcoholic (skull not removed). Number 196b in Rode (1941: 242). Paratype: MNHN A.124/196C: ad ♀, alcoholic (skull not removed). Number 196c in Rode (1941: 242). Paratype: MNHN A.125/196D: ad ♂, alcoholic (skull not removed). Number 196d in Rode (1941: 242). Allotype: MNHN A.122/196A: ♂, alcoholic (skull not removed). Number 196a in Rode (1941: 242). - Comments: Corbet and Hill (1992: 82) and Horácek et al. (2000: 96) mention "Arabia and Egypt" as type locality. Type locality fixed by Koopman (1975: 366) but also Aellen (1957), Anderson and de Winton (1902), Kock (1969a) - See Schlitter and Qumsiyeh (1996: 1). - Etymology: From the Greek "μικρόϛ" (mikrós), meaning "small" and "φύλλον" (phýllon), meaning "leaf" (see Schlitter and Qumsiyeh, 1996: 4; Lanza et al., 2015: 193). 1820. Rhinopoma microphylla: Desmarest, Encyclopédie Méthodique, (Zoologie, Mammalogie), 1: 129. Publication date: 1820. (Current Combination, Alternate Spelling) 1859. Rhinopoma Lepsianum Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 222. Publication date: 1859. Type locality: Sudan: Khartoum [15 40 N 32 35 E] [Goto Description]. Lectotype: ZMB 2578: ad ♂, skull and alcoholic. Collected by: Lepsius; collection date: 1844?. Lectotype designated by Kock (1969a: 54, 57); skull extracted in the 1960's by D. Kock (see Turni and Kock (2008: 22). Paralectotype: ZMB A.3243: ad, skull only. Paralectotype designated by Turni and Kock (2008: 22). - Comments: Type locality mentioned as Blue Nile by Kock (1969a: 58); as White Nile by Koopman (1975: 366), Turni and Kock (2008: 22). See Turni and Kock (2008: 22) for a discussion on the fact that Peters possibly used this skull to describe lepsianum). 1877. Rhinopoma cordofanicum Heuglin, Reise in Nordost Afrika, 2: 24. Publication date: 1877. Type locality: Sudan: Jebel Arashkool: W bank of the White Nile [ca. 14 15 N 32 10 E]. Comments: Allen (1939a: 64) mentions Heuglin and Fitzinger, 1866. Sitzb. K. Akad. Wiss. Wien, 54, sec 1, 547 as author. Nomen Nudum. Included in hardwickei by Kock (1969a: 35). 1969. Rhinopoma microphyllum tropicalis Kock, Abh. Senckenberg. naturforsch. Ges., 521: 58. Type locality: Sudan: Kordofan province: 2 km NE Kadugli: Jebel Talao [11 02 N 29 43 E, 550 m] [Goto Description]. Holotype: SMF 33265: sad ♂, skin and skeleton and skull. Collection date: 12 February 1963. See Kock (1969a: 58): [number 33216 possibly???]. Paratype: SMF 33198: ♂, skull and alcoholic. Collected by: Dr. Dieter Kock; collection date: 6 January 1965. Presented/Donated by: ?: Collector Unknown. Locality: Jebel Talao. Paratype: SMF 33199: ♂, skull and alcoholic. Collected by: Dr. Dieter Kock; collection date: 6 January 1965. Presented/Donated by: ?: Collector Unknown. Locality: Jebel Talao. Paratype: SMF 33200: ♂, skull and alcoholic. Collected by: Dr. Dieter Kock; collection date: 6 January 1965. Presented/Donated by: ?: Collector Unknown. Locality: Jebel Talao. Paratype: SMF 33214: ♂, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 22 October 1962. Presented/Donated by: ?: Collector Unknown. Locality: Jebel Talao. Paratype: SMF 33215: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 16 February 1963. Presented/Donated by: ?: Collector Unknown. Locality: Jebel Talao. - Comments: Most likely a synonym of R. m. microphyllum. If a valid subspecies, lepsianum or cordofanicum may have priority (see Schlitter and Qumsiyeh, 1996: 1). 1985. Rhinopoma mircophylum kineari: Bhatnagar and Parveen, Nat. Acad. Sci. Letters, 8 (2): 64. - Comments: Lapsus for microphyllum. (Lapsus) ? Rhinopoma microphyllum microphyllum: (Name Combination) ? Rhinopoma microphyllum: (Name Combination) TAXONOMY: See Schlitter and Qumsiyeh (1996, Mammalian Species, 542). COMMON NAMES: Arabian: Khafash abu danab kabir, Khaffash. Bengali: Indur-lenji Chamchika. Czech: víkonos velký, nosalec malolupenný, klaponos egyptský, víkonos, víkonos egyptský, víkonos asijský. African Chiroptera Report 2020 Dutch: Grote klapneusvleermuis. English: Larger Mouse-tailed Bat, Long-tailed Bat, Greater Mousetailed Bat, Rat-tailed Bat, Brunnich's Mouse-tailed Bat, Larger Rat-tailed Bat, Greater Rat-tailed Bat. French: Grand Rhinopome, Rhinopome microphylle. German: Ägyptische MausschwanzFledermaus. Hebrew: Atalef Yaznuv Gadol. Indonesian: Kelelawar ekor lembing, Kelelawar Ekor-tikus. Italian: Rinopòma microfìllo. CONSERVATION STATUS: Global Justification Rhinopoma microphyllum is listed as Least Concern (LC ver 3.1 (2001)) as it is widespread and common in at least parts of its range, with no known major threats (Aulagnier and Palmeirim, 2008b; IUCN, 2009; Monadjem et al., 2017bp). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bp). 2008: LC ver 3.1 (2001) (Aulagnier and Palmeirim, 2008b; IUCN, 2009). 2004: LC ver 3.1 (2001) (Aulagnier, 2004b; IUCN, 2004). 1996: LR: lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Human disturbance in roost sites and use of pesticides for locusts affect the species but are not considered major threats at present (Aulagnier and Palmeirim, 2008b; IUCN, 2009; Monadjem et al., 2017bp). CONSERVATION ACTIONS: Aulagnier and Palmeirim (2008b) [in IUCN (2009)] and Monadjem et al. (2017bp) reported that no specific measures are known for this species, but presumably it occurs in some protected areas. A study on the impacts of pesticides is required, especially ways in which the impact might be minimised. GENERAL DISTRIBUTION: The distribution range of Rhinopoma microphyllum extends from northern Africa through southwest Asia to Afghanistan, Pakistan and India. In southeast Asia, there are single old records from Thailand and Sumatra. Recorded from sea level in Egypt up to 1,200 m in Morocco (Aulagnier and Palmeirim, 2008b; IUCN, 2009). Native: Afghanistan; Algeria (Loumassine et al., 2017b: 85); Bangladesh; Benin; Burkina Faso (Kangoyé et al., 2015a: 613); Central African Republic; Chad; Côte d'Ivoire; Djibouti (Pearch et al., 2001: 393 ); Egypt; Eritrea; Ethiopia (Benda et al., 2020; 2584); Ghana; Guinea; India; Indonesia; 379 Iran, Islamic Republic of; Iraq; Israel; Jordan; Lebanon; Liberia; Mali; Mauritania (Benda et al., 2004a: 105); Morocco (El Ibrahimi and Rguibi Idrissi, 2015: 358); Myanmar (see Corbet and Hill, 1992); Niger; Nigeria; Oman; Pakistan; Saudi Arabia; Senegal; Sierra Leone; Sudan; Sumatra; Syrian Arab Republic; Togo; Western Sahara; Yemen. Presence uncertain: Libyan Arab Jamahiriya; Thailand; Tunisia. ECHOLOCATION: Search-phase call-shape (locality not given): CF (multiharmonic) (Schmidt and Joermann, 1983). Frequency of CF-component (second harmonic): 27 - 31 kHz (but each individual emits only one frequency); call-duration "long". When flying in a flight-tunnel, this species emits shorter and slightly frequency-modulated calls (as does R. hardwickii). However, if 3 individuals fly together in the flighttunnel, instead of avoiding jamming by changing to three different frequencies (see R. hardwickii), the main response of these bats is to greatly increase the intensity of their calls (ibid.). Cvikel et al. (2014: 2) showed that R. microphyllum doesn't show the classical jamming avoidance response: they altered their signals as if they were responding to a nearby (silent) object. They also found that individuals exhibit sufficient frequency spread to allow individual recognition without any jamming avoidance response. In Jordan, Benda et al. (2010b: 204) recorded the following parameters for 9 calls: Fstart: 29.6 ± 0.3 (28.9 - 30.0) kHz, Fend: 25.7 ± 0.6 (25.2 - 26.7) kHz, Fpeak: 28.0 ± 0.7 (27.0 - 29.0) kHz, Pulse duration: 11.7 ± 0.8 (10.5 - 13.0) msec, and Inter-pulse Interval: 202.3 ± 46.8 (110.0 - 267.0) msec. Benda et al. (2012a: 182) reported the following values from Iran (hand released): F start: 29.8 ± 0.6 (29.0 - 30.4) kHz, Fend: 25.3 ± 0.2 (25.0 - 25.5) kHz, Fpeak: 27.1 ± 0.4 (26.8 - 27.7) kHz, Pulse duration: 7.9 ± 0.3 (7.6 - 8.3) msec, and inter-pulse interval: 114.6 ± 7.9 (106.5 - 127.2) msec. From Israel the following parameters were reported by Hackett et al. (2016: 223): 116 calls: Pulse duration: 8.37 ± 2.51 msec, Fstart: 30.21 ± 1.95 (28.0 - 31.2) kHz, Fend: 28.99 ± 1.72 (27.0 30.2) kHz, Fpeak: 29.42 ± 1.72 (28.0 - 30.2) kHz. From Algeria, Loumassine et al. (2017b: 87) reported on four harmonics, of which the second one had the most energy: Fstart: 31.9 - 37.9 Khz, Fmax: 28.2 - 31.2 kHz, Fend: 23.1 - 25.2 kHz, and pulse duration: 3.2 - 6.1 msec. 380 ISSN 1990-6471 Davies et al. (2013b: Table S8) report a peakfrequency of 28 kHz and a range between 27 and 30 kHz. Boonman et al. (2013b: 6) determined the detection range to be between 2 and 6.5 m at 20 dB and 5.5 and 14 m at 0 dB. Luo et al. (2019a: Supp.) reported the following data (Free flying bats): F peak: 29.42 kHz, Fstart: 30.21 kHz, Fend: 28.99 kHz, Band width: 1.22 kHz, and duration: 8.37 msec. MOLECULAR BIOLOGY: DNA - Hulva et al. (2007a). Karyotype - Handa and Kaur (1979), and Qumsiyeh and Baker (1985) reported 2n = 42, FN = 66, BA = 26, a metacentric X and an acrocentric Y chromosome. Protein / allozyme - Unknown. ROOST: From northern Israel, Levin et al. (2013: 1) report that R. microphyllum forms sexually-segregated colonies composed of 3,000 to 5,000 individuals during the summer (June to September). At the end of August non-reproductive males left their roost and were occasionally observed in the southern female roosts between September and October. Also in Israel, Levin et al. (2015: 1, 3) found that R. microphyllum hibernates during five months in winter in caves that were geothermally heated to a stable temperature of 20 °C, with almost 100 % relative humidity. The bat's skin temperature was 1 - 3 °C above the average cave temperature. During the same period, the outside temperatures fluctuated between 4 and 22 °C. DIET: In July from Israel, more than 50 % of the bats diet comprised queen carpenter ants (Camponotus felah Dalla Torre, 1893), while other insect groups in the faces comprised mainly Coleoptera and Heteroptera (Levin et al., 2012; Prat and Yovel, 2020: 170). This suggests that the gradual increase in the proportion of queen carpenter ants in the diet during summer results from an increase in this insect’s availability, combined with a high preference for this prey item. To search for these queen swarms, the bats travel dozens of kilometers. Benda et al. (2012a: 196) analyses faeces from two localities in Iran, and found in the pellets from one of the localities only remains from ants (Hymenoptera, Formicoidea), whereas the other sample contained Coleoptera (60 % of volume), Heteroptera (30 %) and small nematoceran Diptera (10 %). POPULATION: Structure and Density:- According to Aulagnier and Palmeirim, 2008b) [in IUCN (2009)] and Monadjem et al. (2017bp) , Rhinopoma microphyllum is uncommon in the western part of the Mediterranean range, but populations probably occur in areas not yet surveyed. Colonies of several thousands have been reported in Egypt. Very rare in Jordan, found in the same caves as R. hardwickei but in much lower numbers (Amr, 2000). In Iran the species is common, the most populated cave is in the Mesopotamian plain and has around 20,000 individuals; the total population of Iran is approximately 30,000 and is considered stable (M. Sharifi pers. comm., 2005). In Africa there are large colonies of up to 5,000 individuals in several areas (GMA Africa Workshop, 2004). Trend:- 2016: Stable (Monadjem et al., 2017bp). 2008: Stable (Aulagnier and Palmeirim, 2008b; IUCN, 2009). R. microphyllum populations within the Asia Minor and Levant region are predicted to have a stable population trend, under climate change scenarios (Bilgin et al., 2012: 433). ACTIVITY AND BEHAVIOUR: A sharp increase in body mass was observed from mid-July (Levin et al., 2012). Levin et al. (2012) also found that the fatty acids composition of R. micophyllum body fat reflects the composition of their main prey item-queen carpenter ants. They accumulate significant fat reserves towards the end of summer (Levin et al., 2012). Levin et al. (2012) found white adipose tissue accumulates in pre-hibernating R. micophyllum. Cvikel et al. (2015: 206) used miniature GPS devices that allowed simultaneous ultrasonic recording and found that one of their tracked bats flew continuously for over 5 hours and that another on flew more than 90 km. All of the bats spent a substantial proportion of their time in high conspecific density. They estimated that bats spent at least 41 ± 14 % of their foraging at less than 150 m from a conspecific and even 8 ± 2 % at less than 12 m from a conspecific PARASITES: Hirst (1923: 979) indicates that Kolenati stated that either R. microphyllum or Pteropus ægyptiacus [=Rousettus aegyptiacus] is the host for Ancystropus zelebori Kolenati, 1856 (Acari). In Iran, Benda et al. (2012a: 196) found the following parasites to be present: bat flies Brachytarsina flavipennis Macquart, 1851 and B. African Chiroptera Report 2020 381 alluaudi (Falcoz, 1923) [= diversa (Frauenfeld, 1857)]. They also refer to Maa (1965), who reported the presence of streblid bat flies Ascodipteron namrui Maa, 1964 and A. rhinopomatos Jobling, 1952 from Israel and Egypt, and to Dusbábek (1970), who reported the Macronyssid mites Macronyssus leucipe (Domrow, 1959) and Steatonyssus afer Radovsky and Yunker, 1963 from Afghanistan (but these came from a mixed collection of R. microphyllum and R. hardwickii). Furthermore, the presence of the chigger mites Sasatrombicula rhinopoma (Vercammen-Grandjean, 1963) and S. multisternalae (Vercammen-Grandjean, 1963) was reported from Afghanistan too. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Burkina Faso, Cameroon, Egypt, Ethiopia, Ghana, Mauritania, Morocco, Niger, Nigeria, Senegal, Sudan. Figure 126. Distribution of Rhinopoma microphyllum †Family TANZANYCTERIDIDAE Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 *2003. Tanzanycterididae Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, Pal. Electr., 5(3): 3. (Current Combination) 2013. Tanzanycteridae: Habersetzer, Engels, Gunnell and Simmons, 16th International Bat Research Conference & 43rd North American Symposium on Bat Research. San José, Costa Rica, 11-15 August 2013, 64. (Lapsus) TAXONOMY: Known genera of the family Tanzanycterididae: †Tanzanycteris Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003. †Genus Tanzanycteris Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 *2003. Tanzanycteris Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, Pal. Electr., 5(3): 3. (Current Combination) TAXONOMY: Using micro-CT scans, Habersetzer et al. (2013: 64) examined the inner ear morphology of T. mannardi and found that this was very similar to the highly sophisticated echolocation system found in current days Hipposideros. They suggest that Tanzanycteris might actually be a Hipposiderid rather than a member of a family by its own. Currently recognized species of the genus Tanzanycteris: †mannardi Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003. †Tanzanycteris mannardi Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, 2003 *2003. Tanzanycteris mannardi Gunnell, Jacobs, Herendeen, Head, Kowalski, Msuya, Mizambwa, Harrison, Habersetzer and Storch, Pal. Electr., 5(3): 5. Type locality: Tanzania: Mahenge: approximately 50 km west of Singida [044750.2S 341554.5E]. Holotype: TNM MP-207: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Partial skeleton including skull, axial skeleton anterior to sacrum, partial left and right humeri, and partial 382 ISSN 1990-6471 left radius. - Etymology: For George W. Mannard who explored the Singida Kimberlite Field and made the first excavation at the type locality in 1957. (Current Combination) TAXONOMY: Ravel et al. (2011: 403) indicate that the phylogenetic position of T. mannardi is still uncertain due to its poor state of preservation, but that it shows many similarities with the Hassianycteridae. Phillips (2015: 9) refers to Gunnell et al. (2003), who identified characters that put T. mannardi within the Yinpterochiroptera, more specifically the Rhinolophoidea. These characters include: extremely enlarged cochlea, broadened first rib, and a dorsally flared iliac blade. However, it remains unclear whether Tanzanycteris is directly linked to the Rhinolophoidea or to the Yinpterochiroptera (which also include the Pteropodidae). Phillips (2015) estimates the Hard Minimum Age for this taxon to be 45.0 miilion years ago and the Soft Maximum Age at 58.9 mya. Ravel et al. (2016: 356) place it in the Middle Eocene. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Middle Eocene (Lutetian - 47.8 - 41.3 mya - Brown et al., 2019: Suppl.) African Chiroptera Report 2020 383 SUBORDER VESPERTILIONIFORMI Van Cakenberghe, Kearney and Seamark, 2007 1821. 1855. 1866. 1866. 1979. 1979. 1985. 1995. 1997. 1998. 1998. 1998. INSECTIVORAE Gray, 299. - Comments: Proposed as an order of bats, for the same content as that of MICROCHIROPTERA Dobson 1875. Originally included the families Noctilionidae J. Gray, 1821 and Vespertilionidae J. Gray, 1821. Not Insectivora Bowdich, 1821 (Mammalia, Lipotyphla) (see Jackson and Groves, 2015: 252). Gymnorhina Giebel, Die Säugethiere in Zoologischer, anatomischer und palaeontologischer Beziehung umfassend dargestellt., xii, 926. - Comments: Originally included the genera Furia F. Cuvier, 1828 [= Furipterus Bonaparte, 1837 [1832–1841]]; Nycticejus [sic = Nycticeius] Rafinesque, 1819; Vespertilio Linnaeus, 1758; Thyroptera de Spix, 1823; Dysopes Illiger, 1811 [= Molossus É. Geoffroy, 1805]; Emballonura Kuhl [= Temminck, 1838]; Diclidurus Wied-Neuwied, 1820; Noctilio (Geoff) [= Linnaeus, 1766]; Taphozous É. Geoffroy, 1818; and Mormops [sic = Mormoops] Leach, 1821] (see Jackson and Groves, 2015: 252). Entomophaga A. Murray, The Geographical Distribution of Mammals., xiv. - Comments: Originally included the Istiophora Giebel, 1855 [= Phyllostomidae Gray, 1825] and Gymnorhini Giebel, 1855 [= Yangochiroptera Koopman, 1985; part]. Not Entomophaga Owen, 1839 (Mammalia, Dasyuromorphia [= Marsupialia Illiger, 1811], nor Entomophaga Owen, 1859 (Mammalia, Didelphimorphia) (see Jackson and Groves, 2015: 252). ENTOMOPHAGA: Murray, xiv. - Comments: Not ENTOMOPHAGA Owen 1859, used for didelphoid marsupials. PHYLLOSTOMATIA Van Valen, Evol. Theory, 4 (3): 103, 109. Publication date: July 1979. - Comments: Proposed as infraorder and originally included the superfamilies Rhinopomatoidea Bonaparte, 1838 (containing of the families Rhinopomatidae Bonaparte, 1838, Emballonuridae Gervais, 1855 and Craseonycteridae Hill, 1974); Rhinolophoidea J. Gray, 1825 (containing the families Rhinolophidae J. Gray, 1825, Nycteridae Van der Hoeven, 1855 and Megadermatidae H. Allen, 1864); and Noctilionoidea J. Gray, 1821 (containing the families Noctilionidae J. Gray, 1821, Mormoopidae de Saussure, 1860, Phyllostomatidae Coues and Yarrow, 1875 [= Phyllostomidae J. Gray, 1825] and Desmodontidae Bonaparte, 1845: 5). Jackson and Groves (2015: 252) considered the name as a synonym of the Yangochiroptera Koopman, 1985, but the listing suggests it might rather be a synonym of the Chiroptera. VESPERTILIONIA Van Valen, Evol. Theory, 4 (3): 103, 109. Publication date: July 1979. - Comments: Proposed as infraorder and originally included the superfamily Vespertilionoidea J. Gray, 1821, containing the families Natalidae J. Gray, 1866, Vespertilionidae J. Gray, 1821, Mystacinidae Dobson, 1875 and Molossidae Gervais, 1855 (see Jackson and Groves, 2015: 252). YANGOCHIROPTERA Koopman, Bat Res. News, 25 (3/4): 26. - Comments: Proposed as Infraorder in the suborder Microchiroptera and originally included the superfamilies Phyllostomoidea J. Gray, 1825 and Vespertilionoidea J. Gray, 1821 (see Jackson and Groves (2015: 252). VESPERTILIIFORMES Zagorodniuk, Godovanets, Pokynchereda and Kyseliuk, Rakhiv (Proceed. Intern. Conf. "ACANAP-1995" 18 - 21 October 1995). - Comments: also Zagorodniuk et al. (1995); Zagorodniuk and Pokynchereda (1997). VESPERTILIFORMES Zagorodniuk. Microchiropteraformes Simmons and Geisler, Bull. Am. Mus. Nat. Hist., 235: 136. Comments: Originally included the fossil genus Eppsinycteris Hooker, 1996; the fossil families Palaeochiropterygidae Revilliod, 1917 and Hassianycteridae Habersetzer and Storch, 1987; and the suborder Microchiroptera Dobson, 1875 [= Chiroptera Blumenbach, 1779; part] (see Jackson and Groves, 2015: 253). Microchiropteramorpha Simmons and Geisler, Bull. Am. Mus. Nat. Hist., 235: 135, 136. Comments: Originally included the fossil genus Australonycteris Hand et al., 1994; the fossil families Icaronycteridae Habersetzer and Storch, 1987 and Archaeonycteridae Revilliod, 1917; and the unranked Microchiropteraformes (see Jackson and Groves, 2015: 253). VESPERTILIONIFORMES Zagorodnyuk. - Comments: Zagorodnyuk (1998a, b and c); Zagorodniuk (1999, 2001a and b, 2002, 2003a and b, 2004). Jackson and Groves (2015: 228) assigned this name to Hutcheon and Kirsch (2004b: 44), who included the families 384 ISSN 1990-6471 Phyllostomidae J. Gray, 1825: 242, Mormoopidae de Saussure, 1860, Noctilionidae J. Gray, 1821, Mystacinidae Dobson, 1875, Emballonuridae Gervais, 1855, Vespertilionidae J. Gray, 1821, Miniopteridae Dobson, 1875, Molossidae Gervais, 1855, Natalidae J. Gray, 1866 and Nycteridae Van der Hoeven, 1855. 2001. YINPTEROCHIROPTERA Springer, Teeling, Madsen, Stanhope and De Jong, Proc. Natl. Acad. Sci. USA, 98 (11): 6423. - Comments: In part. 2004. VESPERTILIONIFORMES Hutcheon and Kirsch, J. mamm. Evolut., 11 (1): ???. Comments: . *2007. VESPERTILIONIFORMI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the Infraorder is based on Vespertilio Linnaeus, 1758 (type of VESPERTILIONIDAE Gray, 1821). (Current Combination) 2008. Microcheroptera: Selim, Nahla and Shelfeh, Tishreen Univ. J. Res. Sci. Stud. - Biol. Sci. Ser., 30 (1): 247. (Lapsus) 2014. Vespertilionimorpha Zagorodniuk, Proc. Theriol. School, 12: 8. 2018. Yinpterachiroptera: Skirmuntt and Katzourakis, Virus Research, 270 (197645): 2. Publication date: 1 July 2018. (Lapsus) TAXONOMY: The suborder name follows the typified name used by Zagorodniuk et al. (1995) - Vespertiliiformes, Zagorodniuk (1997) Vespertiliformes, Zagorodniuk and Pokynchereda (1997) Vespertiliiformes, Zagorodniuk (1998a, 1998b and 1998c) - Vespertilioniformes, Zagorodniuk (1999 (1999, 2001a and 2001b, 2002, 2003a and b, 2004) - Vespertilioniformes and Hutcheon and Kirsch (2004b,2006) - Vespertilioniformes, but with the ending changed to follow the format proposed for standardization of nomenclature of higher taxa by Alonso-Zarazaga (2005). '-formes' indicates ordinal level, while ending '-formi' indicates subordinal level. Currently three Infraorders are recognized within the suborder VESPERTILIONIFORMI from Africa: NOCTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007; NYCTERIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007; VESPERTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007. FUNCTIONAL MORPHOLOGY: Rydell et al. (2019: 1, 2) suggested that whitish wings (as found in 17 species of African emballonurid, vespertilionid and molossid bats) reduce the contrast against the sky and may prevent overheating in bats flying during the day. This would also enable the bats the leave their roosts earlier in the evening and returning later in the morning, allowing them to capture crepuscular and diurnal insects. MOLECULAR BIOLOGY: Volleth (2010: 308) found no cytogenetic synapomorphy for Vespertilioniformes [= Vespertilioniformi], which indicates that this group has a monophyletic origin. INFRAORDER NOCTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 *2007. NOCTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the Superfamily is based on Noctilio Linnaeus, 1766. (Current Combination) TAXONOMY: Currently a single superfamily is known from within the Infraorder NOCTILIONIFORMACEI for Africa: NOCTILIONOIDEA Gray 1821. Superfamily NOCTILIONOIDEA Gray, 1821 *? Noctilionoidea - Comments: The name of the Superfamily is based on Noctilio Linnaeus, 1766. (Current Combination) African Chiroptera Report 2020 TAXONOMY: The placement of the Family Myzodidae, with in the superfamily Noctilionoidea follows Teeling et al. (2005). Simmons et al. (2013: 145) include seven families: Myzopodidae, Mystacinidae, Thyropteridae, Furipteridae, Noctilionidae, Mormoopidae, and 385 Phyllostomidae, which were already distinct by the end of the Eocene. COMMON NAMES: Czech: vampýrovci. Family MYZOPODIDAE Thomas, 1904 *1904. Myzopodidae Thomas, Proc. zool. Soc. Lond., 1904, II, I: 5. Publication date: 1 October 1904. - Comments: Type genus: Myzopoda A. Milne-Edwards and A. Grandidier, 1878. (Current Combination) TAXONOMY: Although they are currently assigned to different superfamilies, karyotypic data suggest that the closest relatives of the Myzopodidae are the Nycteridae Volleth et al. (2020a: 279). Currentl (Simmons and Cirranello, 2020) recognized extant genera: Myzopoda A. MilneEdwards and A. Grandidier, 1878. Known extinct genera: †Phasmatonycteris Gunnell, Simmons and Seiffert, 2014. COMMON NAMES: Czech: přísavkovcovití, myzopodovití, netopýrkové ušatí. Dutch: Madagassische hechtschijfvleermuizen. English: Sucker-footed Bats. French: Myzopodidés. German: Madagassische Haftscheibenfledermaus, Madagaskar-Haftscheibenfledermäuse. Norwegian: Sugeskålflaggermus. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Butler (1978: 67) suggests that the Myzopodidae are probably of African mainland origin, but survive only in Madagascar. Simmons et al. (2013: 146) indicate that one fossil species is known in this family. The family-level stem and crown ages were calculated by Shi and Rabosky (2015: 1537) as 54.1 and 1.1 mya, respectively. MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) mention a 2n value of 26 for the two species in this family. Genus Myzopoda Milne-Edwards and A. Grandidier, 1878 *1878. Myzopoda A. Milne-Edwards and A. Grandidier, Bull. Sci. Soc. Philom. Paris, sér. 7, 2: 220. Publication date: 22 June 1878. - Comments: Type species: Myzopoda aurita A. Milne-Edwards and A. Grandidier, 1878. - Etymology: From the Greek "μύξάω", meaning to suck (not from "μύξα", meaning mucus) and "πούς", meaning foot, referring to the sectional disk on the thumbs and feet (see Palmer, 1904: 446). (Current Combination) 1879. Myxopoda: Dobson, Proc. zool. Soc. Lond., 1878, IV: 871. Publication date: April 1879. - Comments: Lapsus (see Allen, 1939a: 82; Schliemann and Maas, 1978: 1) or emendation (see Palmer, 1904: 445). - Etymology: From the Greek "μύξα", meaning mucus, and "πούς", meaning foot, referring to the sectional disk on the thumbs and feet (see Dobson, 1879a: 281; Palmer, 1904: 445). (Lapsus) TAXONOMY: Currently Simmons and Cirranello, 2020) recognized species of the genus Myzopoda: aurita A. Milne-Edwards and A. Grandidier, 1878; schliemanni Goodman, Rakotondraparany and Kofoky, 2007. COMMON NAMES: Czech: přísavkovci, myzopody. English: Suckerfooted Bats. French: Chauves-souris à ventouses. German: MadagaskarHaftscheibenfledermäuse. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Butler (1978) found a Myzopoda sp. at Olduvai, Kenya, in deposits of early Pleistocene age. 386 ISSN 1990-6471 CONSERVATION STATUS: Assessment History Global Assessed as M. aurita VU (World Conservation Monitoring Centre, 1993); LC ver 3.1 (2001) (Jenkins et al., 2008h; IUCN, 2009). Russell et al. (2008b: 219) suggests that Myzopoda sp. may be among the few Malagasy verterbrate species to benefit from rapid deforestation, due to the species utilizing Ravenala as a day roost. Ravenala is a dominating pioneering species, and dense stands are found in areas of primary lowland forest which have been cleared (Goodman et al., 2007a). BIOGEOGRAPHY: Russell et al. (2008b) using coalescent analysis places the time of divergence of M. aurita and M. schliemanni at approximately 73,500 years ago. DETAILED MORPHOLOGY: Reproductive system - Carter and Enders (2016: 284) found Myzopoda sp. to have a smooth chorioallantois, suggesting that this genus belongs in the emballonurid clade. †Myzopoda africana Gunnell, Butler, Greenwood and Simmons, 2015 *2015. Myzopoda africana Gunnell, Butler, Greenwood and Simmons, Am. Mus. Novit., 3846: 1, 3, figs 2A, 2C, 3. Publication date: 16 December 2015. Type locality: Tanzania: Arusha province: Olduvai Gorge: Bed I, FLK NI, Layer 123. [Goto Description]. - Etymology: The name refers to the African continent as it is the only representative of the genus Myzopoda that is found on continental Africa. SIMILAR SPECIES: M. africana's humeri are on average 21 % larger than those of its living congeners (Gunnell et al., 2015a: 5). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pleistocene (Brown et al., 2019: Suppl.). Myzopoda aurita Milne-Edwards and A. Grandidier, 1878 *1878. Myzopoda aurita A. Milne-Edwards and A. Grandidier, Bull. Sci. Soc. Philom. Paris, sér. 7, 2: 220. Publication date: 22 June 1878. Type locality: Madagascar: "Madagascar". Holotype: MNHN ZM-MO-1997-770: alcoholic (skull not removed). Number 223 in Rode (1941); see Peterson et al. (1995: 139). (Current Combination) 1879. Myxopoda aurita: Dobson, Proc. zool. Soc. Lond., 1878, IV: 879. Publication date: April 1879. (Lapsus) TAXONOMY: See Schliemann and Maas (1978: 116). COMMON NAMES: Czech: přísavkovec ušatý, netopýrek ušatý, myzopoda ušatá. English: Golden Bat, Old World Sucker-footed Bat, Sucker-footed Bat, Madagascar Sucker-footed Bat, Eastern Suckerfooted Bat. French: Chauve-souris malgache à pieds à ventouses, Chauve-souris à pieds à ventouses de Madagascar, Vespertilion Doré, Myzopoda oreillard. German: Östliche Haftscheibenfledermaus. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) because it is widespread, thought to be locally common in areas of anthropogenic disturbance, and there are no obvious major threats (Jenkins et al., 2008h; IUCN, 2009; Monadjem et al., 2017p). Ralisata et al. (2010) refute the suggestion that this species is rare (Hutson et al., 2001). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017p). 2008: LC ver 3.1 (2001) (Jenkins et al., 2008h; IUCN, 2009). 1996: VU A2c ver 2.3 (1994). 1994: VU (Groombridge, 1994). 1990: VU (IUCN, 1990). Regional None known. MAJOR THREATS: There are no known major threats to this species. It is occasionally eaten by people when harvesting Ravenala madagascariensis plants (Jenkins et al., 2008h; IUCN, 2009; Monadjem et al., 2017p). African Chiroptera Report 2020 However, Ralisata et al. (2015: 95) do point out that this harvesting itself might lead to a loss of roosts, and as such has a negative impact on the bats. CONSERVATION ACTIONS: Jenkins et al. (2008h) [in IUCN (2009)] and Monadjem et al. (2017p) reported that M. aurita has only been recorded from a few protected areas: Parc National de Marojejy (Pont and Armstrong, 1990), Tampolo littoral forest (Ifticene et al., 2005) and near to Parc National de Masoala (Russ and Bennet, 1999) and Réserve Spéciale d'Analamazaotra (Russ and Bennet, 1999). Additional study is needed to develop an understanding of local population densities and precise habitat requirements. Ralisata et al. (2010) indicate that it used disturbed patches of vegetation and is not therefore threatened by deforestation, although it may be affected by loss of roosts for building materials. GENERAL DISTRIBUTION: Myzopoda aurita is endemic to the island of Madagascar (Goodman et al., 2007a). It is found in the humid zone of eastern and north-eastern Madagascar and appears to be most common at elevations lower than 500 m (Schliemann and Goodman, 2003) although it has been recorded at Andasibe (ca. 970 m). Native: Madagascar. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: For details, see a.o. Thomas (1904g). DENTAL FORMULA: Inc. 2 - 2 / 4, c 1 - 1 / 1 - 1, pm 3 - 3 / 3 - 3, m 3 - 3 / 3 - 3 (see Dobson, 1879a: 872). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 59) described the baculum from two specimens as being a small structure, triangular in shape, without a central shaft (length: 0.21, 0.26 mm; width: 0.15, 0.16 mm). In a third specimen they found a trace of a circular bone structure. FUNCTIONAL MORPHOLOGY: Schliemann (1970: 397; 1971; 78) and Schliemann and Goodman (2011: 313) studied the adhesive organs of Myzopoda, which are flat, disc-shaped skin duplications, primarily consisting of adipose tissue. He concluded that the static friction between these organs and a vertical surface is sufficient to enable the animals to hold on to this surface. 387 MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003) and Russell et al. (2008b). Karyotype - Richards et al. (2010) examined two females showing a 2n=26 and a Fna=48, with all chromosomes being bi-armed. FISH with Myotis probes revealed a X-autosome translocation. However, the autosome translocated to the X chromosome has not been counted for the Fna. The number of autosomal arms would then be 50 instead of 48. Protein / allozyme - Unknown. HABITS: Compositional analysis revealed that M. aurita males selected coffee plantations, degraded humid forest and wooded grasslands more than other habitats (Ralisata et al., 2010). ROOST: Ralisata et al. (2010) found 133 roosts consisting of the partially unfurled leaves of Ravenala madagascarensis (Sonn, 1782) and housing between nine and 51 individuals, which changed roosts every 1 - 5 days. Ralisata et al. (2015: 95) found that the leaves of the tree became available as roost between one and 19 days after the unfurling started and were inhabited between one and 12 days. DIET: Göpfert and Wasserthal (1995) noted that M. aurita - based on the analysis of fecal pellets of a single individual from north of Tolagnaro, southeast Madagascar - fed on smaller moths ("microlepidoptera"). Ralisata et al. (2010) found the diet comprised mainly of Lepidoptera and Coleoptera. The latter was confirmed by Rasoanoro et al. (2015: 64), who mention the following volume percentages: Aranea: 15.4 ± 2.38, Coleoptera: 26.7 ± 4.60, Diptera: 3.6 ± 3.59, Homoptera: 0.3 ± 0.33, Hymenoptera: 8.4 ± 3.58, Lepidoptera: 40.1 ± 7.31, and Neuroptera: 5.5 ± 5.07. Ramasindrazana et al. (2009: 161) reported the following percentage frequency / percentage volume: Lepidoptera (100 / 60.4 ± 9.54), Coleoptera (54.5 / 19.4 ± 8.44) , Blattaria (54.5 / 19.0 ± 6.32) and Aranea (9.1 / 1.2 ± 1.21). POPULATION: Structure See Russell et al. (2008b: 219). Density Russell et al. (2008b) estimated an effective population size of between 100,054 and 132,742 individuals. In some areas, M. aurita is locally common (P. A. Racey pers. comm.), but it is rarely 388 ISSN 1990-6471 trapped in large numbers during surveys (Russ and Bennet, 1999; Ifticene et al., 2005; Rakotondraparany and Medard, 2005; Jenkins et al., 2007b). Ralisata et al. (2010) caught on average of 5.7 (0 - 15) individuals per mist net night, at Kianjavato, where a total of 138 individiuals were captured on 24 nights of netting. Trend 2016: Unknown (Monadjem et al., 2017p). 2008: Unknown (Jenkins et al., 2008h; IUCN, 2009). UTILISATION: In the area around Tampolo forest, people collecting leaves of the Traveller's palm (Ravenala madagascariensis) for housing materials often encounter M. aurita roosting in the leaves, although no evidence that the bats were deliberately sought, they were collected and taken back to the village where they were cooked and eaten (Jenkins and Racey, 2008). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. ACTIVITY AND BEHAVIOUR: Ralisata et al. (2010) found that the foraging areas used by male bats varied between 7 and 108 ha (100 % minimum convex polygon), where the bats foraged close to the roost for the first hour after emergence, then travelled up to 1.8 km away and returned after 03h00. REPRODUCTION AND ONTOGENY: See Carter et al. (2008). Ralisata et al. (2010) suggest that juvenile males leave their mothers after weaning and join adult males. During October-November, the testes of males appeared larger than at other times of the year, although seasonal variation was not pronounced (Ralisata et al., 2010). PARASITES: Ralisata et al. (2010) did not observe any parasties on the 138 individual males captured. Figure 127. Distribution of Myzopoda aurita Myzopoda schliemanni Goodman, Rakotondraparany and Kofoky, 2007 *2007. Myzopoda schliemanni Goodman, Rakotondraparany and Kofoky, Mamm. biol., 72 (2): 65, 68, figs. 2 - 5. Publication date: 26 March 2007. Type locality: Madagascar: Mahajanga province: Parc National d'Anarafantsika: Ampijoroa, forestry station: Jardin Botanique A [16 19.4 S 46 48.4 E, 160 m] [Goto Description]. Holotype: FMNH 177327: ad ♂, skin and skeleton and skull. Collected by: Steven M. Goodman; collection date: 18 April 2003; original number: SMG-13624. 3 male and 4 female paratypes listed. - Etymology: The species name was chosen to honour Professer Dr. Harald Schliemann, University of Hamburg, in recognition of his long interest in sucker-footed bats, including several important anatomical studies (e.g. Schliemann, 1971; 2003). (Current Combination) 2016. Myz[opoda] schleimanni: Cardiff and Jenkins, Less. Conserv., 6: 90. Publication date: January 2016. (Lapsus) 2016. Myzopoda scliemanni: Amador, Arévalo, Almeida, Catalano and Giannini, J. mamm. Evolut., 23 (4): Suppl.. (Lapsus) 2016. Myzopoda scliemanni: Amador, Moyers Arévalo, Almeida, Catalano and Giannini, J. mamm. Evolut., 25 (1): Suppl. (for 2018). (Lapsus) TAXONOMY: Described by Goodman et al. (2007a) using external measurements and pelage coloration and supported by population genetics (Russell et al., 2008b). COMMON NAMES: Czech: přísavkovec bělobřichý. English: Schliemann's Sucker-footed Bat. French: Chauve-souris à ventouses de Schliemann. German: Schliemanns Haftscheibenfledermaus. African Chiroptera Report 2020 389 ETYMOLOGY OF COMMON NAME: The species name was chosen to honour Professor Dr. Harald Schliemann, University of Hamburg, in recognition of his long interest in sucker-footed bats, including several important anatomical studies (e.g. Schliemann, 1970, 1971). Ankaboka in the north (Goodman et al., 2007a) to Andranomanintsy in the south (Russell et al., 2008b). Goodman et al. (2007a) suggested that the distribution of range of M. schliemanni is broader than currently known and may extend north as far as Ambanja. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) as it has a relatively large geographic range and population size, and like other Myzopoda is thought to do well in highly modified habitats. It is unlikely to have been negatively impacted by deforestation (Jenkins et al., 2008I; IUCN, 2009; Monadjem et al., 2017q). However, it will be important to obtain information on the local abundance of this species (Monadjem et al., 2017q). Native: Madagascar. Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017q). 2008: LC ver 3.1 (2001) (Jenkins et al., 2008I; IUCN, 2009). 2004: Not assessed, but see Goodman et al. (2007a), who discuss the possible conservation status. Regional None known. MAJOR THREATS: The threats to this species are poorly known because of a lack of information on habitat preferences (Jenkins et al., 2008I; IUCN, 2009). Russell et al. (2008b) consider bats of the genus Myzopoda to benefit from deforestation. CONSERVATION ACTIONS: Myzopoda schliemanni has been trapped in Parc National de Namoroka and Parc National d'Ankarafantsika (Goodman et al., 2007a) as well as a recently created protected area in the Lac Kinkony - Mahavay area (Rakotoarivelo and Randrianandrianina, 2007). Based on current information this is a restricted-range species for which there are few data on natural history or ecology and basic field research is clearly needed to obtain information on the local abundance of this species (Jenkins et al., 2008I; IUCN, 2009; Monadjem et al., 2017q). It was not found during surveys of Parc National du Tsingy de Bemaraha (Kofoky et al., 2007), Parc National de Kirindy-Mité (Goodman et al., 2005a; Andriafidison et al., 2006a) or Kirindy CFPF (Goodman et al., 2005a). GENERAL DISTRIBUTION: Myzopoda schliemanni is known from central western Madagascar in lowland areas from GEOGRAPHIC VARIATION: See Goodman et al. (2007a). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Goodman et al. (2007a). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Goodman et al. (2007a). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 59) found in four specimens a triangular-shaped baculum, which was slightly more elongated than in M. aurita (length: 0.26 ± 0.078 mm, width: 0.12 ± 0.017 mm) and in a fifth specimen, the baculum was morphologically similar to that of the latter species. For a description of the crania, teeth, postcranial skeleton, ears and tragus see Goodman et al. (2007a). SEXUAL DIMORPHISM: Goodman et al. (2007a) found there is no sexual dimorphism in M. schliemanni. MOLECULAR BIOLOGY: DNA - Russell et al. (2008b). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: There are few accounts of its foraging habitats, although Rakotoarivelo and Randrianandrianina (2007) mist-netted 16 individuals in relatively disturbed forest. HABITS: See Goodman et al. (2007a). ROOST: Kofoky et al. (2006) observed M. schliemanni roosting in a cave. There are no other published observations of this species' roosting preferences, although it is assumed to also use broad-leaved plants such as Ravenala madagascariensis (Strelitziaceae) (Goodman et al., 2007a; Russell et al., 2008b). 390 ISSN 1990-6471 DIET: Rajemison and Goodman (2007) collected feacal samples from 18 individuals during mid-October from Andranomanitsy Forest, western Madagascar, where they reported four different insect orders. Prey belonging to the orders Lepidoptera and Blattaria were the most common in the samples and were more or less equally represtented (Rajemison and Goodman, 2007). PREDATORS: Goodman et al. (2015c: 78) found the remains of one individual in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. ACTIVITY AND BEHAVIOUR: Rajemison and Goodman (2007) suggest that M. schliemanni is capable of gleaning prey from vegetation (see Diet). REPRODUCTION AND ONTOGENY: See Goodman et al. (2007a) and Carter et al. (2008). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. POPULATION: Structure In Andranomanintsy forest, 8 of the 18 bats captured were males indicating a sex ratio of 1 : 1.25 (Goodman et al., 2007a: 312). Density Estimated effective population size using demographic models from two sites was between 61,453 and 134,630 individuals (Russell et al., 2008b). A roosting colony of 4 individuals was reported from a cave in Parc National de Namoroka (Kofoky et al., 2006). Trend 2016: Unknown (Monadjem et al., 2017q). 2008: Unknown (Jenkins et al., 2008I; IUCN, 2009). Figure 128. Distribution of Myzopoda schliemanni †Genus Phasmatonycteris Gunnell, Simmons and Seiffert, 2014 *2014. Phasmatonycteris Gunnell, Simmons and Seiffert, PLoS ONE, 9 (2): e86712: 6. Publication date: 4 February 2014 [Goto Description]. - Etymology: From the Greek "Phasma(to)" for apparition or spectre, in reference to the long ghost lineage connecting Fayum myzopodids with extant forms, and the Greek "Nykteris" for bat (see Gunnell et al., 2014: 7). - ZooBank: 693EEE77-DD52-408BB404-B689A43706AC. (Current Combination) TAXONOMY: Curently recognized species of the genus Phasmatonycteris: †butleri Gunnell, Simmons and Seiffert, 2014. †phiomensis Gunnell, Simmons and Seiffert, 2014. ZOOBANK: 693EEE77-DD52-408B-B404-B689A43706AC †Phasmatonycteris butleri Gunnell, Simmons and Seiffert, 2014 *2014. Phasmatonycteris butleri Gunnell, Simmons and Seiffert, PLoS ONE, 9 (2): e86712: 1, 7, figs 3A, 4. Publication date: 4 February 2014. Type locality: Egypt: Western Desert: Fayum Depression: Birket Qarun Formation: Fayum Quarry BQ-2 (23 meter level) [Goto Description]. - Etymology: In honour of Percy Butler, in recognition of his work on fossil bats from Africa and of his 75 year publishing career (see Gunnell et al., 2014: 7). - ZooBank: E6B5FFD8-5250-4C2A-B83FCC469DD32557. (Current Combination) African Chiroptera Report 2020 PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: The type specimen was found in the 23 meter level layer at Fayum Quarry BQ-2. This layer is about 37 million year old (Late Eocene, Priabonian - Brown et al., 2019: Suppl.). 391 GENERAL DISTRIBUTION: Egypt ZOOBANK: E6B5FFD8-5250-4C2A-B83F-CC469DD32557 †Phasmatonycteris phiomensis Gunnell, Simmons and Seiffert, 2014 *2014. Phasmatonycteris phiomensis Gunnell, Simmons and Seiffert, PLoS ONE, 9 (2): e86712: 1, 6, figs 2A–D, 3C, E–G. Publication date: 4 February 2014. Type locality: Egypt: Western Desert: Fayum Depression: Fayum Quarry I (242 m level) [Goto Description]. - Etymology: From "Phiom," Greek for the Fayum Region of Egypt’s Western Desert (see Gunnell et al., 2014: 7). ZooBank: 94812861-7911-438E-BD80-AB849541605B. (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: The type material was found in the 242 meter level Fayum Quarry I, in a layer of about 30 million years old (Early Oligocene, Rupelian - Brown et al., 2019: Suppl.). GENERAL DISTRIBUTION: Egypt. ZOOBANK: 94812861-7911-438E-BD80-AB849541605B INFRAORDER NYCTERIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 *2007. NYCTERIFORMACEI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the Infraorder is based on Nycteris G. Cuvier and E. Geoffroy, 1797 (type Family NYCTERIDAE Van der Hoeven, 1855). (Current Combination) GENERAL COMMENTS: We followed the format proposed for standardization of nomenclature of higher taxa by Alonso-Zarazaga (2005), then for typified names this would be standardized, using the connector acei and the ending -iformacei. But, for those without typified names there may well be occasions where incertae-sedis is used. TAXONOMY: Currently a single superfamily is known from within the Infraorder NYCTERIFORMACEI for Africa: NYCTEROIDEA Van der Hoeven, 1855. Superfamily NYCTEROIDEA Van der Hoeven, 1855 *1855. Nycteroidea Van der Hoeven. - Comments: The Superfamily name is based on Nycteris G. Cuvier and E. Geoffroy, 1797 (type Family NYCTERIDAE Van der Hoeven, 1855). (Current Combination) 2005. Emballonuroidea Teeling, Springer, Madsen, Bates, O'Brien and Murphy, Science, 307: 581. TAXONOMY: This name is used contrary to the superfamily name, EMBALLONUROIDEA, used by Teeling et al. (2005), Ravel et al. (2016: 377), and Amador et al. (2016: 22) because, the oldest, typified name is Nycteris G. Cuvier and E. Geoffroy, 1795, rather than Emballonura Temminck, 1838 (or even Emballonuridae Gervais, 1855). 392 ISSN 1990-6471 Family EMBALLONURIDAE Gervais, 1855 *1855. Emballonuridae Gervais, in: F. Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool, (Sec. 7), Vol. 1, pt. 2 (Mammifères): 62 footnote. Publication date: 23 July 1855. Comments: Type genus: Emballonura Temminck. - Etymology: The name of this family is derived from the Greek "έμβάλλω", meaning to throw in and "ούρά", meaning tail, referring to the tail structure of all its members, which resembles a spear thrown into the ground at an angle (see Palmer, 1904: 257; Taylor, 2005). (Current Combination) 1865. Brachyura Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 257. - Comments: Originally included the genera Mystacina J. Gray, 1843; Noctilio Linnaeus, 1766; Taphozous É. Geoffroy, 1818; Emballonura Temminck, 1838; Diclidurus Wied-Neuwied, 1820; and Furia F. Cuvier, 1828 [= Furipterus Bonaparte, 1837 [1832–1841] (see Jackson and Groves, 2015: 253). Pavlinov et al. (1995: 74) mentioned it "In part". Simmons (pers. comm.): "This name is a tough one, no type genus was noted and taxa now placed in 5 families were included. I have left it out because it is not clear just what it would be a junior synonym of. There seems no point in restricting it since it is not the oldest name for any of the family groups now recognized. Fixed". 1866. Diclidurina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 92. Publication date: 1 February 1866. 1866. Emballonurina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 92. Publication date: 1 February 1866. - Comments: Type genus: Emballonura Temminck, 1838. Proposed as tribe, and originally including the genera Urocryptus Temminck, 1838 [= Saccopteryx Illiger, 1811]; Diclidurus Wied-Neuwied, 1820; Saccopteryx Illiger, 1811; Emballonura Temminck, 1838; Proboscidea Spix, 1823 [= Rhynchonycteris Peters, 1867]; Centronycteris J. Gray, 1838; and Furia F. Cuvier, 1828 [= Furipterus Bonaparte, 1837 [1832–1841] (see Jackson and Groves, 2015: 253). 1875. Emballonuridae Dobson, Ann. Mag. nat. Hist., ser. 4, 16 (95): 349. Publication date: 1 November 1875. - Comments: Type genus: Emballonura Temminck, 1838. Originally included the subfamilies Emballonurinae Gervais, 1855 and Molossinae Gervais, 1855 (see Jackson and Groves, 2015: 253). 1893. Emballonurini Winge, Samling af Afhandlinger. E Museo Lundii, 2 (1): 24. - Comments: Proposed as tribe (?), and originally including the genera Mosia J. Gray, 1843; Emballonura Temminck, 1838; Coleura Peters, 1867; Saccopteryx Illiger, 1811; Rhynchonycteris Peters, 1867; Vespertiliavus Schlosser, 1887; Diclidurus Wied-Neuwied, 1820; and Taphozous É. Geoffroy, 1818 (see Jackson and Groves, 2015: 253). 1907. Diclidurinae Miller, Bull. U.S. natl. Mus., 57: 94. Publication date: 29 June 1907. Comments: Type genus: Diclidurus Wied, 1819. 1928. Emballonuroidea Weber, Die Säugetiere., 154. - Comments: Originally included the families Rhinopomatidae Bonaparte, 1838, Emballonuridae Gervais, 1855 and Noctilionidae J. Gray, 1821. Jackson and Groves (2015: 253) considered this name a synonym of the Emballonuridae. 2013. Emballanuridae: Taylor, Sowler, Schoeman and Monadjem, S. Afr. J. Wildl. Res., 43 (1): 18. Publication date: April 2013. (Lapsus) TAXONOMY: Koopman (1993a: 156) mentions Gervais, 1856 as author, whereas Corbet and Hill (1992: 83) mention Dobson, 1875e: 349. Simmons (2005) and Uvizl et al. (2019: 23) recognise two subfamilies: Emballonurinae (including the African genera Coleura and Emballonura), and Taphozoinae (including the African genera Saccolaimus and Taphozous). Goodwin (1942: 120) considers Diclidurinae to be a valid subfamily. Uvizl et al. (2019: 23), however, recognize the New World representatives of the Emballonurinae as differing on tribe level from the Old World forms: Diclidurini versus Emballonurini. Cytogenetic investigations by Volleth et al. (2020b: 268) led them to support the suggestion made by Ruedi et al. (2012: 210) that the Emballonurinae and Taphozoinae should be elevated to family rank, although neither of them do this explicitly. Currently (Simmons and Cirranello, 2020) recognized extant subfamilies, tribes, subtribes, and genera: Emballonurinae Gervais, 1855: Diclidurini Gray, 1866: Diclidurina Gray, 1866: Balantiopteryx Peters, 1867; Cormura Peters, 1867; Cyttarops Thomas, 1913; Diclidurus WiedNeuwied, 1820; Peropteryx Peters, 1867. Saccopterygina Lim, 2007: Centronycteris Gray, 1838; Rhynchonycteris Peters, 1867; Saccopteryx African Chiroptera Report 2020 Illiger, 1811. Emballonurini Gervais, 1855: Coleura Peters, 1867; Emballonura Temminck, 1838; Mosia Gray, 1843; Paremballonura Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, 2012. - Taphozoinae: Saccolaimus Temminck, 1838; Taphozous E. Geoffroy, 1818. COMMON NAMES: Castilian (Spain): Murciélagos de cola envainada. Czech: pochvorepovití, netopýři hladkonosí, embalonurovití. Dutch: Schedestaartvleermuizen. English: Sheath-tailed Bats, Tomb Bats, Sac-winged Bats. Finnish: Hautalepakot. French: Emballonuridés. German: Flattnasenfreischwänze, SackflügelFledermäuse, Glattnasen-Freischwänze. Italian: Emballonùridi. Norwegian: Frihalete flaggermus, gravflaggermus. Russian: Футлярохвостые. Vietnamese: Họ dơi bao. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The Emballonuridae are known from the early Miocene (Butler and Hopwood, 1957; Butler, 1969; 1978). 393 The stem and crown ages for this family were reported as 52.8 and 47.7 MYA by Shi and Rabosky (2015: 1537), and 51 and 45 MYA by Amador et al. (2016: 22). Ravel et al. (2016: 416) indicate that Chambinycteris pusilli is the most basal taxon of this family. PARASITES: The Old World Emballonuridae are host for the genus Ugandobia (Acari: Myobiidae), containing nine species (Fain, 1994: 1280). Two additional species were described from Hipposideridae and Pteropodidae, but Fain (1994: 1280) suggests this was due to contamination of the jars in which they were found, and that the representatives of this genus are actually restricted to the Emballonuridae. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola. †Genus Dhofarella Sigé, Thomas, Sen, Gheerbrant, Roger and Al-Sulaimani, 1985 *1994. Dhofarella Sigé, Thomas, Sen, Gheerbrant, Roger and Al-Sulaimani, Munch. Geowiss. Abh, (A) 26: 35 - 48. - Comments: See Gunnell et al. (2008: 5). (Current Combination) TAXONOMY: Morpholigally considered closer to Taphozous than to Vespertiliavus (Lim, 2007). Timeframe: Late Eocene (Priabonian) - Early Oligocene (Rupelian) (33.9 Mya). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The oldest known emballonurid fossils from Africa. Early Oligocene age (33.9 Mya) (Lim, 2007). GENERAL DISTRIBUTION: Oman (Sigé et al., 1994; Lim, 2007). †Dhofarella sigei Gunnell, Simons and Seiffert, 2008 *2008. Dhofarella sigei Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 5, fig. 6. Type locality: Egypt: Fayum Depression: Quarry L-41: Lower Sequence, Jebel Qatrani Formation. Holotype: CGM 83670:. A left dentary series m1-m3. - Etymology: Named for Bernard Sigé, in recognition of his many contributions to the study of Cenozoic bats (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). SIMILAR SPECIES: See Gunnell et al. (2008). Timeframe: Latest Eocene: Priabonian (37.97 - 33.23 mya), Lower Sequence, Jebel Qatrani formation. Early Oligocene: Rupelian (32.46 - 29 mya) (see Brown et al., 2019: Suppl.). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). 394 ISSN 1990-6471 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). †Dhofarella thaleri Sigé et al., 1994 *1994. Dhofarella thaleri Sigé, Thomas, Sen, Gheerbrant, Roger and Al-Sulaimani, Munich. Geowiss. Abh., (A) 26: 35 - 48. Type locality: Oman. (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Oligocene (see Lim, 2007). Subfamily Emballonurinae Gervais, 1855 *1855. Emballonurinae Gervais, in: F. Comte de Castelnau, Exped. Partes Cen. Am. Sud. Zool. (Sec. 7), Vol. 1, pt. 2 (Mammifères), 62 footnote. Publication date: 23 July 1855. TRIBE Emballonurini Gervais, 1855 *1855. Emballonurini Gervais, in: F. Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool. (Sec. 7), Vol. 1, pt. 2 (Mammifères), 62 footnote. Publication date: 23 Juny 1855. Genus Coleura Peters, 1867 *1867. Colëura Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 479. - Comments: Type species: Emballonura afra Peters. - Etymology: From the Greek "κολεόν" (coleon), meaning "sheath, scabbard" and "ούρά" (urá), meaning tail, referring to the tail being enveloped in the interfemoral membrane as far as the last caudal vertebra (see Palmer, 1904: 195; Lanza et al., 2015: 201). 1939. Coleura: Allen, Bull. Mus. comp. Zool., 83: 65. (Current Spelling) 2018. Coelura: Gunnell and Manthi, J. Hum. Evol., Suppl.. Publication date: 6 April 2018. (Lapsus) TAXONOMY: Key provided in Dunlop (1997, Mammalian Species, 566). Uvizl et al. (2019: 28) indicated that C. afra separated some 6.3 mya from the two other extant species, and the separation between the DjiboutiArabian group and the mainland Tanzanian group occurred some 3.8 mya. Currently (Simmons and Cirranello, 2020) recognized species of the genus Coleura: afra (Peters, 1852); gallarum Thomas, 1915; kibomalandy Goodman, Puechmaille, Friedli- Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, 2012; seychellensis Peters, 1868. Additionally, there is also the extinct †muthokai Wesselman, 1984. COMMON NAMES: Czech: buldočí pochvorepi. English: Sheathtailed bats, Peters's sheath-tailed bats. French: Chauves-souris à queue gainée. German: Schiebeschwanz-Fledermäuse. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Ethiopia, Kenya, Tanzania. African Chiroptera Report 2020 395 †Coleura muthokai Wesselman, 1984 *1984. Coleura muthokai Wesselman, Contributions to vertebrate evolution, 7: 52. - Etymology: The taxon is named in honor of Muthoka, son of Kivingo, a Wakamba tribesman from Kikoko, Kenya, who worked with Henry ("Hank") Barnard Wesselman as his assistant during the field seasons of 1972 and 1973 (Wesselman, 1984: 52). (Current Combination) TAXONOMY: Closely resembles the extant and larger species C. afra (Wesselman, 1984; Lim, 2007). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late Pliocene (2.58 Mya) of southwestern Ethiopia (Lim, 2007, Brown et al., 2019: Suppl.). Coleura afra (Peters, 1852) *1852. Emballonura afra Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 51, pl. 12, pl. 13, fig. 18, 19. Publication date: 1852. Type locality: Mozambique: Lower Zambesi River: Tete [16 10 S 33 35 E, 230 m] [Goto Description]. Syntype: BMNH 1858.6.18.12: ad ♀, alcoholic (skull not removed). Collected by: Prof. Wilhelm Carl Hartwig Peters. See Dobson (1878: 366) and Turni and Kock (2008: 23). Syntype: BMNH 1907.1.1.703:. See Turni and Kock (2008: 23). These latter specimens might possibly be the missing ZMB ones, but I did not find any proof of this (Van Cakenberghe, pers com). Syntype: RMCA [number unknown]:. See Jentink (1888b: 196) in Turni and Kock (2008: 23). These latter specimens might possibly be the missing ZMB ones, but I did not find any proof of this (Van Cakenberghe, pers com). Syntype: ZMB 429:. See Turni and Kock (2008: 23) [missing]. Syntype: ZMB 430:. See Turni and Kock (2008: 23) [missing]. Syntype: ZMB 54845: skull and alcoholic. (Skull damaged) see Turni and Kock (2008: 23). Syntype: ZMB 571a: ♂, skull and alcoholic. see Turni and Kock (2008: 23). Syntype: ZMB 67559: skull and alcoholic. See Turni and Kock (2008: 23). Syntype: ZMB 67561: skull and alcoholic. See Turni and Kock (2008: 23). Syntype: ZMB 67562: skull and alcoholic. See Turni and Kock (2008: 23). Syntype: ZMB 85667: ♂, complete skeleton. See Turni and Kock (2008: 23). Syntype: ZMB 85668: skin only. See Turni and Kock (2008: 23). Syntype: ZMB 85669: ♀. See Turni and Kock (2008). - Etymology: From the feminine Latin adjective àfra meaning "African" (see Lanza et al., 2015: 156). 1915. C[oleura] g[allarum] nilosa Thomas, Ann. Mag. nat. Hist., 8, 15 (90): 576, 577. Publication date: 1 June 1915. Type locality: Sudan: Upper Nile: Bahr-el-Zeraf, near mouth of [ca. 14 42 N 31 14 E] [Goto Description]. Holotype: BMNH 1915.3.6.76: ad ♂, skin only. Collected by: Willoughby P. Lowe; collection date: 3 February 1914; original number: 77. - Comments: Thorn et al. (2009: 41), however, mention 09 25 N 31 10 E for the coordinates. May be valid as a subspecies (Kock, 1969a; Koopman, 1975). In the key presented in the original description on p. 576, the name is given as C. g. nilosa, on the description itself (p. 577) it is mentioned as Coleura gallarum nilosa. 1939. Coleura kummeri Monard, Arq. Mus. Bocage, 10: 55, fig. 1. Publication date: March 1939. Type locality: Guinea-Bissau: Madina Boé [11 45 N 13 13 W] [Goto Description]. Syntype: MHNG 1326.003: ad. Collected by: Dr. Albert Monard. Presented/Donated by: ?: Collector Unknown. Syntype: MNHN ZM-MO-1940-1209 - 201: ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard. Mentioned ad paratype by Rode (1941: 244). This is either number 879, 884, 886 or 887 from Monard (1939: 55). Syntype: NMBA 5277: ad ♀, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 5 March 1938; original number: 881. See http://www.nmb.bs.ch/NaturmuseumBasel/LinksNMB/Sammlung/Kataloge/Mammalia.pdf . Syntype: NMBA 5278: ad ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 5 March 1938; original number: 890. See http://www.nmb.bs.ch/NaturmuseumBasel/LinksNMB/Sammlung/Kataloge/Mammalia.pdf . Syntype: RMCA 15313: ad ♀, alcoholic (skull not removed). Collected by: Dr. Albert Monard; original number: 883. Presented/Donated by: ?: Collector Unknown. Comments: Monard (1939: 55) used 13 specimens and three skulls for the description on this taxon. The whereabouts of specimens 879 ♂, 880 ♀, 882 ♀, 884 ♂, 886 ♂, 887 ♂, 396 ISSN 1990-6471 ? ? ? ? ? 888 ♀, 889 ♂, 891 ♀, 892 (3 skulls) has not yet been determined, although one of the male specimens is the syntype in the Paris Museum. Coleura afra afra: (Name Combination) Coleura afra kummeri: (Name Combination) Coleura afra nilosa: (Name Combination) Coleura afra: (Name Combination, Current Combination) Coleura gallarum nilosa: (Name Combination) TAXONOMY: See Dunlop (1997, Mammalian Species, 566). The recently discovered population in Madagascar could prove to be a distinct species (Goodman et al. (2005a). Vallo et al. (2018: 73) investigated the genetic and morphological variation of various African and Arabian populations and found that specimens from Kenya, Tanzania and Ghana could be assigned to the nominotypical subspecies. The (smaller) specimens from Yemen and Gabon represented two different lineages. Morphologically, these seemed to correspond with gallarum/nilosa (NE Africa + Arabia) and kummeri (W Africa), but this was not confirmed by their molecular analyses. The analyses by Uvizl et al. (2019: 29) suggest that Coleura afra might represent a complex of species rather than one highly variable species. COMMON NAMES: Afrikaans: Skedestertvlermuis. Chinese: 非洲鞘 尾 蝠 . Czech: pochvorep buldočí, netopýr buldočí. English: Southern Sheath-tailed Bat, African Sheath-tailed Bat, Sheath-tailed Bat, Mozambique Sheath-tailed Bat. French: Chauvesouris à queue gainée d'Afrique, Emballonure d'Afrique, Chauve-souris à queue en fourreau d'Afrique. German: Afrikanische Schiebeschwanz-Fledermaus. Italian: Còleura africàna. Portuguese: Morcego de cauda. ETYMOLOGY OF COMMON NAME: The colloquial name refers to the peculiar situation of the tail, which is enclosed by the interfemoral membrane between the back legs (Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008dq; IUCN, 2009; Monadjem et al., 2017n). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017n). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008dq; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004h; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are not well understood. Disturbance of roost sites could potentially be a threat, but the effect of this on the population is not well known (Mickleburgh et al., 2008dq; IUCN, 2009; Monadjem et al., 2017n). Kioko et al. (2015: 19) report a roadkill on the Arusha Highway (Tanzania). CONSERVATION ACTIONS: Mickleburgh et al. (2008dq) [in IUCN (2009)] and Monadjem et al. (2017n) indicate that on the African continent, there are no species-specific conservation measures in place, but there are some protected areas within its range. Both of the known sites for this species in Madagascar are within protected areas (the Ankarana Special Reserve and Namoroka National Park). GENERAL DISTRIBUTION: Coleura afra is found in much of east and easterncentral Africa, with an isolated record from central Mozambique. It is also found in West Africa, with records from Guinea and Guinea Bissau, Sierra Leone, northern Côte d'Ivoire, Ghana, Togo, Benin, and western Nigeria. There is also a small distribution in western Angola. This species was recently discovered in Madagascar (Goodman et al., 2005a) and is known only from the Ankarana Special Reserve in the north of the island and the Namoroka National Park in the west. Harrison (1967: 166) reported a specimen collected in the Strait of Bab el Mandeb, at the southern approach of the Red Sea. However, this might represent C. gallarum. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 535; Taylor et al., 2018b: 62); Benin; Burkina Faso (Kangoyé et al., 2012: 6024; 2015a: 614); Cameroon (Lebreton et al., 2014: 2); Central African Republic; Congo (The Democratic Republic of the); Côte d'Ivoire; Eritrea; African Chiroptera Report 2020 Ethiopia (Lavrenchenko et al., 2004b: 147); Ghana; Guinea; Guinea-Bissau (Monard, 1939; Rainho and Ranco, 2001: 39); Kenya; Madagascar (Goodman et al., 2005a); Mozambique Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 535) ; Nigeria (Happold, 1987); Sierra Leone (Weber et al., 2019: 25 - first record); Somalia; Sudan; Tanzania (including the island of Pemba [see O'Brien, 2011: 287]); Togo (Capo-Chichi et al., 2004: 161); Uganda. Presence uncertain: Burundi; Rwanda. Possibly extinct: Congo (Mickleburgh et al., 2008dq; IUCN, 2009). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Lucati and López-Baucells (2016: Suppl.) report on a piebald (all-white fur/skin patches, eyes always normally coloured) specimen from a cave in Kenya (https://www.inaturalist.org/observations/877909). DETAILED MORPHOLOGY: Baculum - Unknown. Brain - Based on three brains from Kenya, using immunohistochemical methods, Kruger et al. (2010a) describe the nuclear organization of the cholinergic, catecholaminergic and serotonergic systems, and Kruger et al. (2010b) also describe the distribution of Orexin-A immunoreactive cell bodies and terminal networks. SEXUAL DIMORPHISM: Kangoyé et al. (2015a: 614) confirm the sexual dimorphism found by Goodman et al. (2008b): in both body and cranial measurements, the females are larger than the males. ECHOLOCATION: Musila et al. (2018c: 43) [referring to Monadjem et al. (2010d: 246)] mention a narrow band, low-duty pulse call, composed of constant frequency, with peak frequency of 32.9 kHz and a bandwidth of 2.4 kHz. Luo et al. (2019a: Supp.) reported the following data (Hand released bats, 2 calls): Fpeak: 34.2, 31.2 kHz and duration: 10.7, 7.7 msec. 397 that other colonies occur in suitable rocky habitats in the southern Guinea savanna of western Nigeria (Mickleburgh et al., 2008dq). On Madagascar this bat taxon appears to be more varied in its habits, with some roost sites typically very dark and others quite close to the cave entrance. The two known sites on Madagascar are within dry deciduous tropical forest on a karst substrate (Goodman et al., 2005a). ROOST: On mainland Africa, the African sheath-tailed bat is found in well illuminated caves, rocky crevices and cellars of houses (Rosevear, 1965). It preferentially roosts in caves along the sea and lake shores (Kingdon, 1974). Lebreton et al. (2014: 2) report on a colony of about 5,000 specimens roosting in a cave in a rocky, forest-covered hill in Cameroon, which was also occupied by Rousettus aegyptiacus. Kangoyé et al. (2015a: 614) observed thousands of individuals in a cave on a hill at Néguéni, Burkina Faso. MIGRATION: Some authours suggest that Coleura afra migrates (Happold, 1987; Skinner and Smithers, 1990). PREDATORS: Mikula et al. (2016: Supplemental data) mention the Dark chanting goshawk (Melierax metabates Heuglin, 1861) as diurnal avian predator. POPULATION: Structure and Density:- In the African portion of Coleura afra distribution, it roosts in large colonies (in the 1,000s of animals) (Mickleburgh et al., 2008dq; IUCN, 2009; Monadjem et al., 2017n). The population and local abundance of this species in Madagascar is not known, but there is a minimum estimate of 500 individuals in one cave complex in Ankarana, Madagascar. Trend:- 2016: Unknown (Monadjem et al., 2017n). 2008: Unknown (Mickleburgh et al., 2008dq; IUCN, 2009). 2004: Unknown (Mickleburgh et al., 2004h; IUCN, 2004). See also Pye (1980) and Taylor et al. (2005). MOLECULAR BIOLOGY: Karyotype - Robbins (1983b: 35) reported 2n = 38 and FN = 68. HABITAT: In Nigeria, the African sheath-tailed bat is found in localised rocky habitats in southern Guinea savanna (Happold, 1987). Rosevear (1965) stated that they occur in woodland but only where there are acceptable roosting places. It is likely REPRODUCTION AND ONTOGENY: Krutzsch (2000: 113) indicates that C. afra is monoestrous in parts of their range and polyoestrous in others. McWilliam (1987b) observed that this species has two reproductive cycles per year in Kenya. He also showed that early postnatal growth rates in C. afra were higher during the short rainy season than during the long rainy season which followed an extended dry period. This can be attributed to the poor 398 ISSN 1990-6471 condition of females following a pregnancy when food resources were low. PARASITES: BACTERIA Bartonellae Bartonella - Kosoy et al. (2010: 1877) and Bai and Kosoy (2012: 58) reported a prelevence of Bartonella spp. In C. afra from Kenya of 4/9 (44.4 %) cultured from blood samples. Four genotypes of Bartonella spp. Identified in C. afra differed slightly from one another but clustered tightly within a genogroup (Kosoy et al., 2010: 1878). See also Kosoy (2010: 719) for further information. ACARI Fain (1959b: 249, 254) described Chirnyssus africanus (Sarcoptidae) from a C. afra specimen collected on Mount Wago, near Bukwa, Haut-Ituri, DRC and from a "Coleura gallarum" from Moba, DRC. In a subsequent paper, Fain (1959d: 150, 159) described Notoedres (Notoedres) benoiti (Sarcoptidae) from the same hosts and localities. HEMIPTERA Polyctenidae: Eoctenes coleura Maa, 1964 from Sudan (Haeselbarth et al., 1966: 16, host as Coleura gallarum). DIPTERA Streblidae: Raymondia huberi Frauenfeld, 1856 from Mozambique and Ethiopia (Haeselbarth et al., 1966: 102; Shapiro et al., 2016: 254). Raymondi seminuda Jobling, 1954 from Tanzania (Jobling (1954) in Haeselbarth et al. (1966: 104)). Nycteribiidae: Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114, host as Coleura gallarum; Obame-Nkoghe et al., 2016: 5, Gabon); Nycteribia schmidlii scotti Falcoz, 1923 (ObameNkoghe et al., 2016: 5, Gabon), which was infected by the blood parasite Polychromophilus melanipherus (Szentiványi et al., 2019; Suppl.). VIRUSES: Conrardy et al. (2014) tested 2 individuals from Kenya for the presence of adenoviruses, rhabdoviruses and paramyxoviruses, none of which tested positive. Maganga et al. (2014b) tested 94 C. afra samples collected from the Belinga caves of Gabon of which 14 tested positive to an unclassified paramyxovirus named Belinga bat virus (BelPV), one individual presented with haemorrhagic legions at necropsy. In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Coleura afra: Adenoviruses, Astroviruses, Paramyxoviruses. Coronaviridae - Coronaviruses SARS-CoV - During Febuary 2008, Pfefferle et al. (2009) tested 33 fecal samples from two localities in Ghana, and these were negative for coronavirus (CoV) RNA. None of the 85 Kenyan bats tested by Tao et al. (2017: Suppl.) were positive for CoV. Flaviviriduae Pegivirus Nieto-Rabiela et al. (2019: Suppl.) reported the presence of the non-Zoonotic Bat Pegivirus. Nairoviridae Orthonairovirus Six out of 14 C. afra specimens from Gabon tested by Müller et al. (2016: 3) were positive for Crimean Congo hemorrhagic fever virus (CCHFV). Paramyxoviridiae Drexler et al. (2012a: 4) report that morbili-related viruses found in Myotis myotis in Germany were also found in one Coleura afra specimen from Ghana (out of 96 examined). Maganga et al. (2014b) described a new Paramyxovirus (Belinga bat virus, BelPV) from 14 C. afra specimens from the Belinga caves (Gabon). One of these bats died due to several hemorrhagic lesions. None of the other bat species inhabiting the same caves were infected by this virus (Hipposideros cf. ruber, Hipposideros gigas, Miniopterus inflatus, Rousettus aegyptiacus, Rhinolophus alcyone). Picornaviridae Hepatovirus - Zeghbib et al. (2019: Suppl.) mention this virus from a bat from Ghana. Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 30 Kenyan C. afra specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). In their overview table, Maganga et al. (2014a: 8) reported that the following viruses were already found on C. afra: Morbillivirus unclassified. Fagre and Kading (2019: 4) also reported that serologic evidence was found for Crimean-Congo Hemorrhagic Fever Virus (CCHF). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Egypt, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Mozambique, Nigeria, Sierra Leone, Somalia, South Sudan, Sudan, Tanzania, Uganda. African Chiroptera Report 2020 399 Figure 129. Distribution of Coleura afra Coleura gallarum Thomas, 1915 1915. ? C[oleura] gallarum Thomas, Ann. Mag. nat. Hist., 8, 15 (90): 576, 577. Publication date: 1 June 1915. Type locality: Somalia: Zeyla [=Zsyla] [sea level] [Goto Description]. Holotype: BMNH 1911.8.2.4: ad ♀. Collected by: Dr. Ralph Evelyn Drake-Brockman; collection date: 29 October 1910; original number: 336. Presented/Donated by: Dr. Ralph Evelyn Drake-Brockman. - Comments: In the key presented in the original description on p. 576, the name is given as C. gallarum, on the description itself (p. 577) it is mentioned as Coleura gallarum. Coleura afra gallarum: (Name Combination) TAXONOMY: A phylogenetical analysis by Uvizl et al. (2019: 30) indicated that the cyt of the Arabian-Djiboutian lineage of "Coleura afra" differed by 5.0 to 6.3 %, which is similar to the distance between C. seychellensis and C. kibomalandy. This led these authors to raise gallarum in rank from subspecies to full species. GENERAL DISTRIBUTION: Native: southern Yemen, Oman, Djibouti, northwestern Somalia, and eastern Sudan (see Uvizl et al., 2019: 30). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Djibouti, Somalia, Sudan. Figure 130. Distribution of Coleura gallarum Coleura kibomalandy Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, 2012 *2012. Coleura kibomalandy Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, J. Mamm., 93 (6): 1442, 1446, figs 3, 4. Publication date: December 2012. Type locality: Madagascar: Antsiranana province: Ankarana National Park, 2.2 km ESE Amboandriky: Ambatoharanana cave (Crocodile cave) [12.98838S 49.02178E, 20 m] [Goto Description]. Holotype: FMNH 213598: ad ♀, skull and alcoholic. Collected by: Steven M. Goodman and Martinus Corrie Schoeman; 400 ISSN 1990-6471 collection date: 2 November 2010; original number: SMG 16954. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 183864: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 183865: ♂, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 183866: ♂, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 183867: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 183907: skull only. Collected by: Scott G. Cardiff. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213591: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213592: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213593: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213594: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213595: ♀, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213596: ♂, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213597: ♂, skull and alcoholic. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213599: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Martinus Corrie Schoeman. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213600: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Martinus Corrie Schoeman. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213601: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Martinus Corrie Schoeman. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213602: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Martinus Corrie Schoeman. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213603: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Martinus Corrie Schoeman. Presented/Donated by: ?: Collector Unknown. Paratype: UADBA 43497: ♂, alcoholic (skull not removed). Collected by: Richard K.B. Jenkins. Presented/Donated by: ?: Collector Unknown. - Etymology: The name kibomalandy is derived from a northern Malagasy dialect and means white (malandy) and belly (kibo) and refers to this species’ distinctive underside, which distinguishes it from other described species of Coleura, Paremballonura, and Emballonura (see Goodman et al., 2012a: 1446). (Current Combination) TAXONOMY: Goodman et al. (2012a) described a new species of Coleura based on morphology and molecular data. COMMON NAMES: English: Madagascar Sheath-tailed Bat. French: Emballonure de Madagascar. German: Madagaskar-Schiebeschwanzfledermaus. CONSERVATION STATUS: Global Justification Goodman et al. (2012a: 1449) suggest that this is a species of conservation concern due to its restricted distribution and its apparent limited number of individuals. While, Goodman (2017c) justifiy the Data Deficient status as this species is known from two sites in the western half of Madagascar, both within existing protected areas. Even though extensive bat surveys have been conducted in the region of one of these sites, vast areas of limestone karst remain largely unstudied in western Madagascar and we await further field data to properly assess this taxon. Assessment History Global 2016: DD ver. 3.1 (2001) (Goodman, 2017c). Regional None known. MAJOR THREATS: Threats to this species are currently unknown (Goodman, 2017c). CONSERVATION ACTIONS: Goodman (2017c) report that two sites this species is known from include the Ankarana National Park and Namoroka National Park. African Chiroptera Report 2020 GENERAL DISTRIBUTION: This species is known from two sites on Madagascar: the Ankarana National Park and the Namoroka National Park. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 57) described the baculum as a distinctly small structure, which, in dorsal view, has a sort of oblong shape. It is constricted along the midshaft and has a rounded tip that is smaller than the rounded basal portion. In lateral view, it tapers from the proximal to distal side, and it also is slightly constricted along the mid-shaft in this view. There is a slight hook to the tip. Length: 0.12 mm, width: 0.09 mm. 401 Trend:- 2016: Unknown (Goodman, 2017c). PARASITES: BACTERIA Gomard et al. (2016: 5) reported the presence of Leptospira bacteria in one out of three tested bats. VIRUSES: Mélade et al. (2016b: 4) found Paramyxoviruses in two out of six C. kibomalandy specimens they tested. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. ECHOLOCATION: Goodman et al. (2012a: 1448) mention the following parameters: low PF (< 35 kHz), narrow BW (< 7 ms), and short Dur (< 6 ms). HABITAT: Dammhahn and Goodman (2013: 108) indicate that the foraging habitat of this species is unknown, but probably is in open areas or in open forests. POPULATION: Structure and Density:- Goodman et al. (2012a: 1449) indicate that in July 2004, there were about 500 animals in the Ambatoharanana cave, and that no information is available on the population in the other cave. Population size and trend is not known for this species (Goodman, 2017c). Figure 131. Distribution of Coleura kibomalandy Coleura seychellensis Peters, 1868 *1868. Coleura seychellensis Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 367. Publication date: 1869. Type locality: Seychelles. Syntype: ZMB 3470a: ♂. Presented/Donated by: Prof. Edward Percival Wright. University of Dublin, see Turni and Kock (2008: 22). Syntype: ZMB 3470b: ad ♂. Presented/Donated by: Prof. Edward Percival Wright. University of Dublin, see Turni and Kock (2008: 22). Cotype: BMNH 1869.2.19.2: ad ♂. See Thomas (1915b: 579); Turni and Kock (2008: 22) indicate that the BMNH specimen can not be a (co-)type since Peters only used the ZMB specimens for his description, therefore they also reject Thomas' restriction of the type locality to Mahé Island. Comments: Allen (1939a: 65) mentions "1869, for 1868". - Etymology: Referring to the island group where the species occurs. (Current Combination) 1915. C[oleura] silhouettæ Thomas, Ann. Mag. nat. Hist., 8, 15 (90): 576, 578. Publication date: 1 June 1915. Type locality: Seychelles: Silhouette Island [Goto Description]. Holotype: BMNH 1906.3.18.2: ad ♂, skin and skull. Collected by: Prof. J. Stanley Gardiner. Presented/Donated by: Prof. J. Stanley Gardiner. - Comments: In the key on page 576 mentioned as C. silhouettæ, and in the description on p. 578 as Coleura silhouettæ. TAXONOMY: Two subspecies are recognized Coleura seychellensis seychellensis from Mahé and Praslin, and C. s. silhouettae from Silhouette and La Digue (Hill, 1971a: 575; Rocamora and Joubert, 2004; Simmons, 2005). 402 ISSN 1990-6471 COMMON NAMES: Creole: Sousouri banann. Czech: pochvorep seychelský, netopýr seychelský. English: Seychelles Sheath-tailed Bat. French: Chauvesouris à queue gainée des Seychelles, Emballonure des Seychelles, Chauve-souris à queue en fourreau des Seychelles. German: Seychellen Schiebeschwanz-Fledermaus. CONSERVATION STATUS: Global Justification Listed as Critically Endangered (CR C2a(i) ver 3.1 (2001)) because its population size is estimated to number fewer than 250 mature individuals, with no subpopulation greater than 50 individuals, and it is experiencing a continuing decline (Gerlach et al., 2008; IUCN, 2009; Jones et al., 2009b: 506). Bambini et al. (2005) reported that the bats may have become extinct on the island of Praslin within the first five years of the 21th century. While Mondajem et al. (2017da) report that the roosting cave at La Passe contained just 32 individuals in 2003, but more recently only 27 individuals were recorded there. Assessment History Global 2016: CR C2a(i) ver 3.1 (2001) (Mondajem et al., 2017da). 2008: CR C2a(i) ver 3.1 (2001) (Harrison and Lewis, 1961; IUCN, 2009). 2004: CR C2a(I,ii) ver 3.1 (2001) (IUCN, 2004; Mickleburgh et al., 2004g). 2002: CR B1+2cde, C2b, D (Mickleburgh et al., 2002a). 1996: CR (Baillie and Groombridge, 1996). 1994: EN (Groombridge, 1994). 1993 - EN (World Conservation Monitoring Centre, 1993). 1990: EN (IUCN 1990). 1988: EN (IUCN Conservation Monitoring Centre 1988). Collen et al. (2011: 2615) put this species on position 26 of their top 100 mammals that need urgent conservation attention on the basis of a combination of high Evolutionary Distinctivness (ED) and high threat status. Regional None known. MAJOR THREATS: Gerlach (2004) reports that the most catastrophic decline of this species probably occurred in the late 1800s and early 1900s when lowland forests were cleared, converting the mosaic of woodland and open gaps to intensively managed coconut plantations with no shrub layer to support the invertebrate diet of this species. Invasive plants including the Kudzu vine (Pueraria phaseoloides) and coconut scrub from abandoned plantations threaten remaining suitable habitat for this species. Kudzu vine threatens to overgrow roost cave entrances or to change the temperature gradient within caves (Gerlach and Taylor, 2006). Senior et al. (2007: 9) also added that - upon leaving their roosts - all the bats follow the same route, which follows the line with the least vegetation cover. Any encroachment of vegetation will make entry and exit to the roosts more difficult and could be a reason for roost abandonment. The species is sensitive to disturbance of roosting caves and this remains a threat to any active roost sites. Other suggested threats have been the predation of bats by the barn owl, Tyto alba, introduced in 1949, and feral cats (Gerlach et al., 2008; IUCN, 2009). Burgess and Lee (2004: 69) mention that pesticide contamination is probably not a significant threat. CONSERVATION ACTIONS: Gerlach et al. (2008) [in IUCN (2009)] and Mondajem et al. (2017da) report that Rocamora and Joubert (2004) recommend annual census of individuals along established transects; regular sensitive visits to all known roosting localities with additional surveys to locate any additional sites; reevaluation of the species distribution every three or four years; legal protection of all known roosting sites and their immediate surroundings; control of introduced predators (barn owl Tyto alba and cats); habitat protection within known feeding areas; public awareness campaigns; ongoing and additional research into the distribution, ecology and threats to this species. Gerlach and Taylor (2006) also outline the importance of removing invasive vegetation from existing and abandoned roost sites, and restoring lowland forests through the control of coconut and cinnamon, and replanting native vegetation. GENERAL DISTRIBUTION: This species is endemic to the Seychelles, and is currently found on the islands of Silhouette, Mahé (in the northwest of the island) and Praslin. It is thought to have become extinct on La Digue island and Praslin since 1980s (Gerlach et al., 2008). Senior et al. (2007: 3), however, indicated that these bats were only found on Silhouette and Mahé, and that they were last seen on Praslin in 2001 and on La Dique in 1976. The Zanzibar record (Simmons, 2005). is extremely dubious Native: Seychelles, Presence uncertain: Zanzibar. SEXUAL DIMORPHISM: Nicoll and Suttie (1982) [in Ball (2004: 136)] indicate that females are almost 10 % heavier than males: 10.2 grams versus 11.1 grams. African Chiroptera Report 2020 403 ECHOLOCATION: Burgess and Lee (2004: 72) report ultrasounds of 28 to 29 kHz, produced when flying in and out the entrances of their roosting caves. Inside the caves, sounds with a frequency between 24 and 26 kHz were produced. However, Clark (2004: 77) mentions sounds at around 30 - 40 kHz. Joubert (2004: 58) indicates that two harmonic bands are present: one at 40 kHz and one at 25 30 kHz. Ball (2004: 136) mentions an echolocation range from 20 to 50 kHz, but that it is usually around 30 kHz when flying within the roost. colony has increased to 40 in 2009, and that it split up in two parts of approximately 20 individuals in 2010, one remaining at La Passe and the second at Anse Lascars. HABITAT: This species has been recorded from coastal boulder field caves with stable cool temperatures and access into native palm woodland or marsh habitat (Gerlach and Taylor, 2006). Trend:- 2016: Decreasing (Mondajem et al., 2017da). 2008: Decreasing (Gerlach et al., 2008; IUCN, 2009). 2004: Decreasing (Mickleburgh et al., 2004g; IIUCN, 2004). ROOST: It appears to need boulder caves with horizontal ceilings; low, stable temperatures; and clear cave flyways not obscured by vegetation (Gerlach and Taylor, 2006). Abandoned roosts have been recorded from all four islands in the species historical distribution. A detailed study of the roosting site at La Passe is provided by Burgess and Lee (2004). DIET: Joubert (2004: 61) reports the following percentage of prey eaten by C. seychellensis: Diptera (30.5 %), Hymenoptera (25 %), Lepidoptera (19.5 %), Coleoptera (22 %), and Hemiptera (3 %). PREDATORS: Mikula et al. (2016: Supplemental data) mention C. seychellensis to be predated upon by Falco sp. and an unidentified species of diurnal avian predator. POPULATION: Structure and Density:- The species was noted by Wright (1868) to be 'very common in the neighborhood of the town of Port Victoria'. Joubert (2004) mentions that the species was seemingly still abundant up to the 1970s, and that the use of guano deposits as an indicator suggests that the magnitude of decline may have been as high as 90 %. The global population is now clearly very small, and although the precise number is not known, it is believed to be fewer than 100 mature individuals (Rocamora and Joubert, 2004; Gerlach et al., 2008). The roosting cave at La Passe was recorded to contain 32 bats in total in 2003 (Gerlach, 2004; Gerlach et al., 2008). More recently only 27 animals have been recorded roosting at this locality (Gerlach et al., 2008), but Gerlach (2011: 54) mentioned that the size of this Senior et al. (2007: 16) indicate that the population might be higher than estimated IF this species has indeed a harem structure as was suggested by Nicoll and Suttie (1982) as non-harem males would not roost with mating groups. However, this might also have a negative influence due to greater levels of inbreeding. ACTIVITY AND BEHAVIOUR: Joubert (2004) and Gerlach and Taylor (2006) provide specific details on the ecological requirements of this rare species. Smotherman et al. (2016: 537) refer to Gerlach (2009b) in describing the song-like structure for C. seychellensis as consisting of different syllable types in series. REPRODUCTION AND ONTOGENY: Nicoll and Suttie (1982) [in Krutzsch (2000: 113)] indicate that C. seychellensis is a polyoestrous species. MATING: Burgess and Lee (2004: 69) indicate that a harem structure might exist, but their observations were inconclusive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Seychelles. Figure 132. Distribution of Coleura seychellensis 404 ISSN 1990-6471 Genus Paremballonura Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, 2012 *2012. Paremballonura Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, J. Mamm., 93 (6): 1445. Publication date: December 2012 [Goto Description]. - Etymology: From the Greek prefix para-, meaning close to or besides, and the genus Emballonura Temminck, 1838 (see Goodman et al., 2012a: 1445). 2016. Paraemballonura: Amador, Moyers Arévalo, Almeida, Catalano and Giannini, J. mamm. Evolut., 25 (1): 43 (for 2018). Publication date: 24 November 2016. (Lapsus) TAXONOMY: Goodman et al. (2012a: 1440) found this genus to be paraphyletic, which resulted in the description of a new genus for the Malagasy Emballonura clade: Paremballonura. Uvizl et al. (2019: 29) estimate the separation between the two genera took place about 26.0 mya (Upper Oligocene). Curently (Simmons and recognized species of Cirranello, 2020) the genus of Paremballonura: atrata (Peters, 1874); tiavato (Goodman, Cardiff, Ranivo, Russell, and Yoder, 2006). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: There are fossil Paremballonura (refered to as Emballonura) of probable Pleistocene age (1.81 Mya) from Madagascar (McKenna and Bell, 1997; Lim, 2007). Paremballonura atrata (Peters, 1874) *1874. Emballonura atrata Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 693. Type locality: Madagascar: "Interior of Madagascar". Holotype: ZMB 4692: ♀. Collected by: Alfred Crossley. See Dobson (1878: 363) and Peterson et al. (1995: 56). Goodman et al. (2006a: 19) mention that the assumption by Peterson et al. (1995) that J.M. Hildebrandt is the collector, is wrong. Instead, Alfred Crossley is the collector, and they also indicate that the type locality remains unclear. 2012. Paremballonura atrata Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, J. Mamm., 93(6): 1445. (Name Combination, Current Combination) 2016. Paraemballonura atrata: Amador, Moyers Arévalo, Almeida, Catalano and Giannini, J. mamm. Evolut., 25 (1): Suppl. (for 2018). Publication date: 24 November 2016. (Lapsus) TAXONOMY: Following a recent taxonomic revision based on genetic and morphological comparisons, Paremballonura atrata is now confined to the east of Madagascar, with the smaller P. tiavato occurring in the west (Goodman et al., 2006a). Reviewed by Peterson et al. (1995). atrata species group Simmons (2005: 388). COMMON NAMES: English: Madagascar sheath-tailed bat, Peter's Sheath-tailed Bat, Peters's Sheath-tailed Bat. French: Chauve-souris malgache à queue en fourreau, Emballonure de Madagascar. German: Madagaskar-Freischwanzfledermaus. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Sabatier and Legendre (1985: 23) and Gunnell et al. (2014: 3) refer to fossils material from Tsimanampetsotsa, on Madagascar. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) because it is relatively widespread in eastern Madagascar and although its population is certainly decreasing as the humid forest is converted into agricultural areas there is no evidence that the decrease has been rapid enough to warrant Near Threatened or threatened status (Jenkins et al., 2008d; IUCN, 2009; Monadjem et al., 2017r). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017r). 2008: LC ver 3.1 (2001) (Jenkins et al., 2008d; IUCN, 2009). 1996: VU A2c ver 2.3 (1994) (IUCN, 1996). Regional None known. African Chiroptera Report 2020 MAJOR THREATS: Paremballonura atrata is collected by people for bushmeat in one part of eastern Madagascar (D. Andriafidison pers comm., in Jenkins et al., 2008d and Monadjem et al., 2017r). Otherwise, forest clearance for expanding agriculture threatens the foraging and roosting sites for this species (Jenkins et al., 2007b; Jenkins et al., 2008d; IUCN, 2009; Monadjem et al., 2017r). Jenkins et al. (2007b: 219) also point out that fire (started by farmers?) is a potential threat, as traces of fire were found at the entrances of their roosting caves. CONSERVATION ACTIONS: Jenkins et al. (2008d) [in IUCN (2009)] and Monadjem et al. (2017r) note that P. atrata has been recorded from Parc National de Mantadia (Randrianandriananina et al., 2006) and Parc National de Midongy du Sud (Goodman et al., 2006a) as well as from a network of protected sites around the Baie de Antogil (Goodman et al., 2006a). It probably does not require specific conservation action as long as forest vegetation is protected, but local populations should be monitored at roosting sites (Jenkins et al., 2008d; IUCN, 2009; Monadjem et al., 2017r). GENERAL DISTRIBUTION: This species is restricted to the humid zone of eastern Madagascar and has been recorded from around Maroantsetra in the north to Tolagnaro (Fort Dauphin) in the south (Goodman et al., 2006a). It is known from a wide elevational range, from as low as 30 m (Goodman et al., 2006a) to at least 900 m (Randrianandriananina et al., 2006). Native: Madagascar (Goodman et al., 2006a) - Ile Sainte-Marie (Rakotonandrasana and Goodman, 2007: 6). HABITAT: It is considered to be dependent on relatively intact forest (Goodman et al., 2005a; 2006a). ROOST: Paremballonura atrata is usually found roosting in caves and small crevices in or near to relatively intact humid forest (Goodman et al., 2006a; Randrianandriananina et al., 2006; Kofoky et al., 2007; Jenkins et al., 2007b). DIET: Based on droppings from five specimens, Rasoanoro et al. (2015: 64) report the following volume percentages of prey types: Coleoptera: 26.7 ± 8.09, Diptera: 40.9 ± 12.55, Hemiptera: 12.0 ± 1.16, and Hymenoptera: 31.3 ± 8.76. POPULATION: Structure and Density:- There is no information on population size. This species is rarely trapped in flight or detected acoustically and information on populations and distribution is mainly from bats found in caves (Jenkins et al., 2008d; Monadjem et al., 2017r). Colony size in caves rarely exceeds 20 individuals (Jenkins et al., 2007b). Trend:- 2016: Unknown (Monadjem et al., 2017r). 2008: Unknown (Jenkins et al., 2008d; IUCN, 2009). UTILISATION: Paremballonura atrata is collected by people for bushmeat in one part of eastern Madagascar (D. Andriafidison pers comm. in Jenkins et al., 2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. DETAILED MORPHOLOGY: Baculum: The baculum is small and circular in shape (Rakotondramanana and Goodman, 2017: 58); length: 0.09, 0.12 mm; width: 0.09, 0.12 mm. ECHOLOCATION: Kofoky et al. (2009: 379) reported the call from two individuals flown in a flight cage as FM/CF/FM with the most energy in the second harmonic at 52.9 kHz. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003) and Goodman et al. (2006a). Karyotype - Unknown. Protein / allozyme - Unknown. 405 Figure 133. Distribution of Paremballonura atrata 406 ISSN 1990-6471 Paremballonura tiavato (Goodman, Cardiff, Ranivo, Russell and Yoder, 2006) *2006. Emballonura tiavato Goodman, Cardiff, Ranivo, Russell and Yoder, Am. Mus. Novit., 3538: 1, 6, figs, 2, 3, 4, 5. Publication date: 19 October 2006. Type locality: Madagascar: Antsiranana province: Réserve Spéciale d'Ankarana, 2.6 km E Andrafiabe: Andrafiabe cave, in forest near [12 55.9 S 49 03.4 E, 50 m] [Goto Description]. Holotype: FMNH 169705: ad ♂, skin and skull. Collected by: Steven M. Goodman; collection date: 22 January 2001; original number: 11923a. - Etymology: From the Malagasy for "like rocks", referring to the tendency of this bat to occur in areas with exposed rock outcrops and caves (see Goodman et al., 2006a: 6). 2012. Paremballonura tiavato Goodman, Puechmaille, Friedli-Weyeneth, Gerlach, Ruedi, Schoeman, Stanley and Teeling, J. Mamm., 93(6): 1445. (Name Combination, Current Combination) 2016. Paraemballonura tiavato: Amador, Moyers Arévalo, Almeida, Catalano and Giannini, J. mamm. Evolut., 25 (1): Suppl. (for 2018). (Lapsus) TAXONOMY: Described as a new species in a split from "Emballonura atrata" (Goodman et al., 2006a). COMMON NAMES: Czech: pochvorep madagaskarský. English: Western Sheath-tailed Bat. French: Emballonure des rochers. German: TsingyFreischwanzfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) because although this species is declining due to the impacts of ongoing forest loss due to shifting agriculture and logging the reduction of suitable habitat is not thought to be sufficient to warrant listing in a higher category of threat. However, this species is not abundant within its range and requires close population monitoring and a reassessment may be needed in the near future (Jenkins et al., 2008e; IUCN, 2009; Monadjem et al., 2017s). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017s). 2008: LC ver 3.1 (2001) (Jenkins et al., 2008e; IUCN, 2009). Regional None known. MAJOR THREATS: Paremballonura tiavato is threatened by habitat loss due to slash-and-burn agriculture (tavy) and from charcoal collecting and logging (Jenkins et al., 2008e; IUCN, 2009; Monadjem et al., 2017s). Other potential threats come from the disturbance of roosting caves from tourists (Kofoky et al., 2007), fire (Jenkins et al., 2007b) or mining (Cardiff, 2006). CONSERVATION ACTIONS: Jenkins et al. (2008e) [in IUCN, 2009)] and Monadjem et al. (2017s) report that it is known from three protected areas, Parc National Tsingy de Bemaraha, Réserve Spéciale d’Ankarana, Réserve Spéciale d'Analamerana (Goodman et al., 2005a). GENERAL DISTRIBUTION: Endemic to Madagascar (Goodman et al., 2006a). It is restricted to the karstic lowland areas in the western part of the island, extending from the drier areas in the north in Diana Region to Parc National Tsingy de Bemaraha in the west (Goodman et al., 2005a). A record from Toliara in the extreme south has also been attributed to this species (Peterson et al., 1995), but the specimen was not examined as part of the recent reassessment of Emballonura taxonomy on Madagascar (Goodman et al., 2006a). The distribution of this species would be significantly reduced if the Toliara specimen was omitted. It is currently known to range between 10 and 330 m elevation (Goodman et al., 2006a). Native: Madagascar (Goodman et al., 2005a; Goodman et al., 2006a) - Nosy Be (Peterson et al., 1995: 57; Rakotonandrasana and Goodman, 2007: 6) and Nosy Komba (Rakotonandrasana and Goodman, 2007: 6). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Goodman et al. (2006a). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Goodman et al. (2006a). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 58) indicate that the baculum of this species is more African Chiroptera Report 2020 elongated than the rounded form of P. atrata; length: 0.09 mm, width: 0.08 mm. For a description of the crania, teeth, ears and tragus see Goodman et al. (2006a). ECHOLOCATION: Kofoky et al. (2009: 377) reported the call as a FM/CF/FM with up to five harmonics, produced short calls with a medium IPI and a low duty cycle, with a maximum energy at 55.2 kHz. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Unknown. Protein / allozyme - Unknown. 407 DIET: Lepidoptera (Razakarivony et al., 2005; Goodman et al., 2006a). POPULATION: Structure and Density:- Roosting colonies of E. tiavato are usually relatively small (< 20 individuals) (Jenkins et al., 2008e; IUCN, 2009; Monadjem et al., 2017s). Trend:- 2016: Unknown (Monadjem et al., 2017s). 2008: Unknown (Jenkins et al., 2008e; IUCN, 2009). REPRODUCTION AND ONTOGENY: See Goodman et al. (2006a). HABITAT: It appears to be associated with relatively intact forest and is believed to be forest dependent (Goodman et al., 2005a; Goodman et al., 2006a). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Dammhahn and Goodman (2013: 108) indicate that the lower portion of forest is this bat's foraging habitat. HABITS: At night it has been observed resting on buildings (Goodman et al., 2006a). It feeds in forest understorey and over small streams with bank side vegetation (Robinson et al., 2006). ROOST: Usually found roosting in the entrance to caves and rock overhangs that receive weak sunlight (Cardiff, 2006; Goodman et al., 2006a; Kofoky et al., 2007). Figure 134. Distribution of Paremballonura tiavato †Genus Pseudovespertiliavus Ravel, 2016 *2016. Pseudovespertiliavus Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 388. Publication date: 30 September 2016. - Etymology: The name refers to the multiple morphological similarities with the genus Vespertiliavus (see Ravel et al. 2016: 388). †Pseudovespertiliavus parva Ravel, 2016 *2016. Pseudovespertiliavus parva Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 388, figs 16, 17. Publication date: 30 September 2016. Type locality: Algeria: Béchar province: Gour Labiz region (Sahara, Hammada du Dra).: Glib Zegdou [29 42 49 N 04 45 58 W] [Goto Description]. Etymology: From the Latin parva meaning small, referring to the small size of this bat (see Ravel et al., 2016: 388). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene (Lutetian). 408 ISSN 1990-6471 Subfamily Taphozoinae Jerdon, 1867 1867. 1988. Taphozoinae Jerdon, Mammals of India, 30. - Comments: Jackson and Groves (2015: 253) mention 1867 as year of publication. Other authors (e.g. Palmer, 1904: 773) mention 1874 or 1877. These latter dates represent reprints of the "Mammals of India", but the name was already published in the 1867 version. Considered a valid subfamily by McKenna and Bell (1997: 302), Simmons (2005: 381), Jackson and Groves (2015: 253). Taphozoini Robbins and Sarich, J. Mamm., 69 (1): 10. Publication date: 25 February 1988. - Comments: Proposed as tribe and originally included the genera Taphozous É. Geoffroy, 1818 and Saccolaimus Temminck, 1838 (see Jackson and Groves, 2015: 253). Genus Saccolaimus Temminck, 1838 *1838. Saccolaimus Temminck, in: Van der Hoeven, Tijdschr. Nat. Gesch. Physiol., 5: 14. Comments: Corbet and Hill (1992: 85), Grubb et al. (1998: 72) mention "1838: 6 (and 1841a: 277, 279) [C&H]". Type species: Taphozous saccolaimus Temminck, 1838. Etymology: From the Greek "σάκκος", meaning sac and "λαιμός", meaning throat, gullet, referring to the well-developed gular sacs of the type species (see Palmer, 1904: 615; Flannery, 1995a: 328; 1995b: 401). (Current Combination) 1876. Taphonycteris Dobson, Proc. zool. Soc. Lond., 1875, IV: 548. Publication date: April 1876. - Comments: Type species: Taphozous saccolaimus Temminck, 1838, by subsequent designation by Palmer (1904: 661) (see Jackson and Groves, 2015: 254). Etymology: From the Greek "τάφος", meaning grave and "νυκτερίς", meaning bat (see Palmer, 1904: 661). TAXONOMY: Considered a subgenus of Taphozous by Ellerman and Morrison-Scott (1951: 104), Corbet and Hill (1980: 45), Corbet and Hill (1992: 85), and Kock and Dobat (2000: 87), but see Barghoorn (1977: 5), Robbins and Sarich (1988) and Chimimba and Kitchener (1991). Key to the species provided by Chimimba and Kitchener (1991). Currently (Simmons and Cirranello, 2020) recognized species of the genus Saccolaimus: flaviventris (Peters, 1867) – Australia (except Tasmania), southeastern New Guinea (Simmons, 2005: 382); mixtus Troughton, 1925 – southeastern New Guinea, northeastern Queensland (Australia) (Simmons, 2005: 382); peli (Temminck, 1853); saccolaimus (Temminck, 1838) – Bangladesh, India and Sri Lanka through south east Asia (including Burma, Cambodia, Thailand and the Nicobar Islands) to the Philippines, Sulawesi and Borneo, Sumatra, Java, Bali and Timor (Indonesia); New Guinea; New Britain and Bougainville Islands (Papau New Guinea); northeastern Queensland (Australia); Guadalcanal Island (Solomon Islands) (Simmons, 2005: 382). And there is the Wesselman, 1984. extinct African †abitus COMMON NAMES: Czech: vakoví hrobkovci. English: Pouched Bats. French: Saccolaime. German: Taschenfledermäuse. †Saccolaimus abitus (Wesselman, 1984) *1984. Taphozous abitus Wesselman, Contributions to vertebrate evolution, 7: 44. Type locality: Ethiopia: upper member B, Shungura Formation. - Etymology: From the Latin verb abeo meaning to go away, the past participle of which is abitus meaning vanished or departed (Wesselman, 1984). 2004. Saccolaimus abitus: Lim. (Name Combination, Current Combination) 2011. Saccolaimus abditus: Gunnell, Eiting and Geraads, N. Jb. Geol. Paläont. Abh., 260 (1): 70. Publication date: January 2011. (Lapsus) African Chiroptera Report 2020 TAXONOMY: Closest affinity to the extant Saccolaimus peli (Wesselman, 1984; Lim, 2007). 409 Late Pliocene (2.58 Mya) of southwestern Ethiopia (Wesselman, 1984; Lim, 2007, Brown et al. (2019: Suppl.) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: †Saccolaimus kenyensis Gunnell and Manthi, 2018 *2018. Saccolaimus kenyensis Gunnell and Manthi, J. Hum. Evol., p. "4", fig. 5D. Publication date: 6 April 2018. Type locality: Kenya: Nzube's Mandible Site [02 24 00 N 36 06 00 E] [Goto Description]. Holotype: NMK KP-44927: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. The holotype consists of a M 1. - Etymology: Referring to country where the type was found (Gunnell and Manthi, 2018: "5"). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pliocene (Brown et al., 2019: Suppl.). Saccolaimus peli (Temminck, 1853) *1853. Taphozous peli Temminck, Esquisses zoologiques sur la Côte de Guiné. 1e partie, les Mammifères, 82. Type locality: Ghana: Boutry River [=Butre river] [ca. 04 49 N 01 55 W] [Goto Description]. ? Saccolaimus peli: (Name Combination, Current Combination) ? Taphozous (Saccolaimus) peli: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Chinese: 贝 尔 墓 蝠 . Czech: hrobkovec guinejský. English: Pel's Pouched Bat. French: Saccolaime de Pel, Chauve-souris à queue gainée de Pel. German: Pels Taschenfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bl; IUCN, 2009; Monadjem et al., 2017ap). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ap). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bl; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bn; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Selective logging of large trees with cavities results in substantial roost destruction. Parts of S. peli range are fragmented by agricultural activities, but as it lives in clearings, it is not known whether or not this is having a significant negative impact on the species (Mickleburgh et al., 2008bl; IUCN, 2009; Monadjem et al., 2017ap). CONSERVATION ACTIONS: Mickleburgh et al. (2008bl) [in IUCN (2009)] and Monadjem et al. (2017ap) report that it is found in several protected areas. Research is needed to determine the effects of habitat fragmentation on the species. GENERAL DISTRIBUTION: Saccolaimus peli ranges widely through the equatorial forest belt of west and central Africa. In the Upper Guinea region, it occurs in Liberia, Guinea, Côte d'Ivoire and Ghana. Its main range extends from southwestern Nigeria to western Uganda, with isolated records from western Kenya and eastern Angola. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 535); Cameroon; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 535); Côte d'Ivoire; Equatorial Guinea; Gabon; Ghana; Guinea; 410 ISSN 1990-6471 Kenya; Liberia; Nigeria (Capo-Chichi et al., 2004: 161); Uganda. Presence uncertain: Central African Republic; Sierra Leone. Bates et al. (2013: 338) reject the presence of S. peli in the Republic of Congo because Malbrant and Maclatchy (1949) only mention it 'likely' to occur in that country. FUNCTIONAL MORPHOLOGY: Bahlman et al. (2016: 122) indicate that S. peli has long, cartilaginous distal phalanges with tendons running to the distal-most portion of the tip of digits IV and V. They speculate that the flexor muscles and tendons not only flex the terminal IP joint, but also may provide some additional curvature along the length of the cartilaginous phalanges. Trend:- 2016: Unknown (Monadjem et al., 2017ap). 2008: Unknown (Mickleburgh et al., 2008bl; IUCN, 2009). REPRODUCTION AND ONTOGENY: Krutzsch (2000: 113) assigned S. peli to the group of polyoestous species. PARASITES: Nycteribiidae: Tripselia aequisetosa Theodor, 1956 from Liberia and Congo (Haeselbarth et al., 1966: 112). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Liberia, Nigeria, Uganda. MOLECULAR BIOLOGY: Karyotype - Robbins (1983b: 35) reported 2n = 48 and FN = 68, with a metacentric X and an acrocentric Y chromosome. HABITAT: R.B. Woosnam [in Thomas, 1910b: 489] reported that vaste numbers were seen in the Congo Forest, but not anywhere else. He also indicated that they were seen hanging in trees, but never in houses. POPULATION: Structure and Density:- S. peli occurs in low densities across its wide range (Mickleburgh et al., 2008bl; IUCN, 2009; Monadjem et al., 2017ap). Figure 135. Distribution of Saccolaimus peli Genus Taphozous E. Geoffroy St.-Hilaire, 1818 *1818. Taphozous E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 113. Publication date: 1818. - Comments: Type species: Taphozous perforatus E. Geoffroy, 1818, by monotypy (according to Colket and Wilson, 1998: 1) or subsequent designation by Miller (1907: 93) (according to Jackson and Groves, 2015: 255). Mahoney and Walton (1988a: 116) in Grubb et al. (1998: 72) considered the date of Geoffroy's publication to be 1813. Gardner and Hayssen (2004: 12) mention that the correct date should be 1818, not 1813 as mentioned on the publication itself. - Etymology: From the Greek "τάφος" (táfos), meaning "grave" or "tomb" and "ξωός" (zóos), meaning "living" (see Palmer, 1904: 662). (Current Combination) 1821. Thaphosores: Gray, London Med. Repos., 15 (1): 300. - Comments: Jackson and Groves (2015: 256) indicate that this is possibly an incorrect subsequent spelling as Gray refered to "Geoff." as author. Generally, the name is given as "Taphosores", but Gray (1821: 300) actually mentioned Thaphosores. (Lapsus) 1821. Thaphozous: Bowdich, An analysis of the natural classification of mammalia, for the use of students and travellers., 30. (Lapsus) 1822. Thaphozus: Fleming, The Phylosophy of Zoology, 179. (Lapsus) 1922. Liponycteris Thomas, Ann. Mag. nat. Hist., ser. 9, 9 (51): 267. Publication date: 1 March 1922 [Goto Description]. - Comments: Type species: Taphozous nudiventris Cretzschmar, 1830, by original designation (see Colket and Wilson, 1998: 1; Jackson and Groves, 2015: 256). Valid as a subgenus. African Chiroptera Report 2020 1998. 2018. ? 411 Tapozous: Yom-Tov and Kadmon, Diversity Distrib., 4: 67. (Lapsus) Thapozous: Leopardi, Holmes, Gastaldelli, Tassoni, Priori, Scaravelli, Zamperin and De Benedictis, Infec. Gen. Evol., 58: 286. Publication date: 30 January 2018. (Lapsus) Taphozous sp.: TAXONOMY: Some authors (e.g. Allen, 1939a: 66) consider Oken (1816), Lehrbuch d. Naturg., pt. 3., zool., sect 2, p. 926 to be the author of this genus. Meester et al. (1986): Barghoorn (1977), followed by Koopman (1982, and in litt.) regard Saccolaimus as generically distinct from Taphozous, but L.W. Robbins (Abstracts, 4th International Colloquium on Taxonomy and Ecology of African Small Mammals: 33, 1984) states that on the basis of protein electrophoresis and immunology Taphozous and Saccolaimus are closely related. Simmons (2005:382) recognized two subgenera Taphozous and Liponycteris, but this was rejected by Uvizl et al. (2019: 31). Only members of the former subgenus Taphozous occur in Africa and its associated islands. Includes Liponycteris but not Saccolaimus; see Hayman and Hill (1971: 15) and Barghoorn (1977: 5), Robbins and Sarich (1988) and Chimimba and Kitchener (1991), though also see Corbet and Hill (1992). Corbet and Hill (1992: 85) include Saccolaimus as a subgenus. A key is provided by Colket and Wilson (1998: 1, Mammalian Species, 597). Currently (Simmons and Cirranello, 2020) recognized species of the genus Taphozous: achates Thomas, 1915 – Kei, Savu, Roti, Semau and Nusa Penida Isls (Indonesia) (Simmons, 2005: 382); australis Gould, 1854 – northern Queensland (Australia), Torres Strait Isls, southeastern New Guinea (Simmons, 2005: 383); georgianus Thomas, 1915 – northern and western Australia (Simmons, 2005: 383); hamiltoni Thomas, 1920; hildegardeae Thomas, 1909; hilli Kitchener, 1980 – Western Australia, South Australia and Northern Territory (Simmons, 2005: 383); kapalgensis McKean and Friend, 1979 – Northern Territory (Australia) (Simmons, 2005: 383); longimanus Hardwicke, 1825 – Sri Lanka; India and Bangladesh to Burma, Cambodia and Thailand; Peninsular Malaysia; Sumatra, Borneo, Java, Bali, Sumbawa and Flores (Indonesia) (Simmons, 2005: 383); mauritianus E. Geoffroy Saint-Hilaire, 1818. melanopogon Temminck, 1841 – Sri Lanka; India; Burma; Thailand; Laos; Cambodia; Vietnam; southern China; Malay Peninsula and adjacent islands; Borneo; Sumatra, Java, Lombok, Sumbawa, Moyo, Alor, Timor and Sulawesi (Indonesia), Philippines (Simmons, 2005: 384); nudiventris Cretzschmar, 1830; perforatus E. Geoffroy Saint-Hilaire, 1818; theobaldi Dobson, 1872 – central India to Vietnam; Java, Borneo and Sulawesi (Simmons, 2005: 385); troughtoni Tate, 1952 – northwestern Queensland (Australia) (Simmons, 2005: 385). COMMON NAMES: Czech: praví hrobkovci, wečerníkové, netopýři bradožlází. English: Tomb Bats, Pouched Bats. French: Chauve-souris des Tombes, Taphiens. German: Grabfledermäuse, Grabflatterer. Italian: Tafozòi, Pipistrèlli délle tómbe. ETYMOLOGY OF COMMON NAME: The type species of the genus, T. perforatus, was taken from a royal tomb in Egypt (see Taylor, 2005). PARASITES: HEMIPTERA Polyctenidae: Eoctenes intermedius (Speiser, 1904) originally described from a Taphozous species from Egypt (Haeselbarth et al., 1966: 16). DIPTERA Streblidae: Brachytarsina alluaudi (Falcoz, 1923) (Haeselbarth et al., 1966: 101). VIRUSES: Flaviviridae Pegivirus (BPgV) 2 out of 2 unidentified Taphozous specimens from Cameroon were infected by clade G type Pegivirus, and one of them also with clade K type Pegivirus (Quan et al., 2013: Table S5). Paramyxoviridae Mortlock et al. (2015: 1841) reported that three out of 12 examined Taphozous sp. specimens from Cameroon tested positive for Paramyxovirus sequences. Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 1 Kenyan Taphozous sp. specimen, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Chad, Ethiopia, Senegal, South Sudan, Sudan, Tanzania, Uganda, Zambia. 412 ISSN 1990-6471 †Taphozous incognita (Butler & Hopwood, 1957) *1957. Saccolaimus incognita Butler and Hopwood, Fossil Mammals of Africa, Insectivora and Chiroptera from the Miocene rocks of Kenya, 13: 29, fig. 9. Publication date: 1957. Type locality: Kenya: Koru. Holotype: BMGD M.14222:. The left half of a rostrum lacking much of the bone, but showing the interior of the orbit and part of the crowns of P4 and M2. 1978. Taphozous incognita: Butler, Evolution of African Mammals, 65. (Name Combination, Current Combination) TAXONOMY: Most similar to the extant Taphozous nudiventris (Butler and Hopwood, 1957; Butler, 1984; Lim, 2007). Early Miocene (Burdigalian [Brown et al., 2019: Suppl.] - 23.03 Mya) of the Legete Formation in Kenya (Lim, 2007), Koru (Butler, 1978). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya. Taphozous hamiltoni Thomas, 1920 *1920. Taphozous hamiltoni Thomas, Ann. Mag. nat. Hist., ser. 9, 5 (25): 142. Publication date: 1 January 1920. Type locality: Sudan: Equatoria province: Mongalla [05 10 N 31 50 E, 500 m] [Goto Description]. Holotype: BMNH 1919.12.18.1: ad ♀. Collection date: 13 June 1918. Presented/Donated by: The Wellcome Research Laboratories. Formerly Stevenson Hamilton collection, nr 118 (see Thomas, 1920: 143). - Etymology: In honour of Major James Stevenson-Hamilton (1867 - 1957), eminent British regular soldier and conservationist, born in Ireland, considered father of the Kruger National Park. The species is named in his honor as the type specimen came from his collection (see Lanza et al., 2015: 210). (Current Combination) ? Taphozous (Liponycteris) hamiltoni: (Name Combination) TAXONOMY: See Simmons (2005). Regional None known. COMMON NAMES: Chinese: 苏 丹 墓 蝠 . Czech: hrobkovec východoafrický. English: Hamilton's Tomb Bat, Hamilton's Naked-rumped Tomb Bat, Hamilton's Naked-bellied Tomb Bat, Kordofan Sheathtail-bat. French: Taphien de Hamilton. German: Hamiltons Grabfledermaus. Italian: Tafozòo di Hàmilton. MAJOR THREATS: The threats facing this species are unknown. It could be declining due to habitat loss through agriculture, but the data to confirm this are lacking (Mickleburgh et al., 2008x; IUCN, 2009). CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of continuing uncertainties as to its extent of occurrence, status, trends and ecological requirements (Mickleburgh et al., 2008x; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008x; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004x; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). CONSERVATION ACTIONS: Mickleburgh et al. (2008x) [in IUCN (2009)] report that no specific conservation measures are in place. It probably occurs in some protected areas. The distribution and ecological requirements of this species require further investigation. GENERAL DISTRIBUTION: Taphozous hamiltoni is not well known, and there are very few records. It is known mainly from east Africa (including records from Sudan, Somalia, Kenya, and Tanzania), and there is one additional record from southern Chad. Native: Chad; Kenya; Somalia; Sudan; Tanzania; Uganda. African Chiroptera Report 2020 413 POPULATION: Structure and Density:- There is no information, as there are few records of this species (Mickleburgh et al., 2008x; IUCN, 2009). Trend:- 2008: Unknown (Mickleburgh et al., 2008x; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, South Sudan, Sudan. Figure 136. Distribution of Taphozous hamiltoni Taphozous hildegardeae Thomas, 1909 *1909. Taphozous hildegardeæ Thomas, Ann. Mag. nat. Hist., ser. 8, 4 (20): 98. Publication date: 1 August 1909. Type locality: Kenya: Coast province: near Mombassa: Rabai [700 ft] [Goto Description]. Holotype: BMNH 1909.6.12.7: ad ♂, skull and alcoholic. Original number: 613. Collected and presented by Dr. and Mrs. Hinde, (see Thomas (1909a: 99). - Etymology: In honour of Mrs. Hildegarde Beatrice Hinde, wife of the Dr. Sydney Hinde, collector of the type specimen (see Colket and Wilson, 1998: 3; Lanza et al., 2015: 213). ? Taphozous (Taphozous) hildegardeae: (Name Combination) ? Taphozous hidegardeae: (Alternate Spelling) ? Taphozous hildegardeae: (Current Combination, Current Spelling) TAXONOMY: See Colket and Wilson (1998, Mammalian Species, 597). COMMON NAMES: Chinese: 肯 尼 亚 墓 蝠 . Czech: hrobkovec východoafrický. English: Hildegarde's Tomb Bat. French: Taphien de Hildegarde. German: Hildegardes Grabfledermaus. Italian: Tafozòo di Hildegàrde. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B1ab(iii) ver 3.1 (2001)) because its extent of occurrence is less than 20,000 km2, its distribution is severely fragmented, and there is continuing decline in the extent and quality of its forest habitat (Mickleburgh et al., 2008y; IUCN, 2009). Assessment History Global 2008: VU B1ab(iii) ver 3.1 (2001) (Mickleburgh et al., 2008y; IUCN, 2009). 2004: VU C1 ver 3.1 (2001) (Mickleburgh et al., 2004r; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: Disturbance of the caves on which it depends, and loss of forest habitat, could result in population declines (Mickleburgh et al., 2008y; IUCN, 2009). Sagot and Chaverri (2015: 1670) mention roost loss or disturbance, and habitat degradation or loss as major threats for this species. CONSERVATION ACTIONS: Mickleburgh et al. (2008y) [in IUCN (2009)] report that it probably occurs in some protected areas, but stricter conservation of roosting and nesting sites is needed. Research into the cultural importance of the species is being carried out. GENERAL DISTRIBUTION: Taphozous hildegardeae is found on Pemba and Unguja [=Zanzibar] islands (Tanzania), and along the coast of south-east Kenya and north-east Tanzania (from the Lower Tana River south to Dar es Salaam). It has so far been recorded from fewer than ten coastal localities, but is probably present in some places where it has not yet been discovered. 414 ISSN 1990-6471 The two central Kenya records collected by Harrison (1962), are questioned due to their location by Kock (1974a), as these points are not along the East African coast. The questioned locations do, however, coincide with coastal vegetation along the Tana River, which could afford this species suitable habitat (Colket and Wilson, 1998). Native: Kenya; Tanzania. ROOST: McWilliam (1982: 208) found T. hildegardeae roosting in the larger caves of his study area (coastal Kenya), where they were found in smaller chambers or "erosion domes". DIET: In Kenya, McWilliam (1982: 248) reported that T. hildegardeae almost exclusively fed on large (> 10 mm) Lepidoptera and Orthoptera. not result in reproduction, indicating that the male cycle might be a relict of ancestral polyoestry. MATING: T. hildegardeae has a polygonous mating system, where a successful male can father at least seven young per year (McWilliam, 1982: 258). PARASITES: HEMIPTERA Cimicidae: Loxaspis miranda Rothschild, 1912, described from specimens collected at Kilidini,near Mombasa, Kenya, Haeselbarth et al. (1966: 9) suggests that these are from T. hildegardeae. DIPTERA Streblidae: Brachytarsina alluaudi (Falcoz, 1923) in Kenya and probably Tanzania (Haeselbarth et al., 1966: 101). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, Tanzania. POPULATION: Structure and Density:- There are no overall population estimates. However, during a survey in Kenya and Tanzania in 1988, population estimates from three caves surveyed yielded more than 1,500 individuals (1,000+, 300+, 130+) (Susan M. Pont, pers. Obs., in Mickleburgh et al., 2008y [in IUCN, 2009]). It seems to be relatively common in caves. Trend:- 2008: Decreasing (Mickleburgh et al., 2008y; IUCN, 2009). REPRODUCTION AND ONTOGENY: For Kenya, McWilliam (1988b: 433) indicates that male T. hildegardeae have a bimodal reproductive cycle that corresponds to the rainfall-induced seasonality in the coastal area. He also mentioned, however, that the second period (during the short rains in October-December), did Figure 137. Distribution of Taphozous hildegardeae Taphozous mauritianus E. Geoffroy St.-Hilaire, 1818 *1818. Taphozous mauritianus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 127. Publication date: 1818. Type locality: Mauritius: "Mauritius". Holotype: MNHN A.381 - 199: ad ♀, alcoholic (skull not removed). Rode (1941: 243). Comments: Mahoney and Walton (1988a: 116) in Grubb et al. (1998: 72) considered the date of Geoffroy's publication to be 1813. Gardner and Hayssen (2004: 12) mention that the correct date should be 1818, not 1813 as mentioned on the publication itself. Etymology: From the masculine scientific Latin adjective mauritiànus, meaning "Maurician", as the type specimen was collected on the island of Mauritius (see Lanza et al., 2015: 215). (Current Combination) 1835. Taphozous leucopterus Temminck, Monographies de mammalogie ou description de quelques genres de mammifères, dont les espèces ont été observées dans les différens musées de l'Europe. Tome Seconde, 2: 284, pl. 60, fig. 7. Type locality: South Africa: "Interior of South Africa" [Goto Description]. - Comments: Kock (1969a: 71), Meester et African Chiroptera Report 2020 1879. 1900. 1978. 2017. 2020. ? ? 415 al. (1986: 31) and Dengis (1996: 1) mention "1838; Tijdschrift voor natuurlijke geschiedenis en physiologie, Amsterdam 5: 12". Taphozous Dobsoni Jentink, Notes Leyden Mus., 1 (1): 123, note 32. Publication date: April 1879. Type locality: Madagascar: Mohambo [Goto Description]. Holotype: RMNH ?: ♂, alcoholic (skull not removed). Collected by: Jean Baptiste Audebert. Taphozous maritianus var. Cinerascens: Seabra, J. Sci. mat. phys. nat., ser. 2, 6: 77. Publication date: August 1900. Type locality: Angola: Benguela [12 34 S 13 24 E, 160 m]. (Lapsus) Taphozous mauvitanius: Arata and Johnson, Ebola Virus Haemorrhagic Fever, 135. (Lapsus) T[aphozous] mauritiana: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 5. Publication date: 6 April 2017. Taphozous mauritianum: Sow, Seye, Faye, Benoit, Galan, Haran and Brévault, Crop Prot., 132 (105127): 4. Publication date: 5 March 2020. (Lapsus) Taphozous (Taphozous) mauritianus: (Name Combination) Taphozous mauritianus mauritianus: (Name Combination) TAXONOMY: PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Evans and Mourad (2018: 81; Fig. 4, 5) provide images of wall paintings of bats, which look like Taphozous, on the walls of an Egyptian tomb at Beni Hassan, dating back from the Middle Kingdom (c. 2050 - 1650 BC). Figure 138. Taphozous mauritianus caught at Sudwala Caves Resort, Mpumalanga, South Africa. Koopman (1975) believed that a number of subspecies that have been described are of doubtful value. See Dengis (1996, Mammalian Species, 522). COMMON NAMES: Afrikaans: Witlyfvlermuis. Azande (DRC): Fulo. Castilian (Spain): Murciélago de las Tumbas. Chinese: 南 非 墓 蝠 . Czech: hrobkovec mauricijský, netopýr mauricijský, netopýr hrobkový. English: Mauritian Tomb Bat, Tomb Bat, Taphozous bat of Mauritius. French: Taphien de l'Ile Maurice, Taphien de l'Ile de France, Taphien de Maurice, Chauve-souris mauritienne des tombeaux, Chauve-souris des tombes. German: Mauritius-Grabfledermaus. Italian: Tafozòo mauriziàno. Kinyarwanda: Mokombe. Kiluba (DRC): Katutu. Kinande (DRC): Kahazo-komule, Kakorokombe. Portuguese: Morcego das sepulturas das Mauricias. Swahili: Popo. ETYMOLOGY OF COMMON NAME: The specimen from which the species was described came from the island of Mauritius (see Taylor, 2005: 274). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Hutson et al., 2008f; IUCN, 2009; Monadjem et al., 2017bb). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bb). 2008: LC ver 3.1 (2001) (Hutson et al., 2008f; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bl; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016u). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Overall, this species is not significantly threatened. It is locally hunted (by children) in parts of its range (Hutson et al., 2008f; IUCN, 2009; Monadjem et al., 2017bb). CONSERVATION ACTIONS: Hutson et al. (2008f) [in IUCN (2009)] and Monadjem et al. (2017bb) report that this species is known from protected areas within Madagascar including Zombitse National Park and 416 ISSN 1990-6471 Ankarafantsika National Park, as well as from many protected areas on the African continent. GENERAL DISTRIBUTION: Taphozous mauritianus is found throughout much of subsaharan Africa and is also on several islands (including the Tanzanian island of Unguja [=Zanzibar] [see O'Brien, 2011: 287]), Bioko and Pagalu (Equatorial Guinea), and Sao Tomé and Principe). The most northerly records are from the border area between Senegal and Mauritania and at the head of the Nile River in Sudan. It is present throughout southern and eastern Madagascar, but it usually only occurs at lower altitudes and there are no records from the central plateau or above 900 m. It is known from Aldabra (von Brandis (2004: 135), Mauritius, the Comoros and Réunion. Records from the Western Cape region in southern Africa are unverified, with known records extending as far south as Mossel Bay. It is also recorded in an isolated location in the Northern Cape, near Hartwater (Taylor, 2000; Skinner and Chimimba, 2005). In the RSA, the species' distribution is linked with land use/land cover (Babiker Salata, 2012: 50). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,454,841 km2. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 532; Taylor et al., 2018b: 62); Benin (Capo-Chichi et al., 2004: 162); Botswana (Archer, 1977; Monadjem et al., 2010d: 535); Cameroon; Central African Republic; Chad; Comoros (Louette, 2004; Goodman et al., 2010c: 125; Hume and Middleton, 2011: 33); Congo (Bates et al., 2013: 335); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 535); Côte d'Ivoire; Equatorial Guinea (Annobón, Bioko); Ethiopia; Gabon; Gambia (Emms and Barnett, 2005: 50); Ghana; Kenya; Madagascar; Malawi (Happold et al., 1988; Ansell and Dowsett, 1988: 29; Monadjem et al., 2010d: 535); Mauritius; Mayotte (Ramasindrazana et al., 2015a: 109); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 535); Namibia (Monadjem et al., 2010d: 535); Nigeria; Réunion; São Tomé and Principe (Juste and Ibáñez, 1993b: 906; Rainho et al., 2010a: 35); Senegal; Seychelles [Aldabra]; Sierra Leone; Somalia; South Africa (Monadjem et al., 2010d: 535); Sudan; Swaziland (Monadjem et al., 2010d: 535); Tanzania; Togo; Uganda; Zambia (Ansell, 1967; Ansell, 1969; Ansell, 1978;Ansell, 1986; Cotterill, 2004a: 260; Monadjem et al., 2010d: 535); Zimbabwe (Monadjem et al., 2010d: 535). Presence uncertain: Burundi; Guinea; GuineaBissau; Liberia; Mali; Mauritania; Rwanda. GEOGRAPHIC VARIATION: Krutzsch (2000: 114) [referring to Rosevear (1965), Smithers and Lobão Tello (1976) and Happold (1987)] indicates that geographic variation is evident in the gular sac in this species. In Nigeria and Mozambique, it is only present in males, while in West Africa the males have a functioning sac and females have a vestigial pouch. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Fenton (1992: 49) indicates that the grizzled dorsal pelage contributes to the bat's camouflage while resting on natural surfaces. In the resting position, the striking white venter is completely concealed. From western Uganda, Monadjem et al. (2011: 30) reported the following data for 2 specimens: Fa: 60.25 mm, mass: 27.0 g, wing loading: 13.3 N/m 2, aspect ratio: 5.8. ECHOLOCATION: See Taylor (1999b), and Taylor (2000: 63). Rainho et al. (2010a: 21) report the calls of 5 individuals from São Tomé. For 10 calls of specimens from Soutpansberg and Waterberg, South Africa, Taylor et al. (2013b: 18) recorded the following parameters: Fmax: 29.0 ± 1.17 (27.8 - 31.1) kHz, Fmin: 25.3 ± 1.49 (22.1 27.9) kHz, Fknee: 28.3 ± 1.29 (27.0 - 30.2) kHz, Fchar: 26.2 ± 0.94 (24.9 - 27.9) kHz, duration: 2.9 ± 0.64 (2.0 - 3.6) msec. Weier et al. (2020: Suppl.) reported on seven calls from the Okavango River Basin with the following characteristics: Fmax: 28.71 ± 2.38 kHz, Fmin: 23.88 ± 2.10 kHz, Fknee: 26.96 ± 1.67 kHz, Fchar: 24.96 ± 1.72 kHz, slope: 79.41 ± 58.64 Sc, duration: 3.06 ± 0.94 msec. From western Uganda, Monadjem et al. (2011: 32) report the following values: Fmin: 13.13 kHz, Fmax: 13.48 kHz, Fchar: 13.43 kHz, Fknee: 13.18 kHz, duration: 9.1 msec. They also indicate (p. 33) that the Fpeak of 25.6 [actually 25.9] kHz reported by Schoeman and Jacobs (2008: Table S1) is probably for an harmonic. Barataud and Giosa (2013: 150) reported on 9 sequences from Réunion with QFC = 39 kHz and QCF/FM = 18. For Swaziland, Monadjem et al. (2017c: 179) recorded the following parameters: Fmin: 25.1 ± 0.72 (24.3 - 25.9) kHz, Fknee: 28.6 ± 0.60 (28.1 29.5) kHz, Fc: 25.9 ± 1.20 (24.3 - 27.2) kHz, and duration: 2.7 ± 0.27 (2.4 - 3.0) kHz. African Chiroptera Report 2020 Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 25.9 kHz and duration: 7.4 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported for specimens from South Africa, 2n = 42, FN = 64, BA = 24, a metacentric X chromosome, and a submetacentric Y chromosome. Protein / allozyme - Unknown. HABITAT: Open savanna and forst edges in the wetter northern and eastern parts of the subcontinent (Taylor, 2000: 62). HABITS: Fenton and Griffin (1997: 248) recorded one call above 24 kHz, which they attributed to T. mauritianus at an altitude of 500 - 550 m above a Brachystegia woodland in Zimbabwe. ROOST: In the Durban area, Taylor et al. (1999: 67) found most of the roosts (72 %) being associated with the outer walls of houses at a height between 2 and 12 metres. The colony size varied from 1 to 5. In a rural area on Madagascar; Rakotondramanana et al. (2017: 133) found a colony roosting on the trunk of a palm tree (Cocos nucifera), under dried fronds. They left the roost about 30 minutes after sunset. Ravelomanantsoa et al. (2019: 110) report it also roosting in Tamarindus indica. DIET: They feed primarily on moths (Allen et al., 1917); termite alates such as Macrotermes falciger (Happold and Happold, 1988), and other insects (Smithers, 1971). See also (Taylor, 2000: 62; 2005: 275). In Senegal, Sow et al. (2020: 4) used molecular analyses to identify the presence of Heliocheilus albipunctella de Joannis) (Lepidoptera, Noctuidae) in the faeces of T. mauritianus. PREDATORS: Demeler (1981: 133) found remains of T. mauritianus in pellets of Bubo africanus in the Yankari Game Reserve (Nigeria). Mikula et al. (2016: Supplemental data) mention Wahlberg's eagle (Hieraaetus wahlbergi (Sundevall, 1851)) as a diurnal avian predator. 417 Goodman et al. (2015c: 78) found the remains of one individual in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. POPULATION: Structure and Density:- Taphozous mauritianus is not rare and is easy to find. It usually roosts under covering vegetation on the outer bark of trees (Hutson et al., 2008f; IUCN, 2009; Monadjem et al., 2017bb). Trend:- 2016: Unknown (Monadjem et al., 2017bb). 2008: Unknown (Hutson et al., 2008f; IUCN, 2009). REPRODUCTION AND ONTOGENY: Kaudern (1914: 12) reported that he received a female with a very small fetus from villagers in Fenerive (Madagascar) on 6 January 1912. Krutzsch (2000: 113) mentioned that T. mauritianus is monoestrous in parts of its range and polyoestrous in others. Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] mentioned births from October to December and f'rom February to March. In Malawi, births were reported in NovemberDecember and March-April (Happold and Happold, 1990b: 564). PARASITES: HEMIPTERA Polyctenidae: Eoctenes intermedius (Speiser, 1904) from the Congo (Haeselbarth et al., 1966: 16). DIPTERA Nycteribiidae: Tripselia blainvillii (Leach, 1817) distributed over the Madagascan region and tropical Africa from west to east between lat. 10o north and south (Haeselbarth et al., 1966: 113). Hutson (2004a: 129) mentions that Basilia (Tripselia) blainvilli (Nycteribiidae) is a widespread parasite of Taphozous mauritianus. (synonyms: Nycteribia (Acrocholidia) fryeri, Tripselia fryeri). SIPHONAPTERA Ischnopsyllidae: Chiropteropsylla brockmani Rothschild, 1915 widespread (Haeselbarth et al., 1966: 192). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Taphozous mauritianus: Astroviruses, Paramyxoviruses and Rotaviruses. 418 ISSN 1990-6471 Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in one individual from Limpopo Province, South Africa, none were tested positive (0/1). In their overview table, Maganga et al. (2014a: 8) reported that no viruses were ever found in this bat species. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burundi, Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Eswatini, Ethiopia, Gabon, Ghana, Guinea-Bissau, Kenya, Madagascar, Malawi, Mauritius, Mozambique, Namibia, Nigeria, Réunion, Rwanda, Senegal, Sierra Leone, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. Figure 139. Distribution of Taphozous mauritianus Taphozous nudiventris Cretzschmar, 1830 *1830-1831. Taphozous nudiventris Cretzschmar, in: Rüppell, Atlas Reise Nördlichen Afrika, Zoologie Säugethiere, 1: 70, pl. 27b. Publication date: 1830-1831. Type locality: Egypt: Giza province: Giza [30 01 N 31 13 E] [Goto Description]. Lectotype: SMF 4310: ad ♂, skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell; collection date: 1824. Old catalog II.J.1.a; see Mertens (1925: 20), Qumsiyeh (1985: 26). - Comments: Kock (1969a: 83) mentioned the type locality as "Egypt, Nubia" and indicates that Giza is the type locality of the lectotype. Corbet and Hill (1992: 87) only mention "Egypt". Allen (1939a: 67) mentioned 1826 as year of description, whereas Kock (1969a: 83) mentioned 1830. Grubb et al. (1998: 73) mentions "1830, vel 1831". - Etymology: From the masculine Latin adjective nùdus, meaning "naked" and the masculine Latin substantive vènter, meaning"belly", as the fur stops short of the end of the body both dorsally and ventrally (see Lanza et al., 2015: 221). (Current Combination) 1838. Taphozous nudiventer: Temminck, Monographies de mammalogie ou description de quelques genres de mammifères, dont les espèces ont été observées dans les différens musées de l'Europe. Tome Seconde, 2: 280, pl. 60, figs 10 - 12. Publication date: 1838. - Comments: Lapsus (Allen, 1939a: 67). . (Lapsus) 1877. Nycticejus serratus Heuglin, Reise in Nordost Afrika, 2: 35. Publication date: 1877. Type locality: Sudan: ?Abyssinia: on the Araschkol and nearby mountains: "Sherq el Aqabah" [14 15 N 32 10 E] [Goto Description]. - Comments: According to Simmons (2005) this is an enigmatic taxon, which is variously referred to either Taphozous nudiventris (e.g. Allen, 1939a: 67; Koopman, 1993a) or Scotophilus nigrita leucogaster (Allen, 1939a) [implying that it might either be S. dinganii or S. leucogaster], but which may not represent either of those species. Kock (1969a: 83) includes it with a question mark in the synonymy of Taphozous nudiventris, and Happold (2013bm: 434) mentions it as a possible synonym of that species. 1885. Taphozous perforatus var. assabensis Monticelli, Ann. Accad. O. Costa de Aspir. Nat. Napoli, ser. 1, 1: 78, pl. 7. Type locality: Eritrea: Assab [13 01 N 42 43 E]. - Comments: Allen (1939a: 67) indicates that Monticelli considered it a synonym of perforatus, but Senna (1905), who examined the type, says it is T. nudiventris. The type specimen was collected by prof. G.B. Licata. 1964. T(aphozous) mudiuemtris: Walker, Mammals of the World, I: 241. (Lapsus) 2013. Taphozous nudiuentris: Liu, Wang, Zhang, Li, He, Huang, Jiang, Murphy and Shi, Proc. R. Soc. Lond., B 281 (1776): 20132950 3 (for 2014). Publication date: 18 December 2013. (Lapsus) ? Liponycteris nudiventris: (Name Combination) African Chiroptera Report 2020 ? ? 419 Taphozous (Liponycteris) nudiventris: (Name Combination) Taphozous nudiventris nudiventris: (Name Combination) TAXONOMY: Simmons (2005) assigns four subspecies to Taphozous nudiventris: T. n. kachhensis Dobson, 1872; T. n. magnus Wettstein 1913; T. n. nudaster Thomas, 1915; and T. n. zayidi Harrison, 1955. Uvizl et al. (2019: 31) consider zayidi to be a junior synonym of T. n. nudiventris, and also indicate that there is need for further investigation to ascertain the position of the other subspecies. The enigmatic taxon T. n. serratus Heuglin, 1877 has been variously referred to either Taphozous nudiventris (e.g., Allen, 1939a, Koopman, 1993a) or Scotophilus leucogaster (e.g., Allen, 1939a; Koopman, 1993a), although it might not represent either of these species. See Felten (1962), Hayman and Hill (1971), Harrison and Bates (1991), Bates and Harrison (1997) and Barkley (1984) for further information on this taxon. COMMON NAMES: Albanian: Pipistreli i varrezave barkzhveshur. Arabian: Khafash Abu Bouz, Khaffash. Armenian: Մերկփոր պարկաթև. Azerbaijani: Ilpaqqarın məzar yarasası. Basque: Hilobisaguzar gibelsoil. Belarusian: Магільнік галабрухі. Bosnian: Golotrbušni grobni šišmiš. Breton: Logodenn-dall kof noazh ar bezioù. Bulgarian: Голокоремен прилеп. Castilian (Spain): Murciélago de cola libre y vientre desnudo. Catalan (Spain): Ratpenat de tomba de carpó nu. Chinese: 裸 腹 墓 蝠 . "Congolese": Kubukubo, Papo. Croatian: Golotrbi grobni šišmiš. Czech: Netopýr lysobřichý, Hrobkovec lysobřichý. Danish: Nøgenrumpet gravflagermus. Dutch: Kaalbuikgrafvleermuis. English: Naked-bellied Tomb Bat, Naked-rumped Tomb Bat, Egyptian sheath-tailed bat, Nakedrumped bat, Naked-bellied Taphozous. Estonian: Paljaskõht-kääpanahkhiir. Finnish: Arohautalepakko. French: Taphien à croupe nue, Taphien à ventre nu. Frisian: Kealbûktimpelflearmûs. Galician (Spain): Morcego de tumba de cú espido. Georgian: შიშველმუცელა ქარქაშკუდა. German: Nacktbäuchige Tempelfledermaus, NacktbauchGrabfledermaus. Greek: Άτριχη ταφονυχτερίδα. Hebrew: Ashman Gadol. Hungarian: Csupaszhasú sírlakó-denevér. Irish Gaelic: Ialtóg nochtphrompach thuama. Italian: Pipistrello delle tombe a ventre nudo, Tafozòo nudivèntre. Latvian: KailvĒdera sikspārnis. Lithuanian: Plikapilvis maišiasparnis. Luxembourgish: Plakegbauchege Griewerflatterer. Macedonian: Торбокрилест лилјак. Maltese: Farfett il-Lejl Għieri. Montenegrin: Golotrbušni grobni slijepi miš. Norwegian: Nakengumpflaggermus. Polish: Grobownik gołobrzuchy. Portuguese: Morcegode-ventre-nú. Rhaeto-Romance: Pipistrel da fossa a venter niv. Romanian: Liliacul cu burtă golaşa. Russian: Мешкокрыл голобрюхий. Serbian: Голотрби гробнички м[j]ешкокрилаш [=Golotrbi grobnički m[j]eškokrilaš]. Scottish Gaelic: Ialtag nochd-thònach thuaim. Slovak: Hladkonos holobruchý. Slovenian: Golotrebušni mavzolejnik. Swedish: Nakengumpsfladdermus. Tamil: திறந்த ரம் ப் டு கல் லறற வெளொல் , Iṟanta Rampṭu Kallaṟai Veḷāl. Turkish: Çıplakkarınlı Mezar Yarasası. Ukrainian: Мішкокрил голобрюхий. Welsh: Ystlum beddrod tinfoel. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Bates et al., 2008; IUCN, 2009; Monadjem et al., 2017bq). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bq). 2008: LC ver 3.1 (2001) (Bates et al., 2008; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bp; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: This species is tolerant of a certain level of human disturbance. Loss of some roosts (e.g., in buildings) and use of pesticides probably impact some populations negatively, but overall it is not significantly threatened (Bates et al., 2008; IUCN, 2009; Monadjem et al., 2017bq). CONSERVATION ACTIONS: Bates et al. (2008) [in IUCN (2009)] and Monadjem et al. (2017bq) report that there are international legal obligations for protection through the Bonn Convention (Eurobats) in areas to which this applies. However, through most of its range, no specific conservation measures are place for this species. It occurs in many protected areas across its wide range. A study on the impacts of 420 ISSN 1990-6471 pesticides is required, especially ways in which the impact might be minimised. GENERAL DISTRIBUTION: Taphozous nudiventris has a much larger range than previously believed. It has been recorded throughout the southern desert and sub-desert belt of western and central Palaearctic, from Morocco, through the Saharan region across northern Africa to Egypt and north through the Middle East to southern Turkey, and the more arid areas of the Indian subcontinent. The most southerly record is from northern Tanzania. There are two isolated records from Myanmar (the southernmost locality being in the general vicinity of Bago (Pegu) Yoma (Bates et al., 2000)). In South Asia this species is presently known from Afghanistan (Kabul, Kandahar and Nangarhar provinces), Bangladesh, India (Andhra Pradesh, Bihar, Delhi, Gujarat, Karnataka, Madhya Pradesh, Maharashtra, Rajasthan, Sikkim, Tamil Nadu, Uttar Pradesh and West Bengal) and Pakistan (Punjab and Sind) (Molur et al., 2002). It has also been recorded from the Cape Verdian islands (Tranier and de Naurois, 1985: 304) Native: Afghanistan; Algeria (Horácek et al., 2000: 104); Burkina Faso (Kangoyé et al., 2015a: 614); Chad; Congo (The Democratic Republic of the); Djibouti (Pearch et al., 2001: 396); Egypt; Eritrea (Lavrenchenko et al., 2004b: 138); Ghana; Guinea-Bissau (Castella et al., 2000; Rainho and Ranco, 2001: 38); India; Iran, Islamic Republic of; Iraq; Israel; Jordan; Kenya; Macronesian Islands (Cape Verd islands (Tranier and de Naurois, 1985; de Naurois, 1994; Hazevoet, 1995; Masseti, 2010); Mauritania; Morocco; Myanmar; Niger; Nigeria; Pakistan; Palestinian Territory, Occupied; Saudi Arabia; Senegal; Somalia; Sudan; Syrian Arab Republic; Tanzania; Togo; Turkey; United Arab Emirates (Anonymus, 2008: 38); Yemen. Presence uncertain: Ethiopia; Kuwait; Libyan Arab Jamahiriya; Oman; Qatar; Tunisia; Uganda; Western Sahara. The records from Lebanon were revoked by Benda and Engelberger (2016: 17). ECHOLOCATION: In Jordan, Benda et al. (2010b: 204) registered the following parameter for 17 calls: Fstart: 26.1 ± 1.0 (24.4 - 27.6) kHz, Fend: 22.2 ± 0.6 (20.3 - 22.9) kHz, Fpeak: 24.5 ± 0.5 (23.6 - 25.3) kHz, Pulse duration: 15.4 ± 2.6 (11.0 - 19.5) msec, and Inter-pulse interval: 363.9 ± 115.1 (188.0 - 571.0) msec. Luo et al. (2019a: Supp.) reported the following data (Free flying bats): F peak: 23.38 kHz, Fstart: 24.39 kHz, Fend: 22.8 kHz, Band width: 1.59 kHz, and duration: 12.99 msec. From Israel, Hackett et al. (2016: 223) reported the following parameters for 74 calls: Pulse duration: 12.99 ± 3.50 msec, Fstart: 24.39 ± 1.55 (22.3 - 25.3) kHz, Fend: 22.80 ± 1.29 (20.4 - 26.3) kHz, Fpeak: 23.38 ± 1.31 (21.3 - 26.3) kHz. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Ðulic (1984), Yaseen et al. (1994), Hood and Baker (1986), Asan and Albayrak (2007) reported 2n = 42, FN = 64, BA = 24. However, Robbins (1983b: 35) reported 2n = 42 and FN = 68 and Sreepada et al. (1995) 2n = 42, FN = 66, BA = 26 for specimens from India. In Turkey, Karatas and Sözen (2003: 359) and Arslan and Zima (2014: 7) reported 2n = 42, FNa = 64, FN = 68, a meta-/submetacentric X and an acrocentric Y chromosome. The autosomal chromosomes consist of 12 metacentric and submetacentric pairs and 8 acrocentric pairs. Protein / allozyme - Unknown. HABITS: Lucan and Šálek (2012: 235) report on the mobbing behaviour of T. nudiventris on a barn owl (Tyto alba) in the Dakhla Oasis in Egypt. The bats were able to detect the owl when it was on route to their roost, but it could not be determined how the bats were able to do this. There were no alarm or distress calls from other bats, so possibly visual cues or acoustic signals (e.g. sounds made by the feathers of the flying owl) might be involved. ROOST: Davis (2007: 10) found that in the United Arab Emirates, they may roost in groups of 200 to 1,000+ individuals. During late pregnancy and lactation, the females form separate groups. DIET: Benda et al. (2012a: 282) report that the most important food categories in Syria, Sudan, Oman and Turkey are Coleoptera (Scarabaeidae) and Orthoptera. Desai et al. (2012) [in Benda et al., 2012a: 282] found seasonal changes in the food composition of these bats in Gujarat (India): Coleoptera and Orthoptera prevailed in winter, while Isoptera, Lepidoptera, Orthoptera, Neuroptera and Hymenoptera (Vespoidea) were most consumed in summer. PREDATORS: In Algeria, Djilali et al. (2016: 161) found remains of 2 T. nudiventris specimens in 516 preys captured by the Short-eared Owl (Asio flammeus). African Chiroptera Report 2020 POPULATION: Structure and Density:- It is common in some places, and less so in others. It is an uncommon species in the western part of its range: colonies in Africa and the Mediterranean region are generally restricted to a few individuals, although large colonies (dozens to hundreds) have been found in eastern Africa (Bates et al., 2008; IUCN, 2009; Monadjem et al., 2017bq). The species is common in its range in South Asia, however, a declining trend in its population has been observed in recent years (Bates and Harrison, 1997). Trend:- 2016: Stable (Monadjem et al., 2017bq). 2008: Stable (Bates et al., 2008; IUCN, 2009). T. nudiventris populations within the Asia Minor and Levant region are predicted to have a stable population trend, under climate change scenarios (Bilgin et al., 2012: 433). REPRODUCTION AND ONTOGENY: Kurta and Kunz (1987: 82) report that, at birth, the young has a forearm length of about 34.4 % of that of the mother (27.5 against 80.0 mm). 421 as Cerná and Ryšavý (1976) did not provide any proof about the parasite, the identification might be wrong, given the fact that eimerians are reasonably host specific. HEMIPTERA Cimicidae: Leptocimex vespertilionis Ferris and Usinger, 1957 known only from the Sudan (Zeidab, Khartoum, Wad Medani) (Haeselbarth et al., 1966: 13). ACARINA Dermanyssidae: Zumpt and Till (1954: 49) mentions that Hirst (1926) described Steatonyssus sudanensis from a "Liponycteris nudiventris" from near Khartoum, Sudan, but that this mite hasn't been found since. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Burkina Faso, Cameroon, Cape Verde, Congo (Democratic Republic of the), Djibouti, Egypt, Ethiopia, Ghana, Kenya, Niger, Nigeria, Senegal, Somalia, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda. In the United Arab Emirates, Davis (2007: 10) indicates that they arrive in their summer roosts in March, and a single young is born in mid-April. Young are carried by their mothers until 8 weeks of age. In Gujarat (India), Desai et al. (2012: 33) found that pregnancy lasted for 87 days and parturition occurred in August/September. The young reached their full adult size in nine months, when females become sexually mature. The males only become mature at the age of 20 months. MATING: In Gujarat (India), Desai et al. (2012: 33) report that copulation and fertilization takes place during the last week of March and the first week of April. Figure 140. Distribution of Taphozous nudiventris PARASITES: Duszynski (2002: 20) indicates that T. nudiventris might be a host for Eimeria dukei Lavier, 1927, but Taphozous perforatus E. Geoffroy St.-Hilaire, 1818 *1818. Taphozous perforatus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 126. Publication date: 1818. Type locality: Egypt: Kom Ombo [24 26 N 32 57 E]. Holotype: MNHN A.372: ad. Presented/Donated by: ?: Collector Unknown. Comments: Allen (1939a: 66) mentioned "Egypt" only as type locality. Restricted to Ombos (=Kom Ombo), between Edfu and Aswan by Kock (1969a: 74): see Meester et al. (1986: 32). Mahoney and Walton (1988a: 116) in Grubb et al. (1998: 72) considered the date of Geoffroy's publication to be 1813. Gardner and Hayssen (2004: 12) mention that the correct date should be 1818, not 1813 as mentioned on the publication itself. Etymology: From the masculine scientific Latin adjective perforàtus, meaning "perforated", since the tail, as in all Emballonuridae, seems to perforate the uroparagium (see Lanza et al., 2015: 227). (Current Combination) 422 ISSN 1990-6471 1877. 1915. 2014. ? ? Taphozous maritimus Heuglin, Reise in Nordost Afrika, 2: 25. Publication date: 1877. Type locality: Sudan: Suakin [19 06 N 37 22 E]. Taphozous perforatus hædinus Thomas, J. Bomb. nat. Hist. Soc., 24: 62. Publication date: 30 September 1915. Type locality: Kenya: Northern Guaso Nyiro: Chanler Falls [=Chandler's Falls] [00 47 N 38 04 E, 2 000 ft]. Holotype: BMNH 1912.7.1.46: ad ♂, skin only. Collection date: 3 September 1911; original number: 756. Taphozous perforates: Kohl and Kurth, Viruses, 6 (8): 3115. Publication date: 13 August 2014. (Lapsus) Taphozous (Taphozous) perforatus: (Name Combination) Taphozous perforatus haedinus: (Name Combination) TAXONOMY: Harrison (1958b, 1962) regards sudani as a separate species, but is not followed in this by subsequent authors (Rosevear, 1965; Kock, 1969a; Hayman and Hill, 1971; Koopman, 1975). Hayman and Hill (1971) make no attempt to recognize subspecies within perforatus, but Kock (1969a) recognizes three, perforatus, sudani and haedinus. Koopman (1975) does not agree entirely with the geographic limits suggested by Kock, but appears to accept the validity of the names proposed. Simmons (2005) recognized three subspecies: haedinus Thomas, 1916; sudani Thomas, 1915; and senegalensis Desmarest, 1820. Uvizl et al. (2019: 30) could not find any genetic structuring to distinguish haedinus from perforatus and reject the recognition of subspecies in the Middle East and north-eastern Africa. However, they did not examine material from the type locality, nor from any other East African locality, and therefore do not draw any conclusions about the subspecific status of haedinus. Uvizl et al. (2019: 30) also indicate that T. perforatus is probably of Asian origin, and its occurrence in the Middle East and Africa is the result of a rapid westward expansion, which is illustrated by the lack of genetic structure across its populations. COMMON NAMES: Afrikaans: Egiptiese witlyfvlermuis. Arabian: Khafash abu bouz saghir, Khaffash. Chinese: 埃 及 墓 蝠 . Czech: hrobkovec egyptský, wečerník senegalský, wečerník prowrtaný, netopýr slujový. Dutch: Egyptische grafvleermuis. English: Geoffroy's Tomb Bat, Egyptian Tomb Bat, Tomb Bat, Perforated Taphozous bat, African Taphozous. French: Taphien perforé, Taphien des tombeaux, Chauve-souris des tombes egyptiennes. German: Ägyptische Grabfledermaus. Hebrew: Ashman Katan. Italian: Tafozòo perforàto. Portuguese: Morcego das sepulturas do egipto. ETYMOLOGY OF COMMON NAME: The specimen from which this species was described was taken by Geoffroy Saint-Hilaire from a royal tomb in Egypt (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Kock et al., 2008d; IUCN, 2009; Monadjem et al., 2017bo). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bo). 2008: LC ver 3.1 (2001) (Kock et al., 2008d; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004aa; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: NT D2 ver 3.1 (2001) (Richards et al., 2016b). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: Kock et al. (2008d) [in IUCN (2009)] and Monadjem et al. (2017bo) report that human disturbance has been highlighted as a threat to T. perforatus, but overall it is unlikely that this species is significantly threatened across its very wide range. In South Asia it is threatened by clearing of thorn forests for agricultural purposes, from mining and stone quarrying. Roost disturbance due to human interference and development of old buildings for tourism purposes is also considered serious threats (C. Srinivasulu pers. comm., Molur et al., 2002). In Egypt, Mansour et al. (2017: 109) found that liver and kidney tissues of these bats contained 0.39 µg/g wet weights of DDTs (its usage was prohibited in Egypt in 1980) and 0.11 µg/g wet weights of PCBs (banned worldwide in 1990s). These values showed variations between sexes and seasons. African Chiroptera Report 2020 CONSERVATION ACTIONS: Kock et al. (2008d) [in IUCN (2009)] and Monadjem et al. (2017bo) report that this species is present in many protected areas, and no direct conservation measures are currently needed for the species as a whole. The species has not been recorded from any protected areas in Pakistan or India. Being a poorly understood species in the region, more studies on distribution, abundance, reproduction and ecology are recommended (Molur et al., 2002). GENERAL DISTRIBUTION: Taphozous perforatus occurs widely throughout northern and sub-Saharan Africa, the Arabian Peninsula, and Asia, east to the Indian Subcontinent. In sub-Saharan Africa, records extend along the Nile and east to Ethiopia and northern Somalia, and west to Mauritania, Senegal, Gambia, Guinea-Bissau, Ghana, Burkina Faso, Benin, Niger, and northern Nigeria, and south to Kenya (including Lamu island), Tanzania, Democratic Republic of Congo, Zimbabwe, and Botswana. In southwest Asia, it has been recorded from Israel, Saudi Arabia, Yemen and Oman. In South Asia, this species is presently known to be widely distributed from southern Pakistan (Roberts, 1977) and western India (Gujarat, Madhya Pradesh and Rajasthan) (Bates and Harrison, 1997, Molur et al., 2002) and recently reported from central Eastern Ghats in Andhra Pradesh (Chakraborty et al., 2004). It has been recorded up to 200 m. Native: Benin; Botswana (Monadjem et al., 2010d: 535); Burkina Faso (Kangoyé et al., 2015a: 614); Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 535) ; Djibouti (Pearch et al., 2001: 394); Egypt (Benda et al., 2008f: 1); Ethiopia (Lavrenchenko et al., 2004b: 147); Gambia; Ghana; Guinea-Bissau (VeigaFerreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 37); India; Iran, Islamic Republic of; Israel; Kenya; Mali; Mauritania; Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 535); Niger; Nigeria; Oman; Pakistan; Saudi Arabia; Senegal; Somalia; Sudan; Tanzania; Togo; Uganda; Yemen (Benda et al., 2011b: 30); Zambia (Ansell, 1978; Monadjem et al., 2010d: 535); Zimbabwe (Monadjem et al., 2010d: 535). Presence uncertain: Angola; Côte d'Ivoire; Eritrea; Guinea; Namibia (Cotterill, 2004a: 260); Palestinian Territory, Occupied; United Arab Emirates; Zambia. DETAILED MORPHOLOGY: Tongue - El-Mansi et al. (2019: 24) identified two type of papillae: mechanical, subdivided into two 423 types of conicals (small and large), and filiform (pronged and crown-shaped) papillae; and gustatory, subdivided into vallate (containing about 10 ovoid taste buds) and fungiform types. ECHOLOCATION: In Jordan, Benda et al. (2010b: 204) registered the following parameters: Fstart: 31.4 ± 1.5 (29.6 - 34.4) kHz, Fend: 27.6 ± 0.6 (26.6 - 28.8) kHz, Fpeak: 29.2 ± 0.6 (28.5 - 30.0) kHz, duration: 12.4 ± 4.2 (5.8 19.7) msec, and interpulse interval: 225.1 ± 110.4 (58.0 - 426.0) msec. Two sets of data from Iran were presented by Benda et al. (2012a: 182): 1: Fstart: 31.4 ± 1.6 (30.0 - 34.2) kHz, Fend: 29.1 ± 1.7 (28.1 - 31.8) kHz, and Fpeak: 30.4 ± 2.1 (28.9 - 33.4) kHz; 2 (hand released): Fstart: 35.9 ± 0.5 (35.3 - 36.6) kHz, Fend: 29.9 ± 0.5 (29.4 - 30.8) kHz, Fpeak: 32.5 ± 0.3 (32.1 - 33.0) kHz, duration: 4.8 ± 0.3 (4.5 - 5.2) msec, and interpulse interval: 47.3 ± 26.8 (12.5 - 81.3) msec. Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 32.5 kHz, Fstart: 35.9 kHz, Fend: 29.9 kHz, Band width: 6 kHz, and duration: 4.8 msec. Götze et al. (2016: 2) mentioned that T. perforatus doesn't shift the frequency of their echolocation signals (jamming avoidance response - JAR) in the presence of conspecifics. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Ðulic (1984) and Yaseen et al. (1994) reported 2n = 42, FN = 64, BA = 24, a metacentric X chromosome and an acrocentric Y chromosome. Volleth et al. (2020b: 258, 260) reported 2n = 42 and FNa = 66, containing 12 bi-armed, one subtelocentric and seven acrocentric autosomal pairs, a medium-sized metacentric X and a Y chromosome with the same size as the smallest autosomal, which was too small to determine the shape. Protein / allozyme - Unknown. Mansour et al. (2016: 66) investigated the presence of heavy metals in bats from two caves in the Saqqara region of Egypt and found that cobalt was not detected in winter and spring, and at very low concentration in the other seasons (ca. 0.01 µg/g tissue). High concentrations were found for Fe, Ca, Mg, and to some extent also Al and Zn, whereas Ba, Cu and Mn were found at relatively lower concentrations, and Cr, Cd, Mo, Ni and Pb were detected at very low concentrations. Ba, Cd, Co, Cr, Mo, Ni and Pb were not detected in kidney tissue. Kidneys of males showed higher concentrations of Mg than those of females. 424 ISSN 1990-6471 DIET: Rydell and Yalden (1997: 72) report that T. perforatus mainly feeds on Lepidoptera (moths 56 % by volume), but also that Isoptera (14 %), Coleoptera (10 %) and Orthoptera (8 %) are major food categories. Furthermore, Hemiptera, Neuroptera, Hymenoptera and Diptera are eaten. PREDATORS: Mikula et al. (2016: Supplemental data) mention the Fox kestrel (Falco alopex (Heuglin, 1861)) and the Lanner falcon (Falco biarmicus Temminck, 1825) as avian predators. POPULATION: Structure and Density:- It is common in parts of its African range, but is less common elsewhere. It is found in small colonies (between six to eight individuals) in the southern African subregion (Skinner and Chimimba, 2005). In South Asia the abundance, population size and trends for this species are not known, and the species has only been recorded from a few localities (Bates and Harrison, 1997). Trend:- 2016: Stable (Monadjem et al., 2017bo). 2008: Stable (Kock et al., 2008d; IUCN, 2009). REPRODUCTION AND ONTOGENY: Krutzsch (2000: 113) indicates that T. perforatus is a polyoestrous species, although he also mentions (lower on the same page) that it is monoestrous in parts of its range and polyoestrous in others. viruses: Bat astrovirus, Bat Coronavirus, Bat mastadenovirus, Bat papillomavirus, Chuvirus, Rotavirus. Bunyaviridae Willoughby et al. (2017: Suppl.) report the presence of Kaeng Khoi orthobunyavirus. Coronaviridae - Coronaviruses Gortazar and Segalés (2013: 954) indicate that recently a virus was identified from a SaudiArabian T. perforatus specimen that showed 100 % nucleotide identity with MERS-CoV (Middle East Respiratory Syndrome) in a conserved portion of its CoV genome, suggesting that this species might be the host for the MERS-CoV virus. Flaviviridae Flavivirus Dakar bat virus was isolated from the pooled bat organs and salivary glands of this species in 1974 (Williams et al., 1964; Kemp et al., 1974). Reported in this species by de Jong et al. (2011: 10), Luis et al. (2013: Suppl.), Blitvich and Firth (2017: 3), Willoughby et al. (2017: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Botswana, Burkina Faso, Cameroon, Congo (Democratic Republic of the), Djibouti, Egypt, Ethiopia, Ghana, Kenya, Mauritania, Niger, Nigeria, Senegal, South Africa, South Sudan, Sudan, Tanzania, Togo, Uganda, Zambia, Zimbabwe. PARASITES: HAEMOSPORIDA Nycteria medusiformis Garnham and Heisch, 1953 was reported from various countries by Perkins and Schaer (2016: Suppl.). DIPTERA From Iran, Benda et al. (2012a: 275) report on the presence of Raymondia huberi Frauenfeld, 1855 (Streblidae). They also indicate that this bat fly occurs in a broad area from East Africa to the Middle East. Other parasites reported by Benda et al. (2012a: 276) from the north African part of the bat's range are: Argas boueti Roubaud et Colas-Belcour, 1933, Chiropteropsylla aegyptia (Rothschild, 1903), and C. brockmani Rothschild, 1915. VIRUSES: In bats from Saudi Arabia, Mishra et al. (2019: 5) found evidence for the presence of the following Figure 141. Distribution of Taphozous perforatus African Chiroptera Report 2020 425 Taphozous perforatus perforatus E. Geoffroy St.-Hilaire, 1818 *1818. Taphozous perforatus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 126. Publication date: 1818. Type locality: Egypt: Kom Ombo [24 26 N 32 57 E]. Holotype: MNHN A.372: ad. Presented/Donated by: ?: Collector Unknown. Comments: Allen (1939a: 66) mentioned "Egypt" only as type locality. Restricted to Ombos (=Kom Ombo), between Edfu and Aswan by Kock (1969a: 74): see Meester et al. (1986: 32). Mahoney and Walton (1988a: 116) in Grubb et al. (1998: 72) considered the date of Geoffroy's publication to be 1813. Gardner and Hayssen (2004: 12) mention that the correct date should be 1818, not 1813 as mentioned on the publication itself. Etymology: From the masculine scientific Latin adjective perforàtus, meaning "perforated", since the tail, as in all Emballonuridae, seems to perforate the uroparagium (see Lanza et al., 2015: 227). (Current Combination) 1820. Taphozous senegalensis Desmarest, Encyclopédie Méthodique, (Zoologie, Mammalogie), 1: 130. Publication date: 1820. Type locality: Senegal: "Senegal". - Comments: Based on "le lérot-volant" of Daubenton, Mém. Acad. Paris, p. 366, 1759 (see Allen, 1939a: 66). 1958. Taphozous perforatus swirae Harrison, Durban Mus. Novit., 5 (11): 144. Publication date: 15 December 1958. Type locality: Nigeria: N Nigeria: Sokoto [13 02 N 05 16 E]. Holotype: BMNH 1967.1226: ad, skin and skull. Collected by: Mrs. Pamela Swire; collection date: 26 August 1958. Skin made up by D.L. Harrison, formerly in HZM collection 12.2690. - Comments: Grubb et al. (1998: 72) mention the locality as Sokoyo. Considered a valid subspecies by Grubb et al. (1998: 73), but see Kock (1969a: 80 - 81). ? Taphozous perforatus perforatus: (Name Combination, Current Combination) DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burkina Faso, Egypt, Ethiopia, Ghana, Kenya, Mauritania, Niger, Nigeria, Senegal, South Sudan, Sudan, Tanzania, Uganda. Taphozous perforatus sudani Thomas, 1915 *1915. Taphozous Sudani Thomas, Ann. Mag. nat. Hist., 8, 15 (90): 561. Publication date: 1 June 1915. Type locality: Sudan: Equatoria Province: Upper Nile, N of Lado: Mongalla [05 10 N 31 50 E, 500 m] [Goto Description]. Holotype: BMNH 1902.7.4.2: ad ♂. Collected by: W.L.S. Loat Esq. Presented/Donated by: W.L.S. Loat Esq. - Etymology: Referring to the country where the type specimen was collected. 1962. Taphozous sudani australis Harrison, Occ. Pap. Nat. Mus. S. Rhod., 26B: 763. Type locality: Zimbabwe: Junction between Limpopo and Shashi rivers [22 12 S 29 23 E] [Goto Description]. Holotype: [Unknown] ad ♂. Collected by: M.P. Stuart Irwin et al.; collection date: 5 May 1960; original number: 9932. Rhodesian School Exploration Society Sentinel Ranch Expedition. - Comments: Homonym of T. australis Gould, 1854, from Australia (Harrison, 1964a; Kock, 1969a: 74; Meester et al., 1986: 32; Simmons, 2005). 1964. Taphozous sudani rhodesiae Harrison, Arnoldia Rhod., 1 (3): 2. - Comments: Renaming of australis Harrison, 1962, preoccupied. ? Taphozous perforatus sudani: (Current Combination, Current Spelling) PARASITES: Polyctenidae: Eoctenes intermedius (Speiser 1904) from the Congo (Haeselbarth et al., 1966: 16). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), South Sudan, Tanzania, Uganda. †Genus Vespertiliavus Schlosser, 1887 *1887. Vespertiliavus Schlosser, Beitr. Pal. Geol. Österr.-Ung., 6: 70; pl. I, figs 37, 40, 44, 45, 47, 48, 5060,. 426 ISSN 1990-6471 †Vespertiliavus aenigma (Ravel, 2016) *2016. ?Vespertiliavus aenigma Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 384, figs 14, 15. Publication date: 30 September 2016. Type locality: Tunisia: Kassérine governorate: Djebel Chambi National Park: Chambi [35 14 03 N 08 45 29 E, 630 m] [Goto Description]. - Etymology: From the latin aenigma, meaning enigma, puzzle or riddle, referring to the original morphology of the specimens, which makes it difficult to assign them systematically (see Ravel et al., 2016: 384). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. †Vespertiliavus kasserinensis (Ravel, 2016) *2016. Vespertliavus kasserinensis Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 377, figs 11 - 13. Publication date: 30 September 2016. Type locality: Tunisia: Chambi [35 14 03 N 08 45 29 E, 630 m] [Goto Description]. - Etymology: The name refers to the city of Kasserine, at the foot of Djebel Chambi (see Ravel et al., 2016: 377). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Centeno-Cuadros et al., 2019: Suppl.) to early middle Eocene. Family NYCTERIDAE Van der Hoeven, 1855 *1855. Nycteridae Van der Hoeven, Handboek Dierkunde, 2nd ed., 2: 1028. - Comments: Type genus: Nycteris Cuvier and Geoffroy, 1795. - Etymology: The family gets its name from the Greek name for a bat, Nycteris (see Taylor, 2005). (Current Combination) 1866. Nyctericina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 91. Publication date: 1 February 1866. 1910. Petaliidæ Miller, Proc. Biol. Soc. Wash., 22: 90 [Goto Description]. - Comments: Type genus: Nycteris Cuvier and Geoffroy, 1795. Replacement name for Nycteridae Dobson, 1875, due to the fact that Miller (1910) considered the name for the type genus (Nycteris Geoffroy 1803) to be antedated by Nycteris Borkhausen, 1797). 1984. Nycterididae: Butler, Palaeovert., 14 (3): 117, 185. Publication date: 15 November 1984. (Emendation) 2015. Nycteriidae: Anti, Owusu, Agbenyega, Annan, Badu, Nkrumah, Tschapka, Oppong, AduSarkodie and Drosten, Emerg. Inf. Dis., 21 (8): 1419. Publication date: August 2015. (Lapsus) 2018. Nicteridae: Malekani, Musaba, Gembu, Bugentho, Toengaho, Badjedjea, Ngabu, Mutombo, Laudisoit, Ewango, Van Cakenberghe, Verheyen, Asimonyo, Masudi, Bongo and Ngbolua, Nat. Conserv. Res., 3 (1): 68.. Publication date: January 2018. (Lapsus) TAXONOMY: Koopman (1993a) mentions Van der Hoeven (1855) as author whereas Corbet and Hill (1992: 88) mention Dobson (1875e: 348). Flower (1900: 344), Sclater (1901: 120), Medway (1978: 19), Sigé and Legendre (1982), Kock et al. (2002: 82), Hand and Archer (2005: 376), and Lanza et al. (2015: 232) use the name Nycterididae, but we follow Gunnell and Simmons (2005: 212), who state (when referring to the Tanzanycteridae): "This family is a new taxon named by Gunnell et al. (2003) for Tanzanycteris. They spelled the family name Tanzanycterididae, but we follow Simmons and Geisler (1998: 133, footnote 13), who argued that all bat family group names based on generic epithets ending with the Greek root -nycteris should be spelled the same way, i.e., -nycteridae rather than -nycterididae." Simmons and Geisler (1998: 133) state: "Despite Russell and Sigé (1970)'s argument about the nature of the greek root of -nycteris, we think that African Chiroptera Report 2020 it is counterproductive to spell some family-group names in one fashion and others differently. Because Nycteridae is widely accepted as the spelling of the family-group name based on Nycteris, we argue that all family-group names based on genera ending in -nycteris should be spelled in the same fashion." Currently (Simmons and Cirranello, 2020) recognized genus in the family Nycteridae: Nycteris G. Cuvier and E. Geoffroy, 1795. COMMON NAMES: Afrikaans: Langoor-vlermuise. Czech: rýhonosovití, nykteridovití. Dutch: Spleetneusvleermuizen. English: Slit-faced bats, Hollow-faced bats, Hispid bats, Long-eared bats. Finnish: Valkopäälepakot. French: Nyctéridés. German: Schlitznasenfledermäuse, SchlitznasenFledermäuse, Schlitznasen. Norwegian: Hullneser, striflaggermus. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Shi and Rabosky (2015: 1537) calculated the family-level stem age and crown age to be respectively 52.8 and 17.9 MYA. The split off from the Emballonuridae occurred according to Amador et al. (2016: 22) some 51 MYA, and the diversification within the family started about 33 MYA. 427 Ravel et al. (2016: 415, 418) indicate that Koufechia gunnelli represents the basal taxon of the Nycteridae. They also suggest that the family originated in Africa in the Lower Eocene and migrated to Europe in the Oligocene, and even more recently to Asia. MOLECULAR BIOLOGY: The karyotype of the various species in this family has a 2n value between 34 and 42 (Sotero-Caio et al., 2017: 5). VIRUSES: Filoviridae Although no Ebolaviruses have been reported from representatives of this family, Shapiro et al. (2020: 1) indicated that the outbreaks or spillover events of Ebola occurred in areas with high species richness of nycterid bats (and low levels of both anthropogenic disturbance and human population density). They also suggested that more research is required to investigate the role of the Nycteridae in Ebola spillover to humans as up until now the presence of Ebolavirus was only examined in one single species: Nycteris hispida, which turned out to be negative for this virus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". †Genus Khoufechia Ravel, 2016 *2016. Khoufechia Ravel, in: Ravel et al., Geodiversitas, 38 (3): 391. Publication date: 30 September 2016. - Etymology: From the Arabian "Khouféche", a term to designate abat (see Ravel et al., 2016: 391). TAXONOMY: Ravel et al. (2016: 358) indicate that this genus has a basal position within the Nycteridae. †Khoufechia gunnelli Ravel, 2016 *2016. Khoufechia gunnelli Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 391, figs 18 - 20. Publication date: 30 September 2016. Type locality: Tunisia: Kassérine governorate: Djebel Chambi National Park: Chambi [35 14 03 N 08 45 29 E, 630 m] [Goto Description]. - Etymology: In honour of Gregg Gunnell for his numerous contributions to the study of African paleogene bats (see Ravel et al., 2016: 394). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. Genus Nycteris G. Cuvier and E. Geoffroy, 1795 *1795. Nycteris G. Cuvier and E. Geoffroy, Mag. Encyclop. , 1 année 2: 186. - Comments: Type: Vespertilio hispidus Schreber, 1774. Gray et al. (1999: 1) mention Geoffroy and Cuvier, 428 ISSN 1990-6471 1803. 1813. 1838. 1845. 1866. 1866. 1977. 1998. ? 1795 as authors. - Etymology: From the Greek "νυκτορισ", meaning "nocturnal", or from the feminine Greek substantive "νυκτερίς" (nykterís), "bats" (see Lanza et al., 2015: 234). (Current Combination) Nicteris: Desmarest, Nouveau Dictionnaire d'Histoire Naturelle, 15: 501. (Lapsus) Nycterus: G. Fischer, Zoognosia, ed. iii, I: 18. (Lapsus) Petalia Gray, Mag. Zool. Bot., Edinburgh, 2 (12): 494. Publication date: 1 February 1838. - Comments: Type species: Nycteris Javanicus E. Geoffroy Saint-Hilaire, 1813. Nyctoris: ?, London Encyclopedia, 22: 738. (Lapsus) Nycterops Gray, Proc. zool. Soc. Lond., 1866, I (vi): 83. Publication date: May 1866 [Goto Description]. - Comments: Type species: Nycterops pilosa Gray, 1866 (=Vespertilio hispidus Schreber, 1774). - Etymology: Derived from Nycteris and the Greek "οψ", meaning aspect (see Palmer, 1904: 465). Pelatia: Gray, Proc. zool. Soc. Lond., 1866, I (vi): 83. Publication date: May 1866. (Lapsus) Petelia: Lekagul and McNeeley, Mammals of Thailand, 107. (Lapsus) Nyteris: Van Cakenberghe and De Vree, Bonn. zool. Beitr., 48 (2): 123. (Lapsus) Nycteris sp.: TAXONOMY: Meester et al. (1986) state that Andersen (1912b) divided Nycteris into four groups, three of which, viz. hispida, aethiopica (= macrotis - Koopman, 1965, 1975) and thebaica, occur in Southern Africa. The fourth group (arge) is limited to the African forest block. Ellerman and Morrison-Scott (1951 1: 106), Corbet and Hill (1992: 88) mention Nycteris to be a nomen nudum, which was validated by Opinion 111 (Anonymus, 1929) [See also Stiles (1914: 66)]. Revised by Van Cakenberghe and De Vree (1985, 1993a, 1993b, 1999a). A key is provided by Gray et al. (1999: 1, Mammalian Species, 612). Using mitochondrial and nuclear DNA, Demos et al. (2019b) distinguished at least 16 lineages in African Nycteris, 10 of which where wholly or partly occurring in East Africa, which maked them suspect that more lineages will be found in Central and West Africa. Their analyses confirmed the four species groups recognized within the genus. They distinguish the following putative species: arge 1, arge 2, grandis, hispida/aurita, cf. hispida/aurita, macrotis 1, macrotis 2, macrotis 3, nana 1, nana 2, thebaica 1, thebaica 2, thebaica 3, thebaica 4, thebaica 5, thebaica 6, cf. thebaica 1, cf. thebaica 2, cf. thebaica 3. Currently (Simmons and Cirranello, 2020) recognized species of the genus Nycteris: arge Thomas, 1903; aurita (K.Andersen, 1912); gambiensis (K. Andersen, 1912); grandis Peters, 1865; hispida (Schreber, 1774); intermedia Aellen, 1959; javanica E. Geoffroy Saint-Hilaire, 1813 – Java, Nusa Penida (near Bali) and Kangean Isl (Indonesia) (Simmons, 2005: 392); macrotis Dobson, 1876; madagascariensis G. Grandidier, 1937; major (K. Andersen, 1912); nana (K. Andersen, 1912); parisii (De Beaux, 1924); thebaica E. Geoffroy Saint-Hilaire, 1818; tragata (K. Andersen, 1912) – Burma, Thailand, western Malaysia, Sumatra, Borneo (Simmons, 2005: 394); woodi K. Andersen, 1914. COMMON NAMES: Czech: rýhonosi, šerowecové, dutonosecové, nykteridy. English: Hollow-faced bats, Slit-faced bats. French: Nyctères. German: SchlitznasenFledermäuse, Schlitznasen. Italian: Nitterìdi, Nicterìdi. Kiluba (DRC): Kasusu. Kiande (DRC): Kakorokombe. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Avery (2007: 619) reported on the presence of Nycteris sp. in Pleistocene deposits at Wonderwerk Cave, South Africa. The approximate fossil date for the genus was reported by Shi and Rabosky (2015: 1532) to be about 5.332 million years ago. MOLECULAR BIOLOGY: In their study of the FoxP2 gene, Li et al. (2007: 6) found that Nycteris shows the highest rate of nonsynonymous change at exon 17, which they link with the evolution of nasal emission, multiharmonic call structure and prey location by passive listening. PARASITES: HAEMOSPORIDA Perkins and Schaer (2016: Suppl.) reported Nycteria medusiformis Garnham and Heisch, 1953 from various countries in Africa. ACARI Demodecidae: Fain (1960: 84) described Stomatodex cornett from a Nycteris sp. from Astrida (=Butare), Rwanda. Myobiidae: Fain (1994: 1280) indicates that one species of Ewingana has been reported from a Nycteris, but this is possibly due to a museum African Chiroptera Report 2020 contamination. Various species of the genus Nycteris do harbour one species of the monotypic genus Nycterimyobia (also see Fain, 1994: 1280). DIPTERA Nycteribiidae: Penicillida pachymela Speiser, 1901, from Somalia, Sudan, Kenya, Tanzania, the Congo, Guinea and the Cameroon Mts (Haeselbarth et al., 1966: 114). VIRUSES: Coronaviridae None of the 18 Kenyan bats tested positive for CoV (Tao et al., 2017: Suppl.). 429 Paramyxoviridae Mortlock et al. (2015: 1841) reported that one of two examined Nycteris specimens from Kenya tested positive for Paramyxovirus sequences. Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 1 Tanzanian and 1 Kenyan Nycteris sp. specimen, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Chad, Congo, Congo (Democratic Republic of the), Egypt, Ethiopia, Gabon, Kenya, Namibia, Nigeria, Senegal, Somalia, South Africa, South Sudan, Sudan, Tanzania, Uganda, Zambia, Zimbabwe. Nycteris arge Thomas, 1903 *1903. Nycteris arge Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (72): 633. Publication date: 1 December 1903. Type locality: Cameroon: SW Cameroon: Efulen [02 46 N 10 42 E] [Goto Description]. Holotype: BMNH 1904.2.8.2: ad ♂, skull and alcoholic. Collected by: George Latimer Bates Esq. - Etymology: From the Greek "argos" meaning shiny, bright, nimble, swift, rapid. (Current Combination) 1910. Petalia arge: Thomas and Wroughton, Trans. Linn. Soc. Lond., 19 (5) (63): 488. (Name Combination) 2002. Nycteris argae: Fenton and Bogdanowicz, Can. J. Zool., 80: 1008. (Lapsus) 2014. Nycteris arge 2: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. 2019. Nycteris arge 1: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 ((4): 1023. Publication date: 20 August 2019. TAXONOMY: Formerly included intermedia (Hayman and Hill, 1971: 19), but see Van Cakenberghe and De Vree (1985). arge species group (Van Cakenberghe and De Vree, 1985; Simmons, 2005: 391). COMMON NAMES: Castilian (Spain): Murciélago de Cara Hueca. Chinese: 淡色凹脸蝠. Czech: rýhonos guinejský. English: Bates's Slit-faced Bat, Nimble Slit-faced Bat. French: Nyctère de Bates. German: Bates' Schlitznasen-Fledermaus. Kiluba (DRC): Katutu. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bf; IUCN, 2009; Monadjem et al., 2017ar). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ar). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bf; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004by; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: It parts of its range it is presumably threatened by conversion of forest to agricultural land, and the logging of roost trees (Mickleburgh et al., 2008bf; IUCN, 2009; Monadjem et al., 2017ar). CONSERVATION ACTIONS: Mickleburgh et al. (2008bf) [in IUCN (2009)] and Monadjem et al. (2017ar) reported that in view of the species wide range, it is presumably present in some protected areas. Further studies are needed into the distribution and threats to this species. 430 ISSN 1990-6471 GENERAL DISTRIBUTION: Nycteris arge is widely distributed over much of West and Central Africa. It ranges from Sierra Leone in the west, through West Africa and the Congo Basin to western Uganda, southern Sudan, western Kenya, northern Tanzania and Burundi. It is distributed as far south as southern parts of the Democratic Republic of the Congo and northern Angola. Amori et al. (2016: 217) [referring to Grubb et al. (1998: 74)] pointed out that the record from Togo (reported until ACR, 2015) is actually from Ghana. Native: Angola (Hayman, 1963; Simmons, 2005: 391; Monadjem et al., 2010d: 547); Burundi; Cameroon (Simmons, 2005: 391); Central African Republic (Lunde et al., 2001: 537); Congo (Bates et al., 2013: 335); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Simmons, 2005: 391; Monadjem et al., 2010d: 547); Côte d'Ivoire; Equatorial Guinea [= Bioko] (Simmons, 2005: 391); Gabon; Ghana (Decher and Fahr, 2007: 12 - 13); Guinea (Fahr et al., 2006a: 72); Kenya (Webala et al., 2004: 171; Simmons, 2005: 391); Liberia (Monadjem and Fahr, 2007: 50); Nigeria; Rwanda; Sierra Leone (Simmons, 2005: 391); Sudan (Simmons, 2005: 391); Tanzania; Uganda (Kityo and Kerbis, 1996: 60). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data for 11 specimens: Fa: 45.23 ± 0.880 mm, mass: 10.9 ± 0.94 g, wing loading: 5.9 ± 0.50 N/m 2, aspect ratio: 4.8 ± 0.49. GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: Eisenberg and Wilson (1978: 743) indicate that N. arge has a relatively larger cranial volume than its congeners, which they link to differences in feeding strategy and habitat selection. DETAILED MORPHOLOGY: Baculum - The baculum ranges from 3.15 - 4.28 mm in length; the shaft is long, slender, parallelsided and essentially straight; the base is expanded and round; in lateral profile it is sometimes angled ventrally while the tip is simple, with only a small expansion (Thomas et al., 1994: 19). ECHOLOCATION: From western Uganda, Monadjem et al. (2011: 32) reported two different sets of data for two calls each: Fmin: 21.55 kHz, Fmax: 22.11 kHz, Fchar: 21.88 kHz, Fknee: 21.77 kHz, duration: 4.1 msec; Fmin: 46.43 kHz, Fmax: 47.83 kHz, Fchar: 47.23 kHz, Fknee: 47.23 kHz, duration: 5.9 msec. The authors indicate that these different data might represent harmonics. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: R.B. Woosnam [in Thomas, 1910b: 488] reported it to be flying low over the water of the Aruwimi River. Monadjem et al. (2016y: 367) recorded this species widely from forested habitats between 400 and 1,000 m in the Mount Nimba area. POPULATION: Structure and Density:- This is a common species (Mickleburgh et al., 2008bf; IUCN, 2009; Monadjem et al., 2017ar). Trend:- 2016: Stable (Monadjem et al., 2017ar). 2008: Stable (Mickleburgh et al., 2008bf; IUCN, 2009). PARASITES: HAEMOSPORIDA Schaer et al. (2015: 381) found Nycteria cf. eradi parasites in N. arge from Sierra Leone. Landau et al. (2012: 142) and Perkins and Schaer (2016: Suppl.) mention this parasite as Nycteria eradi Rosin, Landau et al., 1978. HEMIPTERA Polyctenidae: Eoctenes nycteridis (Horváth 1910) from Liberia, Congo, Ruanda-Urundi, Tanzania, Uganda and Eritrea (Haeselbarth et al., 1966: 17). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested [as N. argae] between 1986 and 1999, for antibody to SARS-CoV in sera in one individual from Oriental Province, DRC, none tested positive (0/1). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Burkina Faso, Burundi, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea, Kenya, Liberia, Nigeria, Sierra Leone, South Sudan, Tanzania, Uganda. African Chiroptera Report 2020 431 Figure 142. Distribution of Nycteris arge Nycteris aurita (K. Andersen, 1912) *1912. Petalia aurita K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): 547. Publication date: 1 November 1912. Type locality: Kenya: Kilifi [03 38 N 39 51 E] [Goto Description]. Holotype: BMNH 1889.1.11.1: ad ♀, skull and alcoholic. Collected by: G.D. Trevor-Roper. Presented/Donated by: G.D. Trevor-Roper. - Etymology: From the feminine Latin adjective aurìta, meaning "long-eared" (see Lanza et al., 2015: 236). 1924. Nycteris aurita: de Beaux, Atti Sic. Lig. Sci. Lett., n.s., 3 (1): ???. (Name Combination, Current Combination) 1957. Nycteris hispida aurita: Ansell, Ann. Mag. nat. Hist., ser. 12, 10: ???. (Name Combination) 2019. Nycteris cf. hispida/aurita: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1021. Publication date: 20 August 2019. 2019. Nycteris hispida/aurita: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Syst. Evol. Res., 57 (4): 1021. Publication date: 20 August 2019. GENERAL COMMENTS: Lanza et al. (2015: 236) pointed out that we erroneously mentioned Kitui as type locality for this taxon, whereas this should be Kilifi. TAXONOMY: Nycteris aurita is often considered as a synonym or subspecies of N. hispida (Blyth, 1863: 391), but see Van Cakenberghe and De Vree, 1993b). hispida species group Van Cakenberghe and De Vree, 1993b; Simmons, 2005: 391). COMMON NAMES: Chinese: 安 氏 凹 脸 蝠 . Czech: rýhonos východoafrický. English: Andersen's Slit-faced Bat. French: Nyctère d'Andersen, Nyctère à longues oreilles. German: Andersens Schlitznasen-Fledermaus. Italian: Nitterìde orecchiùta, Nicterìde orecchiùta. CONSERVATION STATUS: Global Justification Although this species is known mainly from isolated records from a large area, it is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population (it is common where it has been recorded), and because its savanna habitat is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008r; IUCN, 2009; Monadjem et al., 2017f). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017f). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008r; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004n; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. 432 ISSN 1990-6471 MAJOR THREATS: The threats to this species are poorly known, however, it is likely that there are no major threats to the species, as it has been recorded from an extensive habitat that is not severely declining (Mickleburgh et al., 2008r; IUCN, 2009; Monadjem et al., 2017f). seems to be common where it has been recorded (Mickleburgh et al., 2008r; IUCN, 2009; Monadjem et al., 2017f). CONSERVATION ACTIONS: Mickleburgh et al. (2008r) [in IUCN (2009)] and Monadjem et al. (2017f) report that it is not known if there are any direct conservation measures in place, or if the species is present within any protected areas. Further studies are needed into the distribution, natural history and possible threats to this little known species. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia, Kenya, Rwanda, Somalia, South Sudan, Tanzania. Trend:- 2016: Unknown (Monadjem et al., 2017f). 2008: Unknown (Mickleburgh et al., 2008r; IUCN, 2009). GENERAL DISTRIBUTION: Nycteris aurita is known from scattered records in East Africa. It has been recorded from northern Somalia in the north of its range, through southern Ethiopia and Sudan, into Kenya and Tanzania in the south. Native: Ethiopia (Lavrenchenko et al., 2004b: 147; Simmons, 2005: 391); Kenya (Simmons, 2005: 391); Somalia (Simmons, 2005: 391); Sudan; Tanzania (Simmons, 2005: 391), United Republic of; Uganda; Zambia. Figure 143. Distribution of Nycteris aurita POPULATION: Structure and Density:- There is little information available on the abundance of this species, but it Nycteris cf. parisii De Beaux, 1924 2016. Nycteris cf. parisii Kruskop, Benda, Vasenkov and Lavrenchenko, Lynx, 47: 60. TAXONOMY: Due to the lack of molecular genetic analyses, Kruskop et al. (2016: 60) indicate that taxonomic composition and species delimitations in the N. woodi complex are rather uncertain. Van Cakenberghe and De Vree (1985) suggested that parisii and benuensis are synonyms of N. woodi, but based on bacular morphology, Thomas et al. (1994) suggested that parisii is specifically different from woodi. Kruskop et al. (2016: 60) state that the shape of the baculum of the specimens they report on, is more like that of woodi, and not like that of parisii which would be the species involved if the complex contains more than one species. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia. Figure 144. Distribution of Nycteris cf. parisii African Chiroptera Report 2020 433 Nycteris gambiensis (K. Andersen, 1912) *1912. Petalia gambiensis K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): 548. Publication date: 1 November 1912. Type locality: Senegal: Dialocote [13 25 N 13 20 W] [Goto Description]. Holotype: BMNH 1911.6.10.10: ad, skin and skull. Collected by: G.F. Owen; collection date: 7 May 1910; original number: A5. 1923. P[etalia (Nycteris)] th[ebaica] gambiensis: de Beaux, Atti Soc. ital. Sci. nat., 62: 96. (Name Combination) 1929. Nycteris thebaica gambiensis: Ingoldby, Ann. Mag. nat. Hist., ser. 10, 3: ???. (Name Combination) 1939. Nycteris gambiensis: G.M. Allen, Bull. Mus. comp. Zool., 83: 69. (Name Combination, Current Combination) 1975. [Nycteris] gambianus: Koopman, Bull. Am. Mus. Nat. Hist., 154 (4): 381. (Lapsus) 2007. Nycteris cambiensis: Nel and Markotter, Crit. Rev. Microbiol., 33 (4): 308. (Lapsus) TAXONOMY: Reviewed by Van Cakenberghe and De Vree (1999a). thebaica species group (Van Cakenberghe and De Vree, 1999a; Simmons, 2005: 392). COMMON NAMES: Czech: rýhonos západoafrický. English: Gambian Slit-faced Bat. French: Nyctère de Gambie. German: Gambia SchlitznasenFledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Bergmans and Fahr, 2008; IUCN, 2009; Monadjem et al., 2017ah). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ah). 2008: LC ver 3.1 (2001) (Bergmans and Fahr, 2008; IUCN, 2009). 2004: LC ver 3.1 (2001) (Bergmans and Fahr, 2004; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There appear to be no major threats to this species overall. It is threatened in parts of its range by habitat loss, often resulting from conversion of savanna to agricultural use (Bergmans and Fahr, 2008; IUCN, 2009; Monadjem et al., 2017ah). CONSERVATION ACTIONS: Bergmans and Fahr (2008) [in IUCN (2009)] and Monadjem et al. (2017ah) report that there appear to be no direct conservation measures in place for this species, and it is not known if it occurs in any protected areas. There is a need to protect roosts of this species, and to better determine the eastern limits of the species range. GENERAL DISTRIBUTION: Nycteris gambiensis is a West African species which ranges from Senegal and the Gambia, through most of West Africa, as far east as central Nigeria. There is a single uncertain record from Cameroon which requires verification. A record from Sierra Leone is in error (J Fahr pers. comm. in Simmons, 2005: 392) Native: Benin (Simmons, 2005: 392); Burkina Faso (Simmons, 2005: 392; Kangoyé et al., 2015a: 615); Côte d'Ivoire (Simmons, 2005: 392); Gambia (Simmons, 2005: 392); Ghana (Simmons, 2005: 392); Guinea (Simmons, 2005: 392; Decher et al., 2016: 264); Guinea-Bissau (Monard, 1939; VeigaFerreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 43; Simmons, 2005: 392); Mali; Mauritania; Niger; Nigeria (Simmons, 2005: 392); Senegal (Simmons, 2005: 392); Togo (Simmons, 2005: 392). Uncertain: Cameroon; Sierra Leone (Simmons, 2005: 392). DETAILED MORPHOLOGY: Baculum - The baculum ranges from 2.88-3.42 mm in length; the shaft is slender and parallelsided; the base is expanded and in lateral view is angled ventrally, while the tip is simple with a slight expansion (Thomas et al., 1994:22). POPULATION: Structure and Density:- It can be a locally common species (Bergmans and Fahr, 2008; IUCN, 2009; Monadjem et al., 2017ah). Trend:- 2016: Unknown (Monadjem et al., 2017ah). 2008: Unknown (Bergmans and Fahr, 2008; IUCN, 2009). 434 ISSN 1990-6471 VIRUSES: Willoughby et al. (2017: Suppl.) report the following viruses: Chobar Gorge virus, Lagos bat lyssavirus, Saboya virus. Rhabdoviridae Lyssavirus - Rabies related viruses Lagos bat virus - Isolated in Guinea (Institute Pasteur, 1985). Coronaviridae - Coronaviruses Betacoronavirus Annan et al. (2013: 457) reported that 46 out of 185 (24.9 %) N. cf. gambiensis specimens tested in Ghana were found to be positive for 2c Betacoronavirus. They also found that juvenile bats and lactating females were more likely to be infected than adult males and non-lactating females. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Burkina Faso, Cameroon, Côte d'Ivoire, Ghana, Guinea, Guinea-Bissau, Nigeria, Senegal, The Gambia, Togo. Flaviviridae Flavivirus de Jong et al. (2011: 10) and Luis et al. (2013: suppl.) reported Saboya virus occurring on this species. Calderon et al. (2016: 8) also mention Sokuluk virus. Reoviridae Orbivirus Fomede virus - reported in the CDC’s Arbovirus Catalogue, to have been isolated in 1978 from the Republic of Guinea, from the brain and internal organs of N. cf. gambiensis. Reported by de Jong et al. (2011: 11) and Luis et al. (2013: suppl.). Figure 145. Distribution of Nycteris gambiensis Nycteris grandis Peters, 1865 *1865. Nycteris grandis Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 351, 358. Type locality: Ghana: vicinity of Elmina [05 05 N 01 21 W] [Goto Description]. Paralectotype: RMNH MAM.27348: ad, skin only. Collected by: Colonel C.J.M. Nagtglas; collection date: 1863. - Comments: Grubb et al. (1998: 76) mention the type locality as 'Usually given as "Guinea" but the type, in the Leiden Museum, was collected in 1863 by Nagtglas, and ascribed to Côte d'Or (Jentink, 1888b: 170). The type locality may be further restricted to the vicinity of Elimina [Ghana] which is the only definite locality Jentink cited for any of the Nagtglas material'. The publicatiion covers a meeting held on 13 July 1865. - Etymology: From the Greek "grandis" meaning magnificent (Brown, 1991; Jaeger, 1972, both in Hickey and Dunlop, 2000: 3). (Current Combination) 1866. Nycteris Baikii Gray, in: Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 672 (for 1866). Type locality: "Africa": West Africa: Baikie. - Comments: Meester et al. (1986: 34) mention "Gray in Peters" as author. Nomen nudum. 1912. Petalia grandis: K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): ???. (Name Combination) 1923. Nycteris marica Kershaw, Ann. Mag. nat. Hist., ser. 9, 12 (70): 534. Publication date: 1 October 1923. Type locality: Tanzania: Kilosa: Tindiga [06 55 S 37 20 E] [Goto Description]. Holotype: BMNH 1923.8.2.1: ad, skin and skull. Collected by: Professor Arthur Loveridge; collection date: 24 January 1922. 1925. Nycteris proxima Lönnberg and Gyldenstolpe, Ark. Zool. Stockholm, 17B (9): 1. Publication date: 14 May 1925. Type locality: Congo (Democratic Republic of the): Semliki valley: Kartoushi [00 44 N 29 34 E] [Goto Description]. Holotype: [Unknown] ad ♂. Collection date: 11 March 1921; original number: 813. See Lönnberg and Gyldenstolpe (1925: 2). 1938. [Nycteris] narica: Frechkop, Exploration du Parc National Albert, 50. (Lapsus) 1938. [Nycteris] praxima: Frechkop, Exploration du Parc National Albert, 50. (Lapsus) 1981. N[ycteris] g[randis] marica: Kock, Senckenb. biol., 61 (5/6): 325. (Name Combination) African Chiroptera Report 2020 1991. 2016. 435 Nycteris graudis: Baker, Honeycutt and Van den Bussche, Bull. Am. Mus. Nat. Hist., 206: 46. (Lapsus) Nycteris grande: Bhatnagar, Smith, Rai and Frahm, Anat. Rec., 299: 495. Publication date: 21 January 2016. (Lapsus) TAXONOMY: Reviewed by Van Cakenberghe and De Vree (1993b). N. marica is sometimes recognized as a distinct savanna subspecies, but this does not seem justified based on morphology (Van Cakenberghe and De Vree, 1993b; Hickey and Dunlop, 2000; Simmons, 2005: 392). hispida species group (Van Cakenberghe and De Vree, 1993b; Simmons, 2005: 392). COMMON NAMES: Afrikaans: Groot spleetneusvlermuis. Chinese: 魁凹脸蝠. Czech: rýhonos velký, nycteris velká. English: Large Slit-faced Bat. French: Grande nyctère. German: Große Schlitznase, Große Schlitznasen-Fledermaus. Portuguese: Morcego grande orelhudo. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bg; IUCN, 2009; Monadjem et al., 2017g). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017g). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bg; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cd; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: In general, there appear to be no major threats to this species as a whole. It is threatened in parts of its range by habitat loss, particularly the logging of large trees used for roosting. Some populations may be threatened by overharvesting for subsistence food (Mickleburgh et al., 2008bg; IUCN, 2009; Monadjem et al., 2017g). CONSERVATION ACTIONS: Mickleburgh et al. (2008bg) [in IUCN (2009)] and Monadjem et al. (2017g) report that in view of the species wide range, it is presumably present in a number of protected areas. No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Nycteris grandis is broadly distributed in subSaharan Africa. It ranges from Senegal, through West and Central Africa, to southern Sudan, southeastern Kenya and eastern Tanzania, with scattered records as far south as Zambia, Zimbabwe and Mozambique. Also found on the islands of Zanzibar and Pemba (Simmons, 2005: 392). It is a lowland species. Native: Benin; Burkina Faso (Kangoyé et al. (2012: 6025; 2015a: 615); Cameroon; Central African Republic (Lunde et al., 2001: 537); Congo (Bates et al., 2013: 335); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Simmons, 2005: 392; Monadjem et al., 2010d: 547); Côte d'Ivoire; Equatorial Guinea (Fahr and Ebigbo, 2003: 128); Gabon; Ghana; Guinea; Kenya (Simmons, 2005: 392); Liberia (Fahr, 2007a: 103); Malawi (Happold et al., 1988; Happold and Happold, 1997b: 815; Simmons, 2005: 392; Monadjem et al., 2010d: 547); Mozambique (Smithers and Lobão Tello, 1976; Simmons, 2005: 392; Monadjem et al., 2010d: 547; Monadjem et al., 2010c: 381); Nigeria; Senegal (Simmons, 2005: 392); Sierra Leone; Sudan; Tanzania (Stanley and Goodman, 2011: 43) (including the islands of Pemba and Unguja [O'Brien, 2011: 289]); Togo (Capo-Chichi et al., 2004: 162); Uganda (Kityo et al., 2009b: 137); Zambia (Ansell, 1969; Ansell, 1986; Monadjem et al., 2010d: 547); Zimbabwe (Simmons, 2005: 392; Monadjem et al., 2010d: 547). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Hickey and Dunlop (2000). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Hickey and Dunlop (2000). DETAILED MORPHOLOGY: Baculum - The baculum ranges from 3.45-3.54 mm in length and is small in relation to body size; the shaft is stout and deep, unlike that of any other Nycteris examined, the base is narrow and in lateral profile is usually angled ventrally, while the tip is squared, with a pronounced ventral projection (Thomas et al., 1994: 21). 436 ISSN 1990-6471 For a description of the skull, ears and tragus, and wingshape and aspect ratio see Hickey and Dunlop (2000). SEXUAL DIMORPHISM: N. grandis is not sexually dimorphic in size (Hickey and Dunlop, 2000: 1). POPULATION: Structure and Density:- It appears to be a rare, or rarely recorded, species. The species occurs in small colonies, but is usually found only as single animals or in pairs (Mickleburgh et al., 2008bg; IUCN, 2009; Monadjem et al., 2017g). ECHOLOCATION: See Taylor (1999b) and Hickey and Dunlop (2000). Trend:- 2016: Decreasing (Monadjem et al., 2017g). 2008: Decreasing (Mickleburgh et al., 2008bg; IUCN, 2009). Keeley et al. (2018: 14) indicate that echolocation calls range between 17 and 114 kHz. ACTIVITY AND BEHAVIOUR: See Hickey and Dunlop (2000). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Porter et al. (2010) reported for a single male Gabonese specimen as 2n = 42, with 34 biarmed and 6 acrocentric autosomes, with an X as a large subtelocentric and the Y is a small acrocentric. Protein / allozyme - Unknown. REPRODUCTION AND ONTOGENY: Krutzsch (2000: 115), referring to Verschuren (1957), indicates that N. grandis is polyoestrous with two or more (continuous) cycles per year, a postpartum oestrus and often no anoestrus. HABITAT: This species has been recorded from a variety of lowland habitats, ranging from lowland tropical moist forest (often found near to swampy sites) to drier savanna areas and Miombo woodland (Rosevear, 1965; Fenton et al., 1990; Hickey and Dunlop, 2000; Skinner and Chimimba, 2005). Kangoyé et al. (2015a: 615) report it from a gallery forest in Lera, Burkina Faso. In the Mount Nimba area, Monadjem et al. (2016y: 367) recorded this species from forested and forest edge habitats between 400 and 700 m. HABITS: See Hickey and Dunlop (2000). ROOST: Nycteris grandis generally roosts in hollow trees, but is also found in man made structures such as houses (Fenton et al., 1990; 1993), disused water towers (Fenton et al., 1990; 1993) and culverts (Rosevear, 1965). They may also roost in hollow fallen logs, and holes or small caverns in rocks (Rosevear, 1965; Hickey and Dunlop, 2000). DIET: Fenton et al. (1981: 463; 1990: 2, 4) found that in the Mana Pools National Park, in Zimbabwe, N. grandis primarily fed on frogs (e.g. Ptychadena anchietae, P. mossambica, P. spp., Xenopus laevis), bats (e.g. Nycteris thebaica, Rhinolophus cf darlingi, "Pipistrellus cf. nanus", Hipposideros cf. caffer), arthropods, but sometimes also captured birds (Cisticola fulvicapilla, Apalis flavida) and fish (Tilapia rendalli, Barbus sp., Alestes sp.). Stanley and Goodman (2011: 43), referring to Verschuren (1957), report on a pregnant female collected on 31 July 1993 in the East Usambara Mountains (Tanzania). PARASITES: Little information is available regarding parasites of N. grandis (Hickey and Dunlop, 2000). HAEMOSPORIDA Schaer et al. (2015: 381) found two different forms of Nycteria sp. parasites in N. grandis from Sierra Leone. Karadjian et al. (2016: 9834) mentioned Nycteria medusiformis and Nycteria sp. from the DRC. HEMIPTERA Polyctenidae: Eoctenes nycteridis (Horváth, 1910) from Liberia, Congo, Ruanda-Urundi, Tanzania, Uganda and Eritrea (Haeselbarth et al., 1966: 17). SIPHONAPTERA Pulicidae: Echidnophaga aethiops Jordon and Rothschild, 1906 locality not mentioned (Haeselbarth et al., 1966: 135). VIRUSES: Keymer (1971) reported "broad trypanosomes" from four of six specimens collected from Zambia. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea, Kenya, Liberia, Malawi, Mozambique, Nigeria, Sierra Leone, Tanzania, Togo, Zambia, Zimbabwe. African Chiroptera Report 2020 437 Figure 146. Distribution of Nycteris grandis Nycteris hispida (Schreber, 1774) *1774. Vespertilio hispidus Schreber, Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen, 1 (8): pl. 56 + 1 (9): 169, 188. Publication date: 1774. Type locality: Senegal: "Senegal" [Goto Description]. - Etymology: From the masculine Latin adjective hìspidus, meaning "shaggy", referring to the bristly hair around the facial slit (see Lanza et al., 2015: 242). 1813. Nycteris Daubentonii E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 20: 19. Publication date: August 1813. Type locality: Senegal: "Senegal" [Goto Description]. 1840. Nycteris hispida: Wagner, in: Schreber, Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen, Suppl. 3: ???. (Name Combination, Current Combination) 1843. Nycteris poensis Gray, Catalogue of the species of mammals of the British Museum, 24. Publication date: 13 May 1843. Type locality: Equatorial Guinea: Bioko [formerly Fernando Po] [ca. 03 21 N 08 40 E] [Goto Description]. Holotype: BMNH 1855.12.26.252: ad ♀, skin and skull. Presented/Donated by: Captain E. Downes R.N. Intended specimen = Rhinolophus martini Fraser: see Meester et al. (1986: 33). Comments: Considered a nomen nudum by Dobson (1878: 162) and Allen (1939a: 69). 1843. Rhinolophus Martini Fraser, Proc. zool. Soc. Lond., 1843, V (cxxi): 25. Publication date: July 1843. Type locality: Equatorial Guinea: Bioko [formerly Fernando Po] [ca. 03 21 N 08 40 E] [Goto Description]. 1852. Nycteris villosa Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 48, pl. 11. Publication date: 1852. Type locality: Mozambique: Inhambane [24 00 S 35 28 E] [Goto Description]. Holotype: ZMB 394/85665: ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008) [skull: ZMB 394 + skeleton: ZMB 85665]. - Comments: Considered a valid subspecies by Taylor (1998: 33). 1866. Nycterops pilosa Gray, Proc. zool. Soc. Lond., 1866, I (vi): 83. Publication date: May 1866. Type locality: "Africa": "Africa". 1878. Nycteris hispida villosa: Dobson, Catalogue of the Chiroptera of the collection of the British Museum, 163. (Name Combination) 1883. Nycteris hispidus: Rochebrune, Act. Soc. Linn. Bordeaux, 37: ???. (Name Combination) 1910. Petalia hispida: Thomas and Wroughton, Trans. Linn. Soc. Lond., 19 (5) (63): 488. (Name Combination) 1917. Nycteris pallida J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 425. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Faradje [03 44 N 29 43 E] [Goto Description]. Holotype: AMNH 49144: ad ♂. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 1 March 1912; original number: 1858. Paratype: RMCA 12390: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 22 Novermber 1911; original number: formerly AMNH 49119. Presented/Donated by: ?: Collector Unknown. 438 ISSN 1990-6471 1935. 1935. 1935. 2019. 2019. 2019. Topotype: MCZ 17226: ♂, alcoholic (skull not removed). Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 9 March 1912. Presented/Donated by: ?: Collector Unknown. [Nycteris hispida] martini: Braestrup, Vidensk. Meddr dansk naturh. Foren., 99: 88. (Name Combination) N[ycteris] h[ispida] pallida: Braestrup, Vidensk. Meddr dansk naturh. Foren., 99: 88. (Name Combination) Nycteris hispida hispida: Braestrup, Vidensk. Meddr dansk naturh. Foren., 99: 88. (Name Combination) Nycteris cf. hispida/aurita: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1021. Publication date: 20 August 2019. Nycteris hispida/aurita: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Syst. Evol. Res., 57 (4): 1021. Publication date: 20 August 2019. Nycteris hispidar: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) TAXONOMY: Koopman (1975: 377, 378) includes aurita and pallida. Van Cakenberghe and De Vree (1993b) consider aurita as a separate species. hispida species group (Van Cakenberghe and De Vree, 1993b; Simmons, 2005: 392). COMMON NAMES: Afrikaans: Harige spleetneusvlermuis, Harige Langoorvlermuis. Chinese: 粗 毛 凹 脸 蝠 . Czech: rýhonos štětinatý, šerowec senegalský. English: Hairy Slit-faced Bat, Hairy Long-eared Bat. French: Nyctère hérissée, Nyctère hirsute. German: Rauhaarschlitznase, Gemeine Schlitznasen-Fledermaus, Haarige Schlitznasenfledermaus. Italian: Nitterìde irsùta, Nicterìde irsùta, Nitterìde ìspida, Nicterìde ìspida. Kiluba (DRC): Katutu. Portuguese: Morcego orelhudo piloso. ETYMOLOGY OF COMMON NAME: Although this species is not hairier than any of the other slit-faced bats, the name is applied to it as a translation of the specific name hispida, which is from the Latin hispidus, meaning hairy or bristly, refering to the bristly hair around the facial slit (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008s; IUCN, 2009; Monadjem et al., 2017h). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017h). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008s; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004m; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016c). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: There appear to be no major threats to this species. It is locally threatened in parts of its range by habitat loss, largely through the conversion of forest to agricultural use (Mickleburgh et al., 2008s; IUCN, 2009; Monadjem et al., 2017h). CONSERVATION ACTIONS: Mickleburgh et al. (2008s) [in IUCN (2009)] and Monadjem et al. (2017h) report that in view of the species wide range it is presumably present in a number of protected areas (including the Udzungwa Mountains National Park of Tanzania (Stanley et al., 2005b)). No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: This species has a wide range, encompassing much of sub-Saharan Africa, with the exception of the Horn of Africa and parts of southern Africa. There is an apparently disjunct population in western Mauritania close to the border with Senegal, and an isolated record from central Mali. Also recorded from the island of Unguja [=Zanzibar] (Simmons, 2005: 392; O'Brien, 2011: 289). The records from Cape of Good Hope are probably invalid (Cotterill, 1996a). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 710,458 km2. African Chiroptera Report 2020 Native: Angola (Crawford-Cabral, 1989; Hayman, 1963; Simmons, 2005: 392; Monadjem et al., 2010d: 547); Benin (Capo-Chichi et al., 2004: 162); Botswana (Archer, 1977; Cotterill, 2004a: 261; Simmons, 2005: 392; Monadjem et al., 2010d: 547); Burkina Faso (Kangoyé et al., 2015a: 615); Burundi; Cameroon; Central African Republic (Lunde et al., 2001: 537); Chad; Congo (Bates et al., 2013: 336); Congo (The Democratic Republic of the) (Hayman et al., 1966; Dowsett et al., 1991: 259; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 547); Côte d'Ivoire; Equatorial Guinea [= Bioko] (Simmons, 2005: 392); Ethiopia (Lavrenchenko et al., 2004b: 132); Gabon; Gambia (Simmons, 2005: 392; Emms and Barnett, 2005: 50); Ghana; Guinea (Fahr et al., 2006a: 72); Guinea-Bissau (Bocage, 1892a; Monard, 1939; Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 41); Kenya; Liberia (Fahr, 2007a: 103); Malawi (Happold et al., 1988; Happold and Happold, 1997b: 815; Simmons, 2005: 392; Monadjem et al., 2010d: 547); Mali; Mauritania (Simmons, 2005: 392); Mozambique (Smithers and Lobão Tello, 1976; Simmons, 2005: 392; Monadjem et al., 2010d: 547; Monadjem et al., 2010c: 381); Namibia (Monadjem et al., 2010d: 547); Niger; Nigeria; Rwanda; Senegal (Adam and Hubert, 1976; Simmons, 2005: 392); Sierra Leone; Somalia (Simmons, 2005: 392); South Africa (Monadjem et al., 2010d: 547); Sudan; Tanzania (Stanley et al., 2005b); Togo; Uganda (Kityo and Kerbis, 1996: 60); Zambia (Ansell, 1973; Ansell, 1974; Ansell, 1986; Monadjem et al., 2010d: 547); Zimbabwe (Monadjem et al., 2010d: 547). DETAILED MORPHOLOGY: Baculum - The baculum ranges from 3.33 - 4.16 mm in length, the shaft is long, slender and variably ventrally curved, the base is expanded and in lateral profile is angled ventrally, while the tip is sometimes angled ventrally, it has a projection on the ventral aspect varying between individuals from a slight swelling to a pronounced hook (Thomas et al., 1994: 20). 439 moist forest, into moist savanna, dry savanna, papyrus swamps and marsh. R.B. Woosnam [in Thomas, 1910b: 488] reported it to be most numerous among the acacia trees on the plains around the south end of the Ruwenzori range. Monadjem et al. (2016y: 367) recorded N. hispida sparsely in open savanna on the Liberian side of Mount Nimba. ROOST: Colonies roost in hollow trees, dense bushes, caves, holes in termite colonies and similar habitats. PREDATORS: Mikula et al. (2016: Supplemental data) mention N. hispida to be taken by the Western marsh harrier (Circus aeruginosus (Linnaeus, 1758)). POPULATION: Structure and Density:- Colonies range in size from individual and pairs of animals to up to 20 bats. This is a very common bat (Mickleburgh et al., 2008s; IUCN, 2009; Monadjem et al., 2017h). Trend:- 2016: Stable (Monadjem et al., 2017h). 2008: Stable (Mickleburgh et al., 2008s; IUCN, 2009). REPRODUCTION AND ONTOGENY: Krutzsch (2000: 94) reports that the testes are probably permanently scrotal in N. hispida, according to Matthews (1941). The specieis's reproductive cycle is polyoestrous with two or more (continuous) cycles per year, a postpartum oestrus and often no anoestrus. Matthews (1941) also noticed that copulation followed shortly after parturition. In Burkina Faso, Kangoyé et al. (2015a: 615) captured females carrying young during the month of September. ECHOLOCATION: See Taylor et al. (2005). From Sierra Leone, Weber et al. (2019: 25) reported a lactating female with a young attached on 25 May. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Lee et al. (1989) reported 2n = 42, FN = 78, BA = 38, and a metacentric X and Y. Protein / allozyme - Unknown. PARASITES: Protozoa Kassahun et al. (2015: 168) examined one N. hispida specimen from Awash-Methara (Ethiopia) and found it infected by Leishmania major Friedlin. HABITAT: Nycteris hispida has been recorded from a wide variety of habitats, ranging from lowland tropical ACARI Trombiulidae: Vercammen-Grandjean and Fain (1958: 30) described Trombigastia (Trombigastia) nycteris from N. hispida hispida from the Astrida 440 ISSN 1990-6471 region (=Butare), Rwanda (Stekolnikov, 2018a: 120). Henipavirus, Pneumovirus. HEMIPTERA Polyctenidae: Eoctenes nycteridis (Horváth, 1910) from Liberia, Congo, Ruanda-Urundi, Tanzania, Uganda and Eritrea (Haeselbarth et al., 1966: 17). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Malawi, Mali, Mauritania, Mozambique, Namibia, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. SIPHONAPTERA Ischnopsyllidae: Lagaropsylla idea Smit, 1957, primary host being molossids but recorded from this taxon (Haeselbarth et al., 1966: 190). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - One fecal sample was tested by Pfefferle et al. (2009) in Ghana in February 2008, and this was found to be negative for coronavirus (CoV) RNA. Morbillivirus, Rubulavirus or Hantaviridae (formerly included in Bunyaviridae) Hantavirus Weiss et al. (2012b: 160) report the presence of a new hantavirus, named Magboi virus [MGBV], in one N. hispda specimen collected at the Magboi River, Gola National Park, Sierra Leone. Arai et al. (2019b: 2) and Arai and Yanagihara (2020: 7) tentatively included this virus in the genus Loanvirus. Paramyxoviridae Drexler et al. (2012a: Suppl. Table S1) indicated that the single specimen they examined from Ghana did not test positive for Repirovirus, Figure 147. Distribution of Nycteris hispida Nycteris intermedia Aellen, 1959 *1959. Nycteris intermedia Aellen, Archs Sci. Genève, 12 (2): 218. Type locality: Côte d'Ivoire: NW of Abidjan: Adiopodoumé [05 19 N 04 08 W, 0 - 30 m] [Goto Description]. Holotype: MHNG 923.094: ad ♀, skull and alcoholic. Collected by: Dr. Villy Aellen; collection date: 20 July 1953; original number: A1437. Paratype: MHNG 923.095: ad ♀, skull and alcoholic. Collected by: Dr. Villy Aellen; collection date: 24 July 1953; original number: A1438. Paratype: MHNG 923.096: ad ♀, skull and alcoholic. Collected by: Dr. Villy Aellen; collection date: 19 August 1953; original number: A1439. Presented/Donated by: ?: Collector Unknown. Paratype: ad ♀; alcoholic and skull; Collector: Villy Aellen; Original number: Collection date:. Paratype: MHNG 923.097: ad ♀, skull and alcoholic. Collected by: Dr. Villy Aellen; collection date: 24 July 1953; original number: A1440. Presented/Donated by: ?: Collector Unknown. - Etymology: Referring to its intermediary size between Nycteris arge and Nycteris nana. (Current Combination) TAXONOMY: Formerly included in N. arge, but Van Cakenberghe and De Vree (1985) regard this as a distinct species (Simmons, 2005: 392). COMMON NAMES: Czech: rýhonos prostřední. English: Intermediate Slit-faced Bat. French: Nyctère d'Aellen, Nyctère moyen. German: Intermediäre Schlitznasenfledermaus, Schlitznasen-Fledermaus. Intermediäre CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in some protected areas, and because it is unlikely to be declining fast enough African Chiroptera Report 2020 to qualify for listing in a more threatened category (Mickleburgh et al., 2008bh; IUCN, 2009; Monadjem et al., 2017au). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017au). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bh; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bm; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is threatened in parts of its range by deforestation including loss of roosting trees (Mickleburgh et al., 2008bh; IUCN, 2009; Monadjem et al., 2017au). CONSERVATION ACTIONS: Mickleburgh et al. (2008bh) [in IUCN (2009)] and Monadjem et al. (2017au) report that there do not appear to be any direct conservation measures in place, however, the species has been recorded from some protected areas (e.g. Comoe National Park and Tai National Park both in Côte d'Ivoire). Further research is needed into the distribution, natural history and threats to this species. 441 (Denys et al., 2013: 282); Liberia (Simmons, 2005: 392); Tanzania (Simmons, 2005: 392). MOLECULAR BIOLOGY: Denys et al. (2013: 282) report the karyotype for one female from the Guinean Mount Nimba area as: 2n = 34, NFa = 62, including 15 pairs of large to small metacentric and submetacentric chromosomes and one middle-sized acrocentric pair. The X chromosome is submetacentric. POPULATION: Structure and Density:- This species appears to be relatively rare. It is usually found as small groups of animals (Mickleburgh et al., 2008bh; IUCN, 2009; Monadjem et al., 2017au). Trend:- 2016: Decreasing (Monadjem et al., 2017au). 2008: Decreasing (Mickleburgh et al., 2008bh; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Gabon, Ghana, Guinea, Liberia, Tanzania. GENERAL DISTRIBUTION: Nycteris intermedia is broadly distributed in subSaharan Africa, ranging from Liberia in the west, to western Tanzania in the east, with records as far south as northeastern Angola. The species appears to be known from around 27 localities. It is typically a lowland species. Native: Angola (Van Cakenberghe et al., 1999; Simmons, 2005: 392; Monadjem et al., 2010d: 547); Cameroon; Central African Republic (Lunde et al., 2001: 537); Congo (The Democratic Republic of the); Côte d'Ivoire (Simmons, 2005: 392); Equatorial Guinea; Gabon; Ghana; Guinea Figure 148. Distribution of Nycteris intermedia Nycteris macrotis Dobson, 1876 *1876. Nycteris macrotis Dobson, Monograph of the Asiatic Chiroptera and catalogue of the species of bats in the collection of the Indian Museum Calcutta, 80. Type locality: Sierra Leone: "Sierra Leone" [Goto Description]. Holotype: BMNH 1866.2.2.2: sad ♂, skull and alcoholic. - Etymology: From the Greeek adjective "μακρός" (macròs), meaning "long, large" and the neuter Greek substantive "οὖς" (genitive ώu0964\ός"; "ûs", "ōtós"), meaning "ear", although the ear of the species is not of especially outstanding size (see Lanza et al., 2015: 244). (Current Combination) 1878. Nycteris æthiopica Dobson, Catalogue of the Chiroptera of the collection of the British Museum, xxxviii, 162, 165, pl. 11, fig. 3. Publication date: June 1878. Type locality: Sudan: Kordofan province: Sennaar [15 35 N 33 38 E, 425 m] [Goto Description]. Holotype: BMNH 1847.5.27.32: ad, skin and skull. Collected by: Theodor Kotschy. 1901. Nycteris aethiopica aethiopica: Thomas, Ann. Mag. nat. Hist., ser. 7, 8 (43): 30. 442 ISSN 1990-6471 1901. 1912. 1912. 1912. 1917. 1922. 1923. 1939. 1939. 1942. 1953. 1953. 1960. 1969. 2018. 2019. 2019. 2019. ? ? Nycteris aethiopica luteola Thomas, Ann. Mag. nat. Hist., ser. 7, 8 (43): 30. Publication date: 1 July 1901. Type locality: Kenya: Kitui [01 22 S 38 01 E, 3 500 ft] [Goto Description]. Holotype: BMNH 1901.5.6.4: ad ♀, skin and skull. Collected by: Dr. Sidney Langford Hinde; collection date: 14 February 1901. Petalia aethiopica aethiopica: K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): ???. Petalia aethiopica luteola: K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): ???. Petalia macrotis: K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): ???. (Name Combination) Nycteris major J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 427. - Comments: Not of Andersen (1912b). Nycteris oriana Kershaw, Ann. Mag. nat. Hist., ser. 9, 10 (56): 179. Publication date: 1 August 1922. Type locality: Malawi: Shiré valley: Chiromo [16 32 S 35 09 E, 60 m] [Goto Description]. Holotype: BMNH 1922.4.25.3: ad ♀, skin and skull. Collected by: Rodney Carrington Wood Esq. Collection date: 7 June 1918; original number: 312. Nycteris luteola: Loveridge, Proc. zool. Soc. Lond., 1923, II: 396. Nycteris æthiopica guineensis Monard, Arq. Mus. Bocage, 10: 66, fig. 4. Publication date: March 1939. Type locality: Guinea-Bissau: Mansoa [12 04 N 15 19 W] [Goto Description]. Syntype: [Unknown] ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 1937; original number: 284. Syntype: [Unknown] ♀, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 1937; original number: 282. Syntype: MHNG 1326.005: ad ♂, skull and alcoholic. Collected by: Dr. Albert Monard; collection date: 14 December 1937; original number: 283. Syntype: NMBA 5271: ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 17 Decembre 1937; original number: 319. Syntype: RMCA 15310: ad ♀, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 1937; original number: 285. - Comments: Monard (1939: 66) mentions a larger series: 282 (♀), 283 (♂), 284 (♂), 285 (♀), 319 (♂). Nycteris æthiopica guineensis f. aurantiaca Monard, Arq. Mus. Bocage, 10: 68. Publication date: March 1939. Type locality: Guinea-Bissau: Pitche [12 20 N 13 57 W] [Goto Description]. Nycteris aethiopica oriana: G.M. Allen and Loveridge, Bull. Mus. comp. Zool., 89 (4): 150, 161. Nycteris macrotis macrotis: Ellerman, Morrison-Scott and Hayman, Southern African mammals 1758 to 1951, 54. (Name Combination) Nycteris macrotis oriana: Ellerman, Morrison-Scott and Hayman, Southern African mammals 1758 to 1951, 54. (Name Combination) Nycteris macrotis luteola: Harrison, J. East Afr. Nat. Hist. Soc., 23 (7) (104): 288. Publication date: December 1960. (Name Combination) Nycteris macrotis aethiopica: Kock, Abh. Senckenberg. naturforsch. Ges., 521: 97. (Name Combination) Nycteris macrotis lutreola: Gunnell and Manthi, J. Hum. Evol., Suppl.. Publication date: 6 April 2018. (Lapsus) Nycteris macrotis 1: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris macrotis 2: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris macrotis 3: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris aethiopica: (Alternate Spelling) Nycteris marcrotis oriana: (Name Combination) TAXONOMY: Meester et al. (1986) indicate that both Koopman (1965, 1975, 1982) and Kock (1969a) included the extralimital [to southern Africa] aethiopica Dobson, 1878, and luteola Thomas, 1901, as subspecies in this species. While Hayman and Hill (1971) treat macrotis and aethiopica as separate species, they confirm that the distinction is by no means certain. N. macrotis macrotis Dobson, 1876 may include also Plecotus aethiopicus Fitzinger, 1866, from the White Nile, a nomen nudum (Kock, 1969a). Kock (1969a) regards oriana as a synonym of the extralimital luteola, but Ansell (1978) and Smithers (1983) treat it as a separate subspecies. African Chiroptera Report 2020 Does not include madagascariensis Grandidier, 1937 (see Peterson et al., 1995), or vinsoni Dalquest, 1965 (see Van Cakenberghe and De Vree, 1999a). Simmons (2005: 393) recognises three subspecies - aethiopica Dobson, 1878; luteola Thomas, 1901; and oriana Kershaw, 1922. COMMON NAMES: Afrikaans: Groter spleetneusvlermuis. Chinese: 大耳凹脸蝠. Czech: rýhonos velkouchý, nykteris velkouchá. English: Large-eared Slit-faced Bat, Greater Slit-faced Bat, Big-eared slit-faced bat. French: Nyctère de Dobson, Nyctère à grandes oreilles. German: Großohrige SchlitznasenFledermaus. Italian: Nitterìde di Dòbson, Nicterìde di Dobsòn. ETYMOLOGY OF COMMON NAME: Although the specific name macrotis is derived from the Greek "makros" and "otis" meaning large eared, the species' ears are in fact smaller than in species such as the Egyptian slit-faced bat (N. thebaica) (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008aa; IUCN, 2009; Monadjem et al., 2017k). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017k). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008aa; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004at; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: In general, there are no major threats to this species. In parts of its range it is threatened by general habitat loss, presumably mostly through the conversion of land to agricultural use (Mickleburgh et al., 2008aa; IUCN, 2009; Monadjem et al., 2017k). CONSERVATION ACTIONS: Mickleburgh et al. (2008aa) [in IUCN (2009)] and Monadjem et al. (2017k) report that in view of the species wide range it is presumably present in some protected areas (it has been found in at least one, i.e. Parc National du ‘W’ (Poché, 1975)). No direct conservation measures are needed for this species as a whole. 443 currently GENERAL DISTRIBUTION: Nycteris macrotis is a widespread African species which ranges from Senegal in the west, to Somalia in the east, being recorded as far south as northern Botswana. Found on the island of Zanzibar (Simmons, 2005: 392). It has been recorded between sea level and 2,200 m asl. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,466,326 km2. Native: Angola (Hayman, 1963; Van Cakenberghe and De Vree, 1985; Crawford-Cabral, 1989; Monadjem et al., 2010d: 547); Benin (Capo-Chichi et al., 2004: 162); Botswana (Monadjem et al., 2010d: 547); Burkina Faso (Kangoyé et al., 2015a: 616); Burundi; Cameroon; Central African Republic; Chad; Congo (Bates et al., 2013: 336); Congo (The Democratic Republic of the) (Hayman et al., 1966; Gallagher and Harrison, 1977; Van Cakenberghe and De Vree, 1985; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 547); Côte d'Ivoire; Equatorial Guinea; Ethiopia (Simmons, 2005: 392); Gabon; Gambia (Emms and Barnett, 2005: 50; Simmons, 2005: 392); Ghana; Guinea; Guinea-Bissau (Seabra, 1900a; Monard, 1939; Rainho and Ranco, 2001: 44); Kenya; Liberia (Fahr, 2007a: 103); Malawi (Happold et al., 1988; Van Cakenberghe and De Vree, 1985; Simmons, 2005: 392; Monadjem et al., 2010d: 548); Mali (Meinig, 2000: 105); Mauritania (1st record: Benda et al., 2011a: 267); Mozambique (Lopes and Crawford-Cabral, 1992; Simmons, 2005: 392; Monadjem et al., 2010d: 548; Monadjem et al., 2010c: 381); Niger; Nigeria; Rwanda; Senegal (Simmons, 2005: 392); Sierra Leone; Somalia; Sudan; Tanzania (including the island of Unguja [O'Brien, 2011: 289]); Togo; Uganda (Kityo and Kerbis, 1996: 60); Zambia (Ansell, 1967; Ansell, 1973; Ansell, 1978; Cotterill, 2002b: 6; Monadjem et al., 2010d: 548); Zimbabwe (Simmons, 2005: 392; Monadjem et al., 2010d: 548). Presence uncertain: Namibia (Cotterill, 2004a: 261). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Fain (1953b: 102) reported on the exceptional yellowish-orange colour of some specimens from a cave near the Nyumba Mission (Rwanda). Other specimens from the save cave were uniformly orange or somewhat darker, or yellowish-brown with a few grey hairs, brownishgrey with orange-yellowish hairs, and brownishgrey with a pale belly without any yellowish or orange hairs. These colour differences were not 444 ISSN 1990-6471 found to be linked with either the sexe or the age of the specimens DETAILED MORPHOLOGY: Baculum - The baculum ranges from 3.84 to 5.12 mm in length, the shaft is long, parallel-sided and usually straight, thickening towards the base, the base has two basal lobes of varying development, in lateral view it is sometimes angled ventrally, while the tip is expanded and trifid with three variably developed processes (Thomas et al., 1994: 21). suggest that the Somalian specimen might possibly belong to Nycteris parisii, "aethiopica", or "Nycteris cf. parisii" (sensu Kruskop et al., 2016: 60). Protein / allozyme - Unknown. HABITAT: Monadjem et al. (2016y: 367) reported one specimen from the Mount Nimba area, which was recorded from a disturbed habitat close to a settlement at 500 m. Brain - Based on specimens from Bénin, Amrein et al. (2007) reported that proliferating cells, detected with Ki-67 and MCM2, in the subgranular layer of the dentate gyrus was found to be absent. Amrein et al. (2007) also reported moderate to ample proliferating cells (Ki-67 positive cells) and migrating young neurons (DCX positive cells) in the rostral migratory stream. Furthermore, Amrein et al. (2007) detected few DCX positive cells in the rostral migratory stream and the olfactory bulb. No NeuroD was detected in the hippocampal granule cells (Amrein et al., 2007). ROOST: In the Parc National du 'W' in Niger Republic, 300 km west of Sokoto, they have been found roosting in thatched huts and others were found in abandoned mine shafts (Poché, 1975). Stanley et al. (2007c: 60) refer to Koopman (1975), who mentions that N. macrotis is known to roost in burrows in the ground made by aardvarks (Orycteropus afer) or porcupines (Hystrix cristata). Weber and Fahr (2006: 4) indicate that N. macrotis largely or exclusively depends on the availability of caves as day roosts in Guinea. Femur - Louzada et al. (2019: 136) provided a detailed description of the femur, which has a welldeveloped medial ridge that begins on the shaft of the femur, a short and well-developed lateral ridge, and well-developed trochanters. The latter nearly reach the same level proximally. The femur is greatly sinusoidal and the torsion of the proximal epiphysis is the most conspicuous of all the bat families they examined. POPULATION: Structure and Density:- In the Parc National du ‘W’ in Niger Republic, 300 km west of Sokoto, they have been found roosting in groups of 10-20 individuals (Poché, 1975). Happold (1987) notes that they are solitary or live in small groups. ECHOLOCATION: Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 76.7 kHz and duration: 1.2 msec. REPRODUCTION AND ONTOGENY: Krutzsch (2000: 115) indicates that most nycterid species (including N. aethiopica = N. macrotis) are polyoestrous with two or more (continuous) cycles per year, a postpartum oestrus and often no anoestrus. In the DRC and Rwanda, Anciaux de Faveaux (1978c) reported copulations in August and October. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - In the absence of data on a male of this species (only ane female from Somalia), Lee et al. (1989) described the karyotype as having a diploid number of 2n = 40, FNa = 74, BA = 36, X = M, and suggested that the Y chromosome might be a small metacentric as in the other Nycteris species. Volleth et al. (2020a: 272) reported that a male from Benin had the following karyotype: 2n = 40, FNa = 76, with all of the chromosomes biarmed, consisting of two large submetacentric pairs, 17 medium to small meta- to submetacentric, a medium-sized metacentric X and a very small metacentric Y. Based on the different FNa number, Volleth et al. (2020a: 276) suggest that the two specimens might belong to closely related but different species. They also Trend:- 2016: Unknown (Monadjem et al., 2017k). 2008: Unknown (Mickleburgh et al., 2008aa; IUCN, 2009). Happold and Happold (1990b: 566) reported that, in Malawi, young were born in the wet season and early dry season, suggesting an aseasonal or extended seasonal reproduction, and that females are not in reproductive synchrony. Matthews (1937) [in Krutzsch (2000: 115)] noted that male N. luteola [= N. macrotis] collected at the time of post partum oestrus were reproductively functional. For the same "species", Martin and Bernard (2000: 32) reported that the corpus luteum is only present for the first half of pregnancy. Krutzsch (2000: 94) furthermore reports that the testes are probably permanently scrotal. African Chiroptera Report 2020 445 PARASITES: HAEMOSPORIDA Schaer et al. (2015: 381) found Nycteria sp. parasites in N. macrotis from South Sudan. Flaviviridae In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against non-specific Flaviviruses. HEMIPTERA Polyctenidae: Eoctenes nycteridis (Horváth, 1910) from Liberia, Congo, Ruanda-Urundi, Tanzania, Uganda and Eritrea (Haeselbarth et al., 1966: 17). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Malawi, Mali, Mauritania, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. ACARI Trombiculidae: Stekolnikov (2018b: 280) examined one larva of Trombigastia nycteris Vercammen-Grandjean and Fain, 1958, which was collected from an unknown locality in Rwanda (the slide label refers to "Ada", which might refer to "Astrida" [=Butare]). Stekolnikov (2018a: 162; 2018b: 284) also reported on three larvae of Microtrombicula nycteris (Jadin, VercammenGrandjean and Fain, 1955), collected in Butare. DIPTERA Streblidae: Raymondi alulata Speiser, 1908 from Angola, the Congo, Malawi and Somalia (Haeselbarth et al., 1966: 102). Raymondia scopigera Jobling, 1954 in Cameroon (Haeselbarth et al., 1966: 103, host referred to N. aethiopica), Senegal and Sierra Leone (Shapiro et al., 2016: 255). VIRUSES: Coronaviriddae Joffrin et al. (2020: 6) reported Beta-C coronaviruses from Mozambican bats they identified as "Nycteris thebaica" (but were actually N. macrotis according to Demos et al., 2019b). Figure 149. Distribution of Nycteris macrotis Nycteris madagascariensis G. Grandidier, 1937 *1937. Nycteris madagascariensis G. Grandidier, Bull. Mus. natn. Hist. nat., Paris, sér. 2, 9: 353. Publication date: November 1937. Type locality: Madagascar: N of Diego-Suarez [=Antsiranana]: Rodo valley, N of Pirkana: Ankarana, near the [12 05 S 49 05 E] [Goto Description]. Holotype: MCZ 45433: ♀, skull and alcoholic. Collection date: June 1910. Formerly Grandidier collection; see Helgen and McFadden (2001: 142). Paratype: MCZ 45434: ad ♀, skull and alcoholic. Formerly Grandidier collection; see Helgen and McFadden (2001: 142). - Etymology: Referring to the island of Madagascar, where the species occurs. (Current Combination) TAXONOMY: Earlier included in thebaica, but considered a synonym of macrotis by Van Cakenberghe and De Vree (1985), Koopman (1993a: 162), but a valid species by Peterson et al. (1995: 63), Russ et al. (2001), Simmons (2005: 393). macrotis species group (see Van Cakenberghe and De Vree, 1985). COMMON NAMES: Czech: rýhonos madagaskarský. English: Madagascar Slit-faced Bat. French: Nyctère de Madagascar, Chauve-souris malgache balafrée. German: Madagassische SchlitznasenFledermaus. 446 ISSN 1990-6471 CONSERVATION STATUS: Global Justification This species is listed as Data Deficient (DD ver 3.1 (2001)), as it is only known from two specimens from 1910 in the Irodo River Valley and has not been recorded since. Further extensive surveys are needed for this species, as well as knowledge on its distribution, biology, ecology, and any threats affecting it (Hutson et al., 2008q; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Hutson et al., 2008q; IUCN, 2009). 2000: DD ver 2.3 (1994). Native: Madagascar (Peterson et al., 1995; Simmons, 2005: 393). POPULATION: Structure and Density:- It is known only from two specimens and has not been recorded since 1910 (Hutson et al., 2008q; IUCN, 2009). Trend:- 2008: Unknown (Hutson et al., 2008q; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Regional None known. MAJOR THREATS: The threats to this species are not known (Hutson et al., 2008q; IUCN, 2009). CONSERVATION ACTIONS: Hutson et al. (2008q) [in IUCN (2009)] report that it is not known if the species is present in any protected areas. Further studies are needed into the distribution, abundance, natural history and threats to this species. GENERAL DISTRIBUTION: Nycteris madagascariensis is endemic to Madagascar where it is known only from two specimens caught in northern Madagascar near Analamera in the Irodo River Valley in 1910 (Peterson et al., 1995). Figure 150. Distribution of Nycteris madagascariensis Nycteris major (K. Andersen, 1912) *1912. Petalia major K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): 547. Publication date: 1 November 1912. Type locality: Cameroon: Ja River [ca. 03 00 N 13 00 E] [Goto Description]. Holotype: BMNH 1909.10.2.49: ad ♀, skull and alcoholic. Collected by: George Latimer Bates Esq. Collection date: 23 January 1906. - Etymology: From the Latin "maior" (see Kozhurina, 2002: 16). 1917. Nycteris avakubia J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 426, fig. 3. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Avakubi [01 18 N 27 32 E] [Goto Description]. Holotype: AMNH 49403: ad ♂, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: September 1913; original number: 2623. 1917. Nycteris major: J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 427. Publication date: 29 September 1917. - Comments: The specimens assigned to this species by Allen (1917), however, belong to Nycteris macrotis (see Koopman, 1965). (Name Combination, Current Combination) TAXONOMY: Includes avakubia J.A. Allen, 1917 (see Koopman, 1965; Van Cakenberghe and De Vree, 1985; Simmons, 2005: 393). arge species group (Van Cakenberghe and De Vree, 1985; Simmons, 2005: 393). African Chiroptera Report 2020 COMMON NAMES: Czech: rýhonos kamerunský. English: Ja Slitfaced Bat, Dja Slit-faced Bat; Large Slit-faced Bat. French: Nyctère de Ja. German: DjaSchlitznasenfledermaus, Dja SchlitznasenFledermaus. SIMILAR SPECIES: Eisentraut (1956b: 517) indicated that the claws of N. major are more robust than those of N. arge. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of recent information on its extent of occurrence in Central Africa, natural history, and threats. If the species is found to range more widely in Central Africa it is likely to meet the criteria for Least Concern, as the extent and quality of its habitat are probably not declining at a rate sufficient to justify listing in a higher category (Mickleburgh et al., 2008bi; IUCN, 2009). 447 Republic of the Congo in Central Africa (at Avakubi), with the southern-most record from northern Zambia (close to the border with the Democratic Republic of the Congo). Native: Benin (Capo-Chichi et al., 2004: 162); Cameroon (Simmons, 2005: 393); Central African Republic; Congo (The Democratic Republic of the) (Simmons, 2005: 393); Côte d'Ivoire (Simmons, 2005: 393); Guinea; Liberia (Simmons, 2005: 393); Zambia (Van Cakenberghe and De Vree, 1985; Simmons, 2005: 393; Monadjem et al., 2010d: 548). PREDATORS: Demeler (1981: 133) found remains of N. major in pellets of Bubo africanus in the Yankari Game Reserve (Nigeria). However, as N. macrotis was often mis-identified as N. major and the latter species was not yet reported from that area, the bat's identification still needs to be confirmed. Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008bi; IUCN, 2009). 2004: VU A4c ver 3.1 (2001) (Mickleburgh et al., 2004cg; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). POPULATION: Structure and Density:- This appears to be an extremely rare species, which is only known from a dozen or so specimens. It is not difficult to catch and efforts have been made to locate more specimens (Mickleburgh et al., 2008bi; IUCN, 2009). Regional None known. Trend:- 2008: Unknown (Mickleburgh et al., 2008bi; IUCN, 2009). MAJOR THREATS: The species appears to need rather large trees, and is presumably threatened by deforestation resulting from logging activities and the conversion of forest to agricultural use (Mickleburgh et al., 2008bi; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Côte d'Ivoire, Liberia, Zambia. CONSERVATION ACTIONS: Mickleburgh et al. (2008bi) [in IUCN (2009)] report that although there appear to be no direct conservation measures in place, it has been recorded from the Tai National Park in Côte d'Ivoire. Further studies are needed into the distribution, natural history and threats to this somewhat enigmatic species. There is a need to monitor to presence of this species at its known localities. GENERAL DISTRIBUTION: Nycteris major is distributed in West and Central Africa, where it has been patchily recorded from Guinea, Liberia (at Voinjama) and Côte d'Ivoire in the west, and from Cameroon and the Democratic Figure 151. Distribution of Nycteris major 448 ISSN 1990-6471 Nycteris nana (K. Andersen, 1912) *1912. Petalia nana K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): 547. Publication date: 1 November 1912. Type locality: Congo: [Rio Muni ?] Benito River [ca. 01 31 N 09 52 E] [Goto Description]. Holotype: BMNH 1900.2.5.46: ad ♂, skull and alcoholic. Collected by: George Latimer Bates Esq. 1926. Nycteris nana: Cabrera and Ruxton, Ann. Mag. nat. Hist., ser. 9, 17: xxx. (Name Combination, Current Combination) 1936. Nycteris nana tristis G.M. Allen and Lawrence, Bull. Mus. comp. Zool., 79 (3): 47. Publication date: 30 January 1936. Type locality: Kenya: W Nyanza: Kakamega district: Kaimosi [00 08 N 34 51 E, 2 130 m] [Goto Description]. Holotype: MCZ 31156: ad ♀, skin and skull. Collected by: Professor Arthur Loveridge; collection date: 13 February 1934. See Helgen and McFadden (2001: 142). (Name Combination) 1939. Nycteris nana nana: G.M. Allen, Bull. Mus. comp. Zool., 83: 70. (Name Combination) 2019. Nycteris nana 1: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. 2019. Nycteris nana 2: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. TAXONOMY: Includes tristis Allen and Lawrence, 1936 (Koopman, 1975; Van Cakenberghe and De Vree, 1985; Simmons, 2005: 393). arge species group (Van Cakenberghe and De Vree, 1985; Simmons, 2005: 393). COMMON NAMES: Chinese: 侏 凹 脸 蝠 . Czech: rýhonos drobný. English: Dwarf Slit-faced Bat. French: Nyctère naine. German: Kleinste SchlitznasenFledermaus, Zwerg-Schlitznasenfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bj; IUCN, 2009; Monadjem et al., 2017av). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017av). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bj; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cc; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are poorly known, but it seems likely to be locally threatened by deforestation in parts of its range (Mickleburgh et al., 2008bj; IUCN, 2009; Monadjem et al., 2017av). CONSERVATION ACTIONS: Mickleburgh et al. (2008bj) [in IUCN (2009)] and Monadjem et al. (2017av) report that it is not known if the species is present within any protected areas. Further studies are needed into the distribution, general ecology and threats to this species. GENERAL DISTRIBUTION: Nycteris nana has been widely recorded in West and Central Africa, with some records from East Africa. It ranges from Côte d'Ivoire and Ghana in the west, through south Cameroon and Equatorial Guinea (Rio Muni), to the Democratic Republic of the Congo, Rwanda and Burundi, to Uganda and western Kenya. It ranges as far south as northern Angola. Generally, a lowland species, but can be found up to 2,100 m asl. A record from Tanzania, represents intermedia (J. Fahr pers. comm. in Simmons, 2005: 393). Native: Angola (Crawford-Cabral, 1989; Simmons, 2005: 393; Monadjem et al., 2010d: 548); Cameroon; Central African Republic; Congo (The Democratic Republic of the) (Van Cakenberghe and De Vree, 1985; Monadjem et al., 2010d: 548); Côte d'Ivoire (Simmons, 2005: 393); Equatorial Guinea (Simmons, 2005: 393); Ghana; Kenya (Simmons, 2005: 393); Rwanda; Sudan (Simmons, 2005: 393); Togo; Uganda. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Verschuren (1955) reported on an albino specimen caught in a hollow tree in the DRC. DETAILED MORPHOLOGY: Baculum - The baculum ranges from 2.10 to 3.36 mm in length, smaller than that of N. arge, the shaft is slender, parallel-sided and essentially straight, African Chiroptera Report 2020 the base is expanded and rounded and in lateral profile is often angled ventrally, while the tip is simple, with only a small expansion (Thomas et al., 1994: 20). POPULATION: Structure and Density:- It is considered to be a rather rare bat, colony sizes tend to be small (Mickleburgh et al., 2008bj; IUCN, 2009; Monadjem et al., 2017av). 449 organs of N. cf. nana. Reported by de Jong et al. (2011: 11). Chobar Gorge virus was mentioned by Luis et al. (2013: Suppl.) and Willoughby et al. (2017: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Burundi, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Kenya, South Sudan, Uganda. Trend:- 2016: Unknown (Monadjem et al., 2017av). 2008: Unknown (Mickleburgh et al., 2008bj; IUCN, 2009). PARASITES: HAEMOSPORIDA Schaer et al. (2015: 381) found Nycteria cf. houini in N. nana from South Sudan. Landau et al. (2012: 142) and Perkins and Schaer (2016: Suppl.) mentioned the parasite as Nycterida houini Rosin, Landau and Hugot, 1978, and also reported it from the Republic of Congo and Gabon. VIRUSES: Reoviridae Orbivirus Fomede virus - reported in the CDC’s Arbovirus Catalogue, to have been isolated in 1978 from the Republic of Guinea, from the brain and internal Figure 152. Distribution of Nycteris nana Nycteris parisii (de Beaux, 1924) *1924. Petalia parisii de Beaux, Atti Soc. ital. Sci. nat., 62: 254 (for 1923). Publication date: February 1924. Type locality: Somalia: Balli [=Ballei Uen] [00 42 N 43 04 E] [Goto Description]. Holotype: [Unknown] N 1449: ♀, skull and alcoholic. Collection date: 6 April 1922. See De Beaux (1924: 254), Allen (1939a: 70). - Etymology: In honour of Dr. Bruno Parisi, former head of the zoological section (1921 - 1927) and director (1928 1951) of the Museo Civico di Storia Naturale di Milano (see De Beaux, 1924: 255; Lanza et al., 2015: 250). 1930. Nycteris parisii: Zammarano, Le colonie Italiane, ???. (Name Combination, Current Combination) 1952. Nycteris benuensis Aellen, Mém. Soc. neuchât. Sci. nat., 8: 53. Type locality: Cameroon: Rei Bouba [ca. 08 37 N 14 09 E] [Goto Description]. 1971. N[ycteris] p[arisii] benuensis: Hayman and Hill, Mammals of Africa: Chiroptera, 18. (Name Combination) 1971. N[ycteris] p[arisii] parisii: Hayman and Hill, Mammals of Africa: Chiroptera, 18. (Name Combination) 2015. [Nycteris] parissi: Kaipf, Rudolphi and Meinig, NABU - Biodiversity Assessment in Kafa, Ethiopia, 13. (Lapsus) TAXONOMY: Included in woodi by Van Cakenberghe and De Vree (1985), Koopman (1993a: 162). Thomas et al. (1994: 22) elevated it to species rank (Simmons, 2005: 393). Rosevear (1965: 173) included N. parisii in the aethiopica group. Included in the macrotis group (Simmons (2005: 393) based on Thomas et al. (1994: 22) being closely related to N. macrotis. Simmons (2005: 393) recognises one subspecies benuensis Aellen, 1952. 450 ISSN 1990-6471 COMMON NAMES: Czech: rýhonos somálský. English: Parisi's Slitfaced Bat, Balli Slit-faced Bat, Parissi's Slit-faced Bat. French: Nyctère de Parisi. German: Parisis Schlitznasen-Fledermaus. Italian: Nitterìde di Parìsi, Nicterìde di Parìsi. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of recent information on its extent of occurrence, ecological requirements, threats and conservation status (Mickleburgh et al., 2008ap; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008ap; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004bf; IUCN, 2004). Regional None known. Somalia (Varty and Hill, 1988; Simmons, 2005: 393); Togo? (Amori et al., 2016: 217). DETAILED MORPHOLOGY: Baculum - The shaft is long, parallel-sided and essentially straight, thickening towards the base, the base is expanded, in lateral profile it is angled ventrally, while the tip is expanded and trifid with three variably developed processes (Thomas et al., 1994: 21). POPULATION: Structure and Density:- The abundance of this species is not known, and only a few specimens have ever been collected. It was last recorded in the 1980s. Presumably this is a consequence of limited survey effort within the species known range (Mickleburgh et al., 2008ap; IUCN, 2009). Trend:- 2008: Unknown (Mickleburgh et al., 2008ap; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Ethiopia, Somalia. MAJOR THREATS: The threats to this species are not known (Mickleburgh et al., 2008ap; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008ap) [in IUCN (2009)] report that it is not known if the species is present within any protected areas. Further studies are needed into the distribution, abundance, general ecology and threats to this species. GENERAL DISTRIBUTION: Nycteris parisii has been recorded from southern Somalia, south Ethiopia and from northern Cameroon, close the border with Nigeria. It is a low elevation species. Nycteris parisii parisii may be distributed in East Africa and N. p. benuensis in West Africa. Figure 153. Distribution of Nycteris parisii Native: Cameroon (Aellen, 1952; Simmons, 2005: 393); Ethiopia (Hill, 1975; Simmons, 2005: 393); Nycteris thebaica E. Geoffroy St.-Hilaire, 1818 *1818. Nycteris Thebaicus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 119, pl. 1, No 2. Publication date: 1818. Type locality: Egypt: Qena province: Thebes (near Luxor) [ca.25 41 N 32 39 E] [Goto Description]. Holotype: MNHN 889: ad ♀, skin only. Collected by: Etienne Geoffroy-Saint-Hilaire. Number 172 in Rode (1941: 235), who also mentioned number MNHN 142. - Etymology: From the masculine Latin adjective thebàicus, meaning "native or belonging to Thebes", the ancient capital of Upper Egypt, on the site of modern Luxor, the type locality of the species (see Lanza et al., 2015: 254). (Current Combination) 1820. Nycteris Geoffroyi Desmarest, Encyclopédie Méthodique, (Zoologie, Mammalogie), 1: 127. Publication date: 1820. Type locality: Senegal: 40 mi (= 60 km) east of Podor African Chiroptera Report 2020 1829. 1829. 1834. 1840. 1840. 1852. 1861. 1866. 451 [Goto Description]. - Comments: Included in macrotis by Kock (1969a: 93) [although with a question mark] and Meester et al. (1986: 35). Nycteris affinis A. Smith, Zool. Journ., 4 (16): 434. Publication date: May 1829. Type locality: South Africa: E Cape province: Grahamstown [33 17 S 26 31 E] [Goto Description]. - Comments: Type locality originally as "South Africa", restricted to "Grahamstown, eastern Cape Province, South Africa" by Roberts (1951: 69): see Kock (1969a: 98), Meester et al. (1986: 35), Gray et al. (1999: 2). Nycteris Capensis A. Smith, Zool. Journ., 4 (16): 434. Publication date: May 1829. Type locality: South Africa: SW Cape province: Swellidam [34 01 S 20 26 E] [Goto Description]. Lectotype: BMNH 1863.2.21.6: ad ♂, skin only. Collected by: A. Smith. Paralectotype: RMNH MAM.J1888-171b: ad, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters. - Comments: The type locality was originally mentioned as "Interior of South Africa, as well as upon the Eastern Coase", and restricted to "Swellidam, SW Cape Province, South Africa" by Roberts (1951: 69): see Kock (1969a: 98), Meester et al. (1986: 35), Gray et al. (1999: 2). Ansell and Dowsett (1988: 31) mention the locality as "Swellendam"). Considered a valid subspecies by Taylor (1998: 33). Nycteris thebaica: A. Smith, S. Afr. Quart. J., 2: 59. (Current Spelling) N[ycteris] albiventer Wagner, in: Schreber, Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen, Suppl. 1: 439. Type locality: Sudan: Northern province: Dongola [19 10 N 30 27 E] [Goto Description]. - Comments: Type locality originally given as "Nubia" (see Kock, 1969a: 98), restricted to Dongola, Northern Province, Sudan by Koopman (1975): see Gray et al. (1999: 2). N[ycteris] discolor Wagner, in: Schreber, Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen, Suppl. 1: 440. Type locality: South Africa: Ecklon [Goto Description]. - Comments: Allen (1939a: 68) mentioned the type locality as "Südspitze von Afrika". Kock (1969a: 98), Meester et al. (1986: 35) mention "Southern tip of Africa; Knysna, fide Roberts (1951: 69)."). Nycteris fuliginosa Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 46, pl. 10. Publication date: 1852. Type locality: Mozambique: 12 mi (19 km) NW Quellimane: Boror [17 53 S 36 51 E] [Goto Description]. Syntype: BMNH 1858.6.18.15: ad ♀. See Turni and Kock (2008: 27). Syntype: BMNH 1907.1.1.337: ad ♀, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters. See Turni and Kock (2008: 27). Syntype: ZMB 392: ad ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: March 1846. See Turni and Kock (2008: 27). Syntype: ZMB 3954: ad ♂, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: March 1846. See Turni and Kock (2008: 27). Syntype: ZMB 562: ad ♀, skull and alcoholic. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: March 1846. See Turni and Kock (2008: 27). - Comments: Turni and Kock (2008: 27) refer to Jentink (1888b: 171) in mentioning an additional syntype, which is possibly in the RMNH. Nycteris labiata Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 5. Publication date: 1861. Type locality: Eritrea: Bogos: Cheren [=Keren] [ca. 15 45 N 38 20 E] [Goto Description]. Syntype: SMNS 980a: ad ♂, alcoholic (skull not removed). Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. See Dieterlen et al. (2013: 294). Syntype: SMNS 980b: sad ♂, alcoholic (skull not removed). Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. See: Dieterlen et al. (2013: 294). Syntype: ZMB 2794a: ♂. Collected by: Martin Theodor von Heuglin; collection date: 1852. Formerly SMNS, see Turni and Kock (2008). Syntype: ZMB 2794b: ♂. Collected by: Martin Theodor von Heuglin; collection date: 1852. Syntype ex SMNS, see Turni and Kock (2008: 27). Paralectotype: RMNH MAM.27231: ad, skin and skull. Collected by: Martin Theodor von Heuglin; collection date: November 1861. See Turni and Kock (2008: 27). Turni (pers. comm., 12.04.2007) indicates that this specimen is possibly a syntype, but its collecting date (November 1861) is very close to the publication date. Turni and Kock (2008: 27) mention three additional syntypes: SMNS 980a-b, and one presumably in the RMNH (for the latter, they refer to Jentink, 1888b: 171). Plecotus æthiopicus Heuglin and Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 54 (1) 10: 546. Type locality: Sudan: Bahr el Abiat (White Nile) [ca. 14 00 N 33 00 E] [Goto Description]. - Comments: Meester et al. (1986: 35) mention Fitzinger only as author. 452 ISSN 1990-6471 1868. 1871. 1871. 1881. 1900. 1900. 1900. 1904. 1908. 1909. 1910. 1911. 1912. 1912. 1912. 1912. Nomen nudum (see Allen, 1939a: 68). Included in macrotis by Kock (1969a: 93), but rejected as nomen nudum. Nycteris Geoffroyi Var. Senegalensis Hartmann, Z. Gesel. f. Erdk., 3 (28/29): 44. Type locality: Sudan: Sennaar [15 35 N 33 38 E, 425 m] [Goto Description]. Nycteris angolensis Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 903 , fig. 5 (for 1870). Type locality: Angola: Biballe and Rio Coroca: Caconda [13 42 S 15 05 E, 1 900 m] [Goto Description]. Syntype: BMNH 1879.1.21.2:. Syntype: ZMB 3633: juv ♂, skull and alcoholic. From Rio Corvea, Angola; ex Lisbon Museum; see Turni and Kock (2008). Syntype: ZMB 3637: ♂, skull and alcoholic. Collected by: José Alberto de Oliveira Anchieta. Ex Lisbon Museum; this specimen was collected at Caconda, Biballe (see Turni and Kock (2008)). Peters (1871: 904) mentions Anchieta as collector for the type specimen. - Comments: Type locality restricted by Hill and Carter (1941): see Gray et al. (1999: 2). The publication covers a meeting held on 22 December 1870. Nycteris damarensis Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 905, fig. 7 (for 1870). Type locality: Namibia: Damaraland: Otjimbingue [22 15 S 16 10 E] [Goto Description]. Syntype: BMNH 1838.2.8.4: ad, skin and skull. Collected by: Captain Alexander; original number: 1079. Presented/Donated by: ?: Collector Unknown. Syntype: NRM. See Turni and Kock (2008: 27). Syntype: ZMB 3298: ♂, skull and alcoholic. Collected by: Missionary Hahn. See Turni and Kock (2008: 27). Syntype: ZMB 3887: ad ♂, skull and alcoholic. Collected by: Higgins. See Turni and Kock (2008: 27) [missing]. - Comments: Part of the original series is in the NRM (see Turni and Kock (2008: 27). The publication covers a meeting held on 22 December 1870. Peters (1871: 905) also mentions that the name damarensis was already used in the British Museum and in the Catalogue of Mammalia from 1843. Nycteris Revoilii Robin, Bull. Soc. Philom. Paris, (7) 5: 90. Type locality: Somalia: north of 10 ° N [Goto Description]. Holotype: MNHN ZM-MO-1881-4 (174): ad ♂, alcoholic (skull not removed). Collected by: Revoil. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1881-1 (174a): ad ♂. Collected by: Revoil. Presented/Donated by: ?: Collector Unknown. Paratype: MNHN ZM-MO-1881-3 (174b): ad ♂. Collected by: Revoil. Presented/Donated by: ?: Collector Unknown. Cotype: MCZ 14929: ♀, alcoholic (skull not removed). Collected by: Revoil. Presented/Donated by: ?: Collector Unknown. - Comments: Type locality originally "Somalia", restricted to "Somalia, north of 10 °N" by Moreau et al. (1946: 399): see Gray et al. (1999: 2). Allen (1931: 235) mentions a "cotype" in MCZ (14929), collected by Revoil in Somaliland in 1880. Nycteris thebaica var. angolensis: Seabra, J. Sci. mat. phys. nat., ser 2, 6 (22): 18. (Name Combination) Nycteris thebaica var. damarensis: Seabra, J. Sci. mat. phys. nat., ser 2, 6 (22): 19. (Name Combination) Nycteris thebaica var. Fuliginosa: Seabra, J. Sci. mat. phys. nat., ser 2, 6 (22): 18. (Name Combination) Nycteris thebaica capensis: Trouessart, Catalogus Mammalium tam viventium quam fossilium, supplement, ???. (Name Combination) Petalia capensis: Thomas and Wroughton, Proc. zool. Soc. Lond., ???. Nycteris capensis var. fuluginosa: Seabra, Ann. Scient. Acad. Politecn. Porto, 4: ???. Petalia thebaica: Thomas, Ann. Mag. nat. Hist., ser. 8, 6: ???. (Name Combination) Petalia revoili: G.M. Allen, Bull. Mus. comp. Zool., 54 (9): 322. Petalia damarensis brockmani K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): 548. Publication date: 1 November 1912. Type locality: Somalia: Upper Sheikh [4 300 ft] [Goto Description]. Holotype: BMNH 1910.3.27.4: ad ♀, skin and skull. Collected by: Dr. Ralph Evelyn Drake-Brockman; collection date: 11 January 1910. Petalia damarensis damarensis: K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): ???. Petalia damarensis media K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): 548. Publication date: 1 November 1912. Type locality: Ethiopia: Harar province: Harer [09 18 N 42 08 E, 2 000 m] [Goto Description]. Holotype: BMNH 1912.2.28.1: ad, skin and skull. Collected by: G. Kristensen; collection date: 19 January 1912. Presented/Donated by: Nathanial Charles Rothschild. Petalia thebaica thebaica: K. Andersen, Ann. Mag. nat. Hist., ser. 8, 10 (59): ???. (Name Combination) African Chiroptera Report 2020 1918. 1923. 1933. 1934. 1934. 1934. 1936. 1936. 1938. 1939. 1944. 1956. 1971. 1975. 1998. 1998. 1999. 2018. 2019. 2019. 2019. 2019. 2019. 2019. 2019. 2019. 2019. ? ? ? ? 453 Nycteris thebaica thebaica: Wettstein, Denkschr. k. Akad. Wiss. Wien, Math.-Naturw. Kl., 94: ???. (Name Combination) Petalia (Nycteris) thebaica aurantiaca de Beaux, Atti Soc. ital. Sci. nat., 62: 91. Publication date: July 1923. Type locality: Kenya: Northern Guaso Nyiro: Archer's Post [00 39 N 37 41 E, 2 000 ft] [Goto Description]. Holotype: [Unknown] N 1432: ♀, skull and alcoholic. Collected by: L. Franchetti and L. Tonker; collection date: January 1920. Nycteris damarensis damarensis: G.M. Allen and Loveridge, Bull. Mus. comp. Zool., 75: ???. Nycteris capensis capensis: Shortridge, Mammals of South West Africa, 55. (Name Combination) Nycteris capensis damarensis: Shortridge, Mammals of South West Africa, 56. P[etalia (Nycteris)] theb[aica] adana: de Beaux, Atti Soc. ital. Sci. nat., 62: 93. (Name Combination) Nycteris damarensis brockmani: G.M. Allen and Lawrence, Bull. Mus. comp. Zool., 79 (3): 35, 49. Nycteris thebaica revoili: G.M. Allen and Lawrence, Bull. Mus. comp. Zool., 79 (3): 35, 49. (Name Combination) Nycteris thebaica aurantiaca: Frechkop, Exploration du Parc national Albert, Fasc. 10: ???. (Name Combination) Nycteris damarensis media: G.M. Allen, Bull. Mus. comp. Zool., 83: 69. Nycteris thebaica albiventer: Aharoni, Bull. Zool. Soc. Egypt, 6: 26. (Name Combination) Nycteris thebaica brockmani: Toschi, Atti Soc. ital. Sci. nat., 95: ???. (Name Combination) Nycteris thebaica labiata: Hill and Morris, Bull. Br. Mus. (nat. Hist.) Zool., 21 (2): 34. (Name Combination) Nycteris thebiaca: Koopman, Bull. Am. Mus. Nat. Hist., 154 (4): 379. - Comments: Lapsus (actually Koopman uses this name consequently in this publication). (Lapsus) Nyteris angolensis: Van Cakenberghe and De Vree, Bonn. zool. Beitr., 48 (2): 140. Publication date: October 1998. (Lapsus) Nyteris thebaica: Van Cakenberghe and De Vree, Bonn. zool. Beitr., 48 (2): 123. Publication date: October 1998. (Name Combination, Lapsus) Nycteris thabaica labiata: Van Cakenberghe and De Vree, Bonn. zool. Beitr., 48 (2): 147. (Lapsus) Nycteris thebica capensis: Gunnell and Manthi, J. Hum. Evol., Suppl.. Publication date: 6 April 2018. (Lapsus) Nycteris cf. thebaica 1: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris cf. thebaica 2: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 2023. Publication date: 20 August 2019. Nycteris cf. thebaica 3: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1026. Publication date: 20 August 2019. Nycteris thebaica 1: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris thebaica 2: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris thebaica 3: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris thebaica 4: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris thebaica 5: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023. Publication date: 20 August 2019. Nycteris thebaica 6: Demos, Webala, Kerbis Peterhans, Goodman, Bartonjo and Patterson, J. Zool. Syst. Evol. Res., 57 (4): 1023.. Publication date: 20 August 2019. Nycteris capensis: Nycteris thebaica angolensis: (Name Combination) Nycteris thebaica damarensis: (Name Combination) Nycteris thebaica revoilii: (Alternate Spelling) 454 ISSN 1990-6471 TAXONOMY: Bat. French: Nyctère de la Thébaïde, Nyctère égyptienne. German: Großohrhohlnase, Ägyptische Schlitznasen-Fledermaus, Stirngrubenfledermaus. Hebrew: ‫לילן‬, leilan, Leylan. Italian: Nitterìde tebàna, Nicterìde tebàna, Nitterìde di Tèbe, Nicterìde di Tèbe. Portuguese: Morcego orelhudo do Egipto. Kiluba (DRC): Kasusu. Yao (Malawi): Liputiputi (applied to all small bats). ETYMOLOGY OF COMMON NAME: So named because they were first described from tombs in Thebes, Egypt (see Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: N. thebaica was found in upper Pleistocene samples at Border Cave, South Africa (Avery, 1991: 6). Butler (1984: 185) reported on a distal end of a humerus found at Chamtwara (Kenya). Figure 154. Nycteris thebaica (SMG-15760) caught at Marloth Park, Mpumalanga, South Africa. Meester et al. (1986) state that the subspecies classification of thebaica is uncertain. Hayman and Hill (1971), while acknowledging that the species may include valid subspecies, do not attempt to recognize any. Kock (1969a), however, lists seven subspecies, and Koopman (1975), while not providing a complete synonymy of the species, appears to come to broadly similar conclusions to Kock concerning those names that he does consider. Simmons (2005: 393) recognises seven subspecies - adana K. Andersen, 1912; angolensis Peters, 1870; brockmani K. Andersen, 1912; capensis A. Smith, 1829; damarensis Peters, 1870; labiata Heuglin, 1861; and najdiya Nader and Kock, 1982. Van Cakenberghe and De Vree (1999a) suggest that the status of brockmani and damarensis remains unclear as these forms may represent distinct species (Harrison and Bates, 1991; Gray et al., 1999). thebaica species group (Van Cakenberghe and De Vree, 1999a; Simmons, 2005: 393). COMMON NAMES: Afrikaans: Gewone-spleetneusvlermuis, Egiptiese spleetneusvlermuis, Kaapse Langoorvlermuis. Arabian: Khafash Teiba, Khaffash. Castilian (Spain): Rinolofo egipcio. Chinese: 非洲凹脸蝠. Czech: rýhonos egyptský, šerowec tébský, dutonosec, netopýr koptový, nykteris egyptská. Dutch: Thebaanse spleetneusvleermuis. English: Egyptian Slit-faced Bat, Common Slitfaced Bat, Geoffroy's Nycteris, Cape Long-eared CONSERVATION STATUS: Global Justification This species has a large range and faces no major threats, hence is listed as Least Concern LC ver 3.1 (2001) (Mickleburgh et al., 2008bk; IUCN, 2009; Monadjem et al., 2017aw). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017aw). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bk; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ce; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016d). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: Habitat destruction and degradation affect the species. Roost disturbance and degradation are the main threats in the Mediterranean region. These are not considered major threats at present (Mickleburgh et al., 2008bk; IUCN, 2009; Monadjem et al., 2017aw). In the Limpopo province, South Africa, Toussaint and McKechnie (2012: 1138) indicate that the cool baobab tree cavities appear to be critical for the presence of N. thebaica in that area and may buffer this species from future increases in extreme maximum ambient temperatures. Aerial hawking and water stress are believed to be the major risk factors linked to climatic change (Sherwin et al., 2012: 174). African Chiroptera Report 2020 Collinson et al. (2015: 311) report on two roadkills in the Greater Mapungubwe Transfrontier Conservation Area (GMTFCA), South Africa. CONSERVATION ACTIONS: Mickleburgh et al. (2008bk) [in IUCN (2009)] and Monadjem et al. (2017aw) report that it may occur within some protected areas. No specific conservation measures are known. Protection of the roost sites is required and also legal protection of the species should be improved. Further research is required on the population size and trends. GENERAL DISTRIBUTION: Nycteris thebaica is broadly distributed across savanna and riparian zones. Mostly found in subSaharan Africa; also found in Morocco, Libya, Egypt (primarily down the Nile River valley, but also into Sinai) and the Middle East (Israel, Palestine and Jordan). Recorded form the islands of Unguja [=Zanzibar] and Pemba [see Simmons (2005: 393), although not from the latter by O'Brien (2011: 289)]. Elevation from sea level to 2,000 m. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,704,632 km2 (covering 27 % of the total surface). In South Africa, its distribution is mainly affected by the mimimum temperature of the coldest month (Babiker Salata, 2012: 49). Corbet (1984: 8) states that the inclusion of Corfu (Greece) in the range could be the result of a vagrant specimen from N Africa (for details see Van Cakenberghe and De Vree, 1999a). Native: Angola (Crawford-Cabral, 1989; Cotterill, 2004a: 261; Monadjem et al., 2010d: 548); Benin (Simmons, 2005: 393); Botswana (Archer, 1977; Monadjem et al., 2010d: 548); Burkina Faso (Simmons, 2005: 393; Kangoyé et al., 2015a: 617); Burundi; Cameroon; Central African Republic; Chad; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 548); Côte d'Ivoire; Djibouti (Pearch et al., 2001: 397; Simmons, 2005: 393); Egypt (Simmons, 2005: 393); Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 133); Gambia; Ghana (Simmons, 2005: 393); Guinea (Konstantinov et al., 2000: 144; Simmons, 2005: 393; Decher et al., 2016: 264); Guinea-Bissau; Israel (Simmons, 2005: 393); Jordan (Al-Omari et al., 2000: 5); Kenya (Simmons, 2005: 393); Libyan Arab Jamahiriya; Malawi (Ansell and Dowsett, 1988: 31; Happold et al., 1988; Monadjem et al., 2010d: 548); Mali (Simmons, 2005: 393); Morocco (Simmons, 2005: 393; Benda et al., 2010a: 157; El 455 Ibrahimi and Rguibi Idrissi, 2015: 358); Mozambique (Smithers and Lobão Tello, 1976; Lopes and Crawford-Cabral, 1992; Monadjem et al., 2010d: 548; Monadjem et al., 2010c: 382); Namibia; Niger (Simmons, 2005: 393; Monadjem et al., 2010d: 548); Nigeria (Simmons, 2005: 393); Palestinian Territory, Occupied; Rwanda; Saudi Arabia; Senegal (Simmons, 2005: 393); Sierra Leone; Somalia (Simmons, 2005: 393); South Africa (Simmons, 2005: 393; Monadjem et al., 2010d: 549); Sudan; Swaziland (Monadjem, 2001, Monadjem, 2006b; Simmons, 2005: 393; Monadjem et al., 2010d: 549); Tanzania (Stanley and Goodman, 2011: 43); Togo (Amori et al., 2016: 218); Uganda (Kityo and Kerbis, 1996: 60); Yemen; Zambia (Ansell, 1969; Ansell, 1973; Ansell, 1978; Ansell, 1986; Monadjem et al., 2010d: 549); Zimbabwe (Monadjem et al., 2010d: 550). Presence uncertain: Lebanon; Lesotho; Liberia; Mauritania. Bates et al. (2013: 338) reject the presence of N. thebaica in the Republic of Congo as the locality (Kemo) mentioned by Pousargues (1896: 259) and Van Cakenberghe and De Vree (1999a: 143) is actually located in the Central African Republic. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Schoeman and Jacobs (2002: 157) mention the following wing parameters for 8 specimens from the Algeria Forestry Station (RSA): Fa: 48.2 ± 0.8 mm, Wing area: 155.4 ± 7.4 cm 2, Wing span: 28.7 ± 1.1 cm, Wing loading: 8.3 ± 0.9 N/m 2, and aspect ratio: 5.3 ± 0.3. Four piebald (all-white fur/skin patches, eyes always normally coloured) specimens were reported from a mine in Tanzania (Howell, 1980; Lucati and López-Baucells, 2016: Suppl.). DETAILED MORPHOLOGY: Baculum - The baculum ranges from 2.49 to 2.97 mm in length, the shaft is slender and parallelsided, the base is expanded, in lateral profile it is usually angled ventrally, while the tip is simple with a small expansion (Thomas et al., 1994: 22). Brain - From specimens collected in Bénin, Amrein et al. (2007) reported that proliferating cells, detected with Ki-67 and MCM2, in the subgranular layer of the dentate gyrus was found to be absent. Amrein et al. (2007) also reported moderate to ample proliferating cells (Ki-67 positive cells) and migrating young neurons (DCX positive cells) in the rostral migratory stream. Amrein et al. (2007) could not detect any antigen in the antibody against DCX, and no NeuroD was detected in the hippocampal granule cells. 456 ISSN 1990-6471 Reproduction system - Carter and Enders (2016: 284) found that N. thebaica has an extensive smooth chorioallantois, and its luminal epithelium is incomplete. Gastro-intestinal tract (GIT): Aylward et al. (2019: 1108) reported for six specimens an average weight of 10.17 ± 1.71 g and a GIT of 0.61 ± 0.14 g (6.02 ± 1.31 %). FUNCTIONAL MORPHOLOGY: Zhuang et al. (2012: 1) showed that the pit in the noseleaf of Nycteris thebaica has an effect on focusing the acoustic near field as well as shaping the radiation patterns and therefore enhancing the directionality. Toussaint et al. (2012) investigated the patterns of heterothermy exhibited by N. thebaica during the austral winter in a tropical semi-arid environment. They expected that it would use pronounced heterothermy during the warm, dry winter months, but found this not to be the case. As possible reasons for this, they suggest the relatively high risk of predation, the communal roosting, the species' reproduction cycle (although they find this highly unlikely as the results for both sexes were similar), and the relatively long nightly foraging period. SEXUAL DIMORPHISM: In Swaziland, Monadjem (2001: 49) found that females were heavier and had longer forearms than males. ECHOLOCATION: Schoeman and Waddington (2011: 291) mention a peak frequency of 90.3 ± 0.7 kHz and a duration of 1.7 ± 0.1 msec for specimens from Durban, South Africa. Davies et al. (2013b: Table S8) report a peak frequency of 88.23 kHz and a range between 55.97 and 75.5 kHz. However, also in the Durban area, Naidoo et al. (2011: 23) mention a peak frequency of 65.8 ± 20.1 kHz, and a duration of 1.7 ± 1.0 msec. From the Algeria Forestry Station (RSA), Schoeman and Jacobs (2002: 157) report the type of call to be "Low duty echolocation dominated by frequency modulated calls", with Fpeak: 77.4 ± 2.7 kHz, and a low intensity. Parker and Bernard (2018: 57) recorded one call at the Mapungubwe National Park (RSA): Fchar: 62.50 kHz, Fmax: 83.40 kHz, Fmin: 62.40 kHz, Fknee: 63.10 kHz, duration: 1.68 msec and 11.63 calls/sec. Although very common in their study area in Swaziland, Monadjem et al. (2017c: 181) were unable to detect this species using an Anabat Express Detector due to its whispering (low db) calls. Even at distances of 0.1 m, they could not detect a call. For Morocco, Disca et al. (2014: 226) recorded the following information: call type: FM multi H, Fstart: 137.0 ± 1.2 kHz, Fend: 19.6 ± 0.5 kHz, Fpeak: 84.5 ± 3.9 kHz, Fmax all harmonics: 27.9 ± 0.2 kHz, Fmax first harmonic: 56.1 ± 0.7 kHz, Fmax third harmonic: 105.4 ± 3.2 kHz, bandwidth: 117.4 ± 1.5 kHz, duration: 2.4 ± 0.3 msec. This information is based on one single sequence they were not able to identify in hand. Hackett et al. (2016: 223) reported the following values for 13 calls from Israel: Pulse duration: 1.35 ± 0.13 msec, Fstart: 78.45 ± 3.08 kHz, Fend: 63.20 ± 1.99 kHz, Fpeak: 70.18 ± 2.00 kHz. Luo et al. (2019a: Supp.) reported the following data (Free flying bats): F peak: 70.18 kHz, Fstart: 78.45 kHz, Fend: 63.2 kHz, Band width: 15.25 kHz, and duration: 1.35 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Both Peterson and Nagorsen (1975) and Rautenbach et al. (1993) reported 2n = 42, FN = 78, BA = 38, with a submetacentric X chromosome, and a metacentric Y chromosome. Protein / allozyme - Unknown. HABITAT: This species has a wide distribution and is found in most habitats throughout its range. Cooper-Bohannon et al. (2016: Table S2) indicate that - in southern Africa - it occurs primarily in Savanna (63 %) areas, followed by SW arid (18 %) and Afromontane (10 %) habitats. Based on the study of stable isotope ratios of carbon, nitrogen and hydrogen, Voigt et al. (2013: 755) suggest that N. thebaica might potentially be a migratory bat species, as far as elevational migration along the slopes of Mount Kilimandjaro is concerned. However, they also point out that the very low wing loading and aspect ratio is atypical for a migratory species, and therefore still categorize it as a sedentary species. Foord et al. (2018: 5), furthermore, indicate that N. thebaica moves less than one km per night between its roost and foraging sides. HABITS: In Swaziland male N. thebaica exhibit less roost site fidelity than females: there is greater roost movement in males than females (Monadjem, 2006b). Stanley et al. (2007b: 209) report on a specimen in a Victor Rat trap, because it probably chased insects attracted to the bait used in the trap. African Chiroptera Report 2020 Thomas and Jacobs (2013: 125) found that, at De Hoop Nature Reserve (South Africa), N. thebaica emerged significantly earlier during mid-summer than during early and late summer. They also reported that bats feeding on Lepidoptera (N. thebaica and Rhinolophus capensis) emerged earlier than bats feeding on Diptera (Tadarida aegyptiaca). ROOST: Roosting in caves, mine audits and various other hollow sites (Churchill et al., 1997), road culverts (Monadjem, 1998a; 2001; 2006b), under houses (LaVal and LaVal, 1980; Seamark and Bogdanowicz, 2002). Avery et al. (1990: 7) indicate that cool and moist caves or cave-like structures are used as daytime roosts. In Swaziland, Monadjem et al. (2015: 15) found that roost switching occurs frequently, with more transitions to roosts where large numbers of bats were staying. They also found that moving to another roost was related to the distance of the new roost from the original one. MIGRATION: The species is not known to undergo migration (Monadjem, 2006a), and Laycock (1973) and Wingate (1983) suggest that N. thebaica does not have any homing ability. Monadjem (2006b) states that bat numbers vary greatly with season, suggesting that some movement is taking place. Monadjem (2006b) also reported that a banded female in Swaziland was discovered 107 km from where it was originally banded. DIET: Casual observations in Israel (Makin, 1979), Sudan (McLellan, 1986) and Namibia (Felten, 1956) reported that scorpions formed a stable part of N. thebaica diet. Yerbury and Thomas, 1895) examined cull-parts in Arabia, identifying various species of Orthoptera. Whereas Chapman (1958, in Tanzania), Roer (1975, in Namibia), and LaVal and LaVal (1980, in South Africa) found that Orthoptera, Lepidoptera, and Coleoptera were the primary prey orders consumed by N. thebaica. LaVal and LaVal (1980) presented results for early autumn (April), mid-winter (July), spring (November), and summer (January to March), listing a monthly breakdown of all prey orders consumed and finding that Orthoptera predominated in the study. In Zimbabwe, Fenton (1975) combined the results of his wet season collection (November 1969 to March 1970), finding only Orthoptera and Lepidoptera wings. Fenton et al. (1977a) and Fenton and Thomas (1980) examined the faeces of N. thebaica in Zimbabwe, and found that in the wet season (January and 457 February 1976) Lepidoptera, Diptera and Coleoptera formed the bats' diet, while in the dry season (June 1977) he found only Lepidoptera (100 percent volume). In Zambia, Whitaker and Black (1976) examined the stomach contents of 33 N. thebaica which they separated into summer (November to March) and winter (May to September) categories. They found similar prey orders in N. thebaica diet, but also found many wingless insects. At Farm Laatstgevonden, South Africa, Taylor et al. (2012b: 62) found the following prey groups: Lepidoptera (31.5 %), Coleoptera (24.75 %), Hemiptera (19.5 %), Orthoptera (12.75 %), and other (11.5 %). Bowie et al. (1999) at Mkuzi Game Reserve, KwaZulu Natal, South Africa, found that the diet consisted mainly of non-volant prey, primarily orthopterans and arachnids. Feldman et al. (2000), in Israel, found they fed on Lepidoptera and Diptera. Seamark and Bogdanowicz (2002) showed that prey eaten by N. thebaica varies significantly by order but not by season, and there was a significant interaction between the prey category and season, suggesting that these two factors are not independent from each other. Coleoptera (49.6 %) in the culled parts, (calculated as percent composition) dominated in spring (SeptemberNovember), Orthoptera (38.8 %) in summer (December-February), Hemiptera (42.8 %) in autumn (March-May) and Lepidoptera (36.3 %) in winter (June-August). The diet also included a frog and small fish. From the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) reported the following prey groups based on 51 faecal pellets from 8 bats (in volume percent): Lepidoptera (34.9 ± 27.3), Orthoptera (23.5 ± 19.6), Coleoptera (21.3 ± 17.6), Arachnida (13.5 ± 27.4), Diptera (3.8 ± 10.6), Hymenoptera (1.6 ± 4.6), and Hemiptera (1.5 ± 4.5). Weier et al. (2019b: 6) examined droppings from this species in the Levubu region (Limpopo, RSA) and found DNA sequences of Cryptophlebia peltastica (Meyrick, 1921) (Lepidoptera) and Nezara viridula (Linnaeus, 1758) (Hemiptera). Using Next Generation Sequencing (NGS), (Taylor et al. (2017b: 243, 252 - 254) reported on the 28 food items found in faecal pellets in the Limpopo province (RSA): Lepidoptera (occurred in 100 % of analysed pellets - Toxocampinae tathorhynchus + Eutelia polychorda + Acanthovalva inconspicuaria + Isturgia roraria + Scopula rufisalsa + Hyblaea puera + Sena sp. + Sena prompta + Enmonodia capensis + Leucania BioLep10 + Pandesma robusta + Pericyma atrifusa + Argyrogrammatini 458 ISSN 1990-6471 sp. + Thysanoplusia indicator + Tycomarptes sp. + Garella nilotica + Maurilia arcuata + Phycitinae sp. + Arotrura JFL201 + unmatched), Blattodea (50 % - Pycnoscelus + unmatched), Orthoptera (33 %), Coleoptera (17 % - Cybister tripunctatus + unmatched); Hemiptera (17 % - Macrorhaphis acuta + Nezara viridula); Orthoptera (Tettigoninae sp.). PREDATORS: Perrin (1982: 16) and Avery et al. (2005: 1054) found this species in pellets from Tyto alba in South Africa. Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure:- Monadjem (2001: 50) mentions that sex ratio was highly skewed with adult females outnumbering adult males by 2.8:1, even though the sex ratio of juveniles was 1:1. Sex ratios of adults varied seasonally, being highly skewed in July/August, October and December. Monadjem (2001: 50) suggests that this might be due to the females being resident and males joining them in late summer to mate. Density:- Fairly common through most of its range. However, an uncommon species in the Mediterranean region. In Morocco only three small colonies (usually 10 to 30 and occasionally up to 100 individuals) are known (Stéphane Aulagnier pers. comm. 2007 in Mickleburgh et al., 2008bk; IUCN, 2009; Monadjem et al., 2017aw). Trend:- 2016: Unknown (Monadjem et al., 2017aw). 2008: Unknown (Mickleburgh et al., 2008bk; IUCN, 2009). LIFESPAN: In Swaziland, Monadjem (2006b) found that for both males and females, survivorship was low in the first year, but increased thereafter. While 15 % of females and 10 % of males banded as juveniles in 1998 and 1999 survived to three years of age. Monadjem (2006a) reported the age of N. thebaica in Swaziland to reach at least 7 to 8 years. Monadjem et al. (2015) studied the effect of time, age and sex class and climatic variables on survival from 1450 marked individuals from 16 roosts over a 10-year period. They found that the annual survival rate was highest in adult males and lowest in juvenile females. Climatological factors did not seem to have an influence. ACTIVITY AND BEHAVIOUR: Female N. thebaica in Swaziland appear to be resident year-round in the culverts, which suggests that males join females in late summer for the purpose of mating (Monadjem, 2001). Lindeque (1987) describes the inflight behaviour of mating, where the active pursuit of the female by the male bat can be equated to courtship phase, with the butting phase inducing the female bat to hover and allowing the male to copulate, the brief frontal approach and head-neck biting by the male could be relict of some incomplete behaviour sequence. Korine et al. (2015: 5) mention that the activity of N. thebaica could be underestimated because of the whispering nature of its echolocation calls. REPRODUCTION AND ONTOGENY: Martin and Bernard (2000: 32) mention that the corpus luteum is only present for the first half of pregnancy, and is subsequently completely regressed (Badwaik and Rasweiler, 2000: 276). Bernard (1982a) [in Krutzsch (2000: 114)] indicated that this species is monoestrous in the subtropical, temperate regions of South Africa with a longer than usual gestation of five months and a three-month anoestrus. Male and female cycles are synchronized; spermatogenesis commences in late summer and the epididymides contain mature sperm from autumn to summer (see Bernard, 1976). In Malawi, females are in reproductive synchrony, give birth at the end of the dry season and lactate their young for at least 1.5 months (Happold and Happold, 1990b: 566). A pregnant female was collected on 22 July 1992 in the West Usambara Mountains (Tanzania) (Stanley and Goodman, 2011: 43). MATING: Mating system is not known (Monadjem, 2001). PARASITES: BACTERIA Four out of 49 specimens tested at Mlawula (Swaziland) by Dietrich et al. (2016b: 3) tested positive for Bartonella and one was positive for Rickettsia. Dietrich et al. (2016a: 4) found Leptospira in the urine of a bat from South Africa, and Coxiella in its faeces. For the RSA, these bacteria were further identified as Leptospira borgpetersenii, whereas in bats from Swaziland sequences belonging to L. interrogans were found. FUNGI Brennan et al. (2015: 145) found no antibodies against Pseudogymnoascus destructans (White- African Chiroptera Report 2020 nose syndrome) in blood samples from bats from Botswana. HAEMOSPORIDA Schaer et al. (2015: 381) found Nycteria sp. parasites in N. thebaica from South Sudan. Garnham and Heisch (1953: 357) described Nycteria medusiformis from a "Nycteris capensis". 24 of the 36 southern African N. thebaica specimens reported by Clément et al. (2019: 5) were infected by Trypanosoma cf. livingstonei_likeA, likeB (5), likeC (5), likeD, likeE, likeF, likeG, likeI, likeJ, likeK (3), likeL, likeM, likeN (2). NEMATODA Spiruridae: An infection by Spirura nycterisi Khalil, 1975 in Tanzania was reported by PeraltaRodríguez et al. (2012: 1006). DIPTERA Streblidae: Raymondia alulata Speiser, 1908, for which N. thebaica is the typical host, occurs in many parts of southern Africa from the Cape northwards to the Kaokoveld in Namibia and eastwards to Bukawayo in Zimbabwe (Haeselbarth et al., 1966: 102). Shapiro et al. (2016: 249) reported it for the first time from Swaziland. Raymondia hardyi Fiedler, 1954 from South Africa (Shapiro et al., 2016: 254). Raymondia planiceps Jobling, 1930 (Haeselbarth et al., 1966: 102). Raymondia seminuda Jobling, 1954 from southern Botswana (Haeselbarth et al., 1966: 103; Shapiro et al., 2016: 255). Nycteribiidae: Nycteribia scissa rhodesiensis Theodor, 1957 from Zimbabwe and Malawi (Haeselbarth et al., 1966: 108). Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114). SIPHONAPTERA Pulicidae: Echidnophaga aethiops Jordan and Rothschild, 1906 described from Klipfontein, Namaqualand, South Africa and also found in Namibia, Botswana, Tanzania, Kenya and Somalia (Haeselbarth et al., 1966: 135). Ischnopsyllidae: Oxparius isomalus (Waterston, 1915) locality not stated (Haeselbarth et al., 1966: 188). ACARI Laelapidae: Metaspinolaelaps aelleni Till ([May] 1958) was described from Nycteris thebaica capensis from Zululand. Till (1960: 223) suggests that this possibly might be a synonym of Bewsiella fledermaus Domrow, [March] 1958, but this view doesn't seem to be followed (e.g. https://en.wiki2.org/wiki/Metaspinolaelaps). 459 Macronyssidae: Steatonyssus longipes Radovsky and Yuker, 1953 was described from an Egyptian slit-faced bat (Negm et al., 2018: 728). Trombiculidae: Vercammen-Grandjean and Fain (1958: 24) mention the presence of Trombigasta (Trombigastia) mounti (Radford, 1954) on "N. capensis damarensis" in Asmara, Eritrea. Stekolnikov (2018a: 50, 119, 160, 173) reported Whartonia oweni Vercammen-Grandjean and Brennan, 1957, Trombigastia mounti (Radford, 1954) (from "N. thebaica damarensis"), Microtrombicula minutissima (Oudemans, 1910) (from "N. thebaica capensis"), Microtrombicula tanzaniae Goff, 1982. VIRUSES: Astroviridae Four out of 14 Mozambican bats tested by Hoarau et al. (2018: 2) were positive for this group of viruses. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in six individuals from Limpopo Province, South Africa, none were tested positive (0/6). Paramyxoviridae Mortlock et al. (2015: 1841) reported that one out of twelve examined South African N. thebaica specimens tested positive for Paramyxovirus sequences. Rhabdoviridae Lyssavirus - Rabies related viruses Duvenhage Virus: In 1986, Foggin (1988) recovered the Duvenhage (DUVV) virus from a Nycteris thebaica in Zimbabwe (see Paweska et al., 2006; Hayman et al., 2011a: 88; van Eeden et al., 2011: 67). Markotter et al. (2013: 1000) report on antibodies for DUVV that were recovered from bats from Swaziland. Rabies: Oelofsen and Smith (1993) tested 12 individual brains, from three localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burkina Faso, Cameroon, Central African Republic, Congo (Democratic Republic of the), Djibouti, Egypt, Eritrea, Eswatini, Ethiopia, Ghana, Guinea, Kenya, Malawi, Mali, Morocco, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Uganda, Zambia, Zimbabwe. 460 ISSN 1990-6471 Figure 155. Distribution of Nycteris thebaica Nycteris vinsoni Dalquest, 1965 *1965. Nycteris vinsoni Dalquest, J. Mamm., 46 (2): 254, 256. Publication date: May 1965. Type locality: Mozambique: 212 km SSW Beira, S bank Save river: Zinave [21 26 S 33 52 E] [Goto Description]. Holotype: KU 105221: ad ♀, skin and skull. Collected by: Prof. Dr. Walter Woelberg Dalquest; collection date: 8 October 1963; original number: MWUC 18739. Presented/Donated by: ?: Collector Unknown. See Jones and Genoways (1969: 130). - Etymology: In honour of J. Vinson, who sponsored an expedition to Mozambique during which the first specimens were collected (see Dalquest, 1965; Taylor, 2005). (Current Combination) TAXONOMY: Bronner et al. (2003) state that is was originally described from Mozambique (Dalquest, 1965). Kock (1969a) considered Nycteris vinsoni a synonym of N. macrotis luteola, based on the size and position of the second lower premolar. Hayman and Hill (1971) suggest that N. vinsoni is a variant of N. aethiopica, itself considered a member of the N. macrotis group (Koopman, 1965; Kock, 1969a). Koopman (1975) argued that the size and position of the second lower premolar is extremely variable, and instead recognised vinsoni as a distinct species closely related to N. thebaica, based on the presence of a pyriform tragus (and see Koopman, 1982). He later concluded that the tragus of the holotype is actually semilunate and therefore synonymised vinsoni under N. macrotis oriana (Koopman, 1992, 1993a). Swanepoel et al. (1980),Meester et al. (1986), Corbet and Hill (1992) and Simmons (2005: 394) retained vinsoni as a distinct species, a treatment endorsed by limited morphometric comparisons (Van Cakenberghe and De Vree, 1999a). thebaica species group (Van Cakenberghe and De Vree, 1999a; Simmons, 2005: 394). COMMON NAMES: Afrikaans: Vinson se spleetneusvlermuis. Czech: rýhonos mosambický. English: Vinson's Slitfaced Bat, Mozambian Slit-faced Bat. French: Nyctère de Vinson, Nyctère du Mozambique. German: Vinsons Schlitznasen-Fledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient in view of the absence of recent information on its extent of occurrence, ecological requirements, threats and conservation status (Mickleburgh et al., 2008aq; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh 2008aq; IUCN, 2009). 2004: DD ver 3.1 (Mickleburgh et al., 2004be; IUCN, 2004). LR/lc (Hilton-Taylor, 2000); CR (IUCN, 1996: CR [assessed as Nycteris macrotis] and Groombridge, 1996). et al., (2001) 2000: 2000). (Baillie Regional None known. MAJOR THREATS: The threats to this species are not known (Mickleburgh et al., 2008aq; IUCN, 2009). African Chiroptera Report 2020 CONSERVATION ACTIONS: Mickleburgh et al. (2008aq) [in IUCN (2009)] report that the collection locality appears to be within Zinave National Park, Mozambique. Further studies are needed to better determine the geographic range, natural history and possible threats to this poorly known bat. 461 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Mozambique. GENERAL DISTRIBUTION: Nycteris vinsoni is known only from the type locality of "the south bank of the Save River, 212 km SSW Beira, Mozambique" (Dalquest, 1965). Native: Mozambique (Dalquest, 1965; Simmons, 2005: 394). POPULATION: Structure and Density:- It is known only from the type series of two animals (Mickleburgh et al., 2008aq; IUCN, 2009). Figure 156. Distribution of Nycteris vinsoni Trend:- Unknown (Mickleburgh et al., 2008aq; IUCN, 2009). Nycteris woodi K. Andersen, 1914 *1914. Nycteris woodi K. Andersen, Ann. Mag. nat. Hist., ser. 8, 13 (78): 563. Publication date: 1 June 1914. Type locality: Zambia: Chilanga [15 33 S 28 11 E, 4 100 ft] [Goto Description]. Holotype: BMNH 1914.4.22.2: ad, skin and skull. Collected by: Rodney Carrington Wood Esq. Collection date: November 1913. - Etymology: In honour of Rodney C. Wood, a well-known naturalist, who collected the original specimens at Chilanga, near Lusaka, Zambia (see Smithers, 1983: 118; Taylor, 2005). (Current Combination) 1946. Nycteris woodi sabiensis Roberts, Ann. Transv. Mus., 20 (4): 304. Type locality: Zimbabwe: Melsetter district: Sabi River: Birchenough Bridge [19 54 S 32 14 E] [Goto Description]. Holotype: TM 8578: ad ♀, skin and skull. Collected by: A.G. White; collection date: 11 January 1938. (Name Combination) 1946. Nycteris woodi woodi: Roberts, Ann. Transv. Mus., 20 (4): 304. (Name Combination) 2018. Nycteris woodii: Parker and Bernard, Mammalia, 83 (1): 53. Publication date: 14 April 2018. (Lapsus) TAXONOMY: Van Cakenberghe and De Vree (1985) included parisii and benuensis. Thomas et al. (1994) elevated parisii to full species rank, followed by Simmons (2005: 393), who included benuensis as a subspecies of parisii. Simmons (2005: 394) recognised one subspecies sabiensis Roberts, 1946. COMMON NAMES: Afrikaans: Wood se spleetneusvlermuis, Woodspleetneusvlermuis, Woodse Langoorvlermuis. Czech: rýhonos jihoafrický. English: Wood's Slitfaced Bat, Wood's Long-eared Bat. French: Nyctère de Wood. German: Woods Schlitznasen-Fledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in some protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ab; IUCN, 2009; Monadjem et al., 2017w). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017w). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008ab; IUCN, 2009). 2004: NT ver 3.1 (2001) 462 ISSN 1990-6471 (Mickleburgh et al., 2004ap; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional South Africa:- 2016: NT B1ab(ii,iii,iv, v) ver 3.1 (2001) (Schoeman et al., 2016b). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: It is possibly locally threatened in parts of its range by habitat loss (particularly through conversion of land to cotton farming), pesticide use, and disturbance of roosting sites (Mickleburgh et al., 2008ab; IUCN, 2009; Monadjem et al., 2017w). CONSERVATION ACTIONS: Mickleburgh et al. (2008ab) [in IUCN (2009)] and Monadjem et al. (2017w) report that it has been recorded from some protected areas (e.g. Kruger National Park, South Africa). Additional studies are needed into the distribution, natural history and possible threats to this poorly known species. GENERAL DISTRIBUTION: Nycteris woodi has been recorded from Zimbabwe, South Africa, to Zambia, north-west Mozambique and Malawi. It may range into southwestern Tanzania, however, it has yet to be recorded in this country. It is generally a lowland species. South of 8° S, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 308,190 km2. ECHOLOCATION: See Taylor (1999b). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported 2n = 42, FN = 78, BA = 38, a metacentric X chromosome, and an acrocentric Y chromosome, for a specimen from South Africa. Protein / allozyme - Unknown. POPULATION: Structure and Density:- The abundance of this species is not well known. It has been recorded in colonies of a few dozen individuals, otherwise animals have been recorded individually (Mickleburgh et al., 2008ab; IUCN, 2009; Monadjem et al., 2017w). Trend:- 2016: Decreasing (Monadjem et al., 2017w). 2008: Decreasing (Mickleburgh et al., 2008ab; IUCN, 2009). REPRODUCTION AND ONTOGENY: Although data were inconclusive, Happold and Happold (1990b: 566) indicate that young were born early in the wet season in Malawi. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Malawi, Mozambique, South Africa, Tanzania, Zambia, Zimbabwe. Native: Malawi (Happold et al., 1988; Ansell and Dowsett, 1988: 30; Monadjem et al., 2010d: 550); Mozambique (Simmons, 2005: 394; Monadjem et al., 2010d: 550); South Africa (Simmons, 2005: 394; Monadjem et al., 2010d: 550); Zambia (Ansell, 1967; Ansell, 1986; Simmons, 2005: 394; Monadjem et al., 2010d: 550); Zimbabwe (Cotterill, 2004a: 261; Monadjem et al., 2010d: 550). Presence uncertain: Tanzania (Simmons, 2005: 394). DETAILED MORPHOLOGY: Baculum - The baculum ranges from 2.49 to 2.73 mm in length, the shaft is straight and parallelsided, the base is expanded, in lateral profile it is angled ventrally, while the tip is simple and unexpanded, unlike N. macrotis and N. parisii (Thomas et al., 1994:21). Figure 157. Distribution of Nycteris woodi INFRAORDER VESPERTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, 2007 *2007. VESPERTILIONIFORMACEI Van Cakenberghe, Kearney and Seamark, African Chiroptera Report. Publication date: July 2007. - Comments: The name of the Infraorder African Chiroptera Report 2020 ? 463 is based on Vespertilio Linnaeus, 1758 (type of VESPERTILIONIDAE Gray, 1821). (Current Combination) Yangochiroptera TAXONOMY: Currently two superfamily is known from within the Infraorder VESPERTILIONIFORMACEI for Africa: MOLOSSOIDEA Gervais, 1856; VESPERTILIONOIDEA Gray, 1821. COMMON NAMES: Czech: netopýrotvaří, netopýři. Superfamily MOLOSSOIDEA Gervais, 1856 *1856. Molossoidae Gervais. (Current Combination) 1998. Molossoidea: Simmons, Interfamilian relationships in bats, ???. - Comments: Originally included the families Molossidae Gervais, 1855 and Antrozoidae Simmons, 1998 [= Antrozoinae Miller, 1897]. Considered a synonym of Molossidae by Jackson and Groves (2015: 257). Family MOLOSSIDAE Gervais, 1856 *1856. Molossidae Gervais, in: F. Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool, (Sec. 7), Vol. 1, pt. 2 (Mammifères): 53 footnote. Publication date: 23 July 1856. - Comments: Type genus: Molossus E. Geoffroy St. Hillaire, 1805. - Etymology: The name of the family is derived from the Greek "molossus", a kind of dog used by Greek shepherds, in ancient times (see Taylor, 2005). (Current Combination) 1865. Molossi Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, p. 258. - Comments: Type genus: Molossus E. Geoffroy, 1805. Originally included the genera Dysopes Illiger, 1811 [= Molossus É. Geoffroy, 1805]; Molossus É. Geoffroy, 1805; Promops Gervais, 1855; Mormopterus Peters, 1865; Nyctinomus É. Geoffroy, 1818 [= Tadarida Rafinesque, 1814]; and Chiromeles [= Cheiromeles] Horsfield, 1824 (see Jackson and Groves, 2015: 257). 1866. Molossina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 93. Publication date: 1 February 1866. - Comments: Type genus: Molossus E. Geoffroy, 1805. Proposed as tribe and originally included the genera Molossus É. Geoffroy, 1805; Promops Gervais, 1855 and Nyctinomus É. Geoffroy, 1818 [= Tadarida Rafinesque, 1814] (see Jackson and Groves, 2015: 257). 1872. Molossidae Gill, Smiths. Misc. Coll., 11 (1): 17. - Comments: Type genus: Molossus E. Geoffroy, 1805. Originally included the genus Molossus É. Geoffroy, 1805 (see Jackson and Groves, 2015: 257). (Current Combination) 1889. Gymnuridae Ameghino, Actas Acad. Nac. Ciencias, Argentine e Cordoba, p. 351. Comments: See Taylor (1934). 1893. Molossini Winge, Samling af Afhandlinger. E Museo Lundii, 2 (1): 24. - Comments: Type genus: Molossus E. Geoffroy, 1805. Proposed as tribe(?) and originally included the genera Mystacina J. Gray, 1843; Nyctinomus É. Geoffroy, 1818 [= Tadarida Rafinesque, 1814]; Chiromeles [= Cheiromeles] Horsfield, 1824; and Molossus É. Geoffroy, 1805 (see Jackson and Groves, 2015: 257). TAXONOMY: Koopman (1993a: 232) mentioned Gervais, 1856, whereas Corbet and Hill (1992: 156) and Peterson et al. (1995: 142) mentioned Gill 1872: 17 as author for this family. Revised by Freeman (1981b). For a very different arrangement see Legendre (1984). See also Lamb et al. (2011). Pavlinov et al. (1995: 90) recognise two subfamilies: Cheiromelinae and Tadaridinae. 464 ISSN 1990-6471 Most recent authors (Hoofer and Van Den Bussche, 2003: 2; Simmons, 2005; Lamb et al., 2011: 1) accept that the Molossidae consists of two subfamilies: Molossinae and Tomopeatinae. The latter subfamily used to be assigned to the Vespertilionidae, but Hoofer and Van Den Bussche (2003: 2) clearly indicate that this subfamily belongs to the Molossidae. Representatives of this subfamily only occur in South America and are therefore not included here. Бульдоговые. Tanoboase (Ghana): afrifraa, frede frede. Vietnamese: Họ dơi thò đuôi. Based on 102 morphological characters from the skull, dentition, postcrania, external morphology, tongue, and penis, Gregorin and Cirranello (2015: 2) distinguish two clades within the Molossidae, both of which include taxa from the Old and the New World: One clade is comprised of Mormopterus, Platymops, Sauromys, Neoplatymops, Molossops, Cynomops, Cheiromeles, Molossus, and Promops, whereas the second includes Tadarida, Otomops, Nyctinomops, Eumops, Chaerephon, and Mops. The relation of Myopterus to these two clades remains unclear. They also found that genera such as Tadarida, Chaerephon and Molossops sensu lato do not appear to be monophyletic. Shi and Rabosky (2015: 1532) report that the approximate fossil date for the Molossidae is 37.2 million years ago. The family-level stem ages/crown ages are respectively 53.8 and 45.2 mya. Ammerman et al. (2012: 12) indicate that the split between the Old and New World Molossid bats took place about 29 Mya. Ammerman et al. (2012: 12) distinguish 16 - 17 genera, including 110 species worldwide. Currently (Simmons and Cirranello, 2020) recognized genera within the family MOLOSSIDAE: Austronomus Troughton, 1941); Cabreramops Ibãñez, 1980; Cheiromeles Horsefield, 1824; Cynomops Thomas,1920; Eumops Miller, 1906; Molossops Peters, 1866; Molossus E. Geoffroy, 1805; Mops Lesson, 1842; Mormopterus Peters, 1865; Myopterus E. Geoffroy, 1818; Neoplatymops Peterson, 1965; Nyctinomops Miller, 1902; Otomops Thomas, 1913; Platymops Thomas, 1906; Promops Gervais, 1856; Sauromys Roberts, 1917; Tadarida Rafinesque, 1814 and Tomopeas Miller, 1900. However, they completely overlooked the species belonging to the genus Chaerephon Dobson, 1874 in their overview. COMMON NAMES: Castilian (Spain): Murcielagos cola de raton o de cola libre. Czech: morousovití, netopýři buldočí, tadaridovití. Dutch: Buldogvleermuizen. English: Free-tailed Bats, Wrinkled-lipped Bats, Bulldog Bats, Mastiff Bats, Bonneted Bats, Wolfdog Bats. Finnish: Doggilepakot. French: Molossidés. German: Bulldoggfledermäuse, Bulldogg-Fledermäuse. Italian: Molòssidi. Norwegian: Buldoggflaggermus, foldeleppede flaggermus. Russian: Свободнохвостые, PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Lavocat (1961) found an indetermined Molossid species from the Mio-Pliocene at Beni Mellal. While Butler (1978: 66) found four indetermined species from the early Pleistocene at Olduvai, Avery (1991: 6) found the remains of two different sized molossids in the upper Pleistocene at Border Cave, South Africa. MOLECULAR BIOLOGY: Across the family, the 2n value varies between 34 and 52 (Sotero-Caio et al., 2017: 5). ROOST: López-Baucells et al. (2018: 30) investigated roost selection by molossid bats in rural Madagascar and found that they predominantly roosted in modern public buildings (e.g. schools, offices and libraries), built with cement and bricks. Small, traditional housed made from mud and wood were much less used by molossids, but housed small colonies of vespertilionids. PREDATORS: Mikula et al. (2016: Supplemental data) mention unidentified Molossidae to be taken by Grey kestrels (Falco ardosiaceus Vieillot, 1823) and Dickinson's kestrel (Falco dickinsoni Sclater, 1864), Red-necked falcon (Falco ruficollis Daudin, 1800), Eastern chanting goshawk (Melierax poliopterus (Cabanis, 1868)) [the latter three preying on "Tadarida" sp, where they also include Mops and Chaerephon]. PARASITES: HEMIPTERA Cimicidae: Crassicimex sexualis Ferris and Usinger, 1957 described from a single female taken 50 miles SE of Torit, Uganda (Haeselbarth et al., 1966: 12, host referenced as a Tadarida sp.). SIPHONAPTERA Ischnopsyllidae: Araeopsylla scitula (Rothschild, 1909) found in southern Africa Cape Province, Lesotho, Natal, Transvaal and Botswana, where it has been collected from Free-tailed bats which are its true host (Haeselbarth et al., 1966: 189). Lagaropsylla Jordan and Rothschild, 1921 almost African Chiroptera Report 2020 all records of hosts are from Free-tailed Bats (Haeselbarth et al., 1966: 190). Lagaropsylla hoogstraali Smit, 1957 from Sudan host recorded as a molossid (Haeselbarth et al., 1966: 190). Lagaropsylla idea Smit, 1957 recorded mainly from free-tailed bats, with distributions in tropical Africa (Haeselbarth et al., 1966: 190). Lagaropsylla oblique Smit 1957 known to infest molossid species from Sierra Leone, Cameroon and the Congo (Haeselbarth et al., 1966: 191). 465 Lagaropsylla tauffliebi Smit 1962 a series was taken from a free-tailed bat (Tadarida sp.) near Bangui, Central African Republic (Haeselbarth et al., 1966: 191; host referenced as a Tadarida sp.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Ethiopia, Guinea, Kenya, Madagascar, Mozambique, Namibia, South Africa, South Sudan, Tanzania, Uganda, Zambia, Zimbabwe. Subfamily Molossinae Gervais, 1856 *1856. Molossinae Gervais, in: F. Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool, (Sec. 7), Vol. 1, pt. 2 (Mammifères): 53 footnote. Publication date: 23 July 1856. - Comments: Type genus: Molossus E. Geoffroy St. Hilaire, 1805. (Current Combination) 1866. Molossina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 93. Publication date: 1 February 1866. - Comments: Type genus: Molossus E. Geoffroy, 1805. Proposed as tribe and originally included the genera Molossus É. Geoffroy, 1805; Promops Gervais, 1855 and Nyctinomus É. Geoffroy, 1818 [= Tadarida Rafinesque, 1814] (see Jackson and Groves, 2015: 257). 1984. Cheiromelinae Legendre, Rev. suisse Zool., 91 (2): 425. Publication date: June 1984. Comments: Type genus: Cheiromeles Horsfield, 1824. Pavlinov et al. (1995: 90) recognise this as a subfamily, but see R(3345: 432) and Jackson and Groves (2015: 257). 1984. Molossinae: Legendre, Rev. suisse Zool., 91 (2): 399, 425. Publication date: June 1984. - Comments: Type genus: Molossus E. Geoffroy, 1805. Originally included the genera Molossus É. Geoffroy, 1805; Eumops Miller, 1906; Molossops Peters, 1865 [including the subgenera Cynomops Thomas, 1920 and Neoplatymops Peterson 1965]; Myopterus É. Geoffroy, 1818 and Promops Gervais, 1855 (see Jackson and Groves, 2015: 258). 1984. Tadaridinae: Legendre, Arch. Zool. Mus., Moscow State Univ., 91 (2): 399, 426. Comments: Type genus: Tadarida Rafinesque, 1814. Originally included the genera Tadarida Rafinesque, 1814; Mormopterus Peters, 1865a; Nyctinomops Miller, 1902; Otomops Thomas, 1913; and Rhizomops Legendre, 1984 [= Tadarida Rafinesque, 1814 (see Jackson and Groves, 2015: 257). (Lapsus) 1995. Tadarinae: Peterson, Eger and Mitchell, Faune de Madagascar, 84: 143. (Lapsus) TAXONOMY: Bronner et al. (2003) state that the genera Mops and Chaerephon have often been included as subgenera of Tadarida (Ellerman et al., 1953; Hayman and Hill, 1971; Swanepoel et al., 1980; Meester et al., 1986; Corbet and Hill, 1992). Roberts (1951) elevated these taxa to generic rank, an approach followed by Freeman (1981b), Koopman (1982, 1993a) and Simmons (2005). In southern Africa, Mops is therefore represented by two species (M. midas and M. condylurus); Chaerephon by five (C. bivittata, C. ansorgei, C. nigeriae, C. chapini and C. pumilus); and Tadarida by four (T. aegyptiaca, T. lobata, T. ventralis and T. fulminans). Other African genera are Mormopterus, Myopterus, Otomops, Platymops and Sauromys (Lamb et al., 2011: 2), with Myopterus, Platymops and Sauromys being endemic to mainland Africa. Ammerman et al. (2012: 23) suggest that this subfamily should be subdivided into four tribes: Molossini containing the New World taxa (Molossus, Eumops, Molossops, Cynomops, Neoplatymops, Nyctinomops, and Promops), tribe Tadarini containing Old World taxa (Tadarida, Chaerephon, Mops, Platymops, Sauromys, Myopterus, and Otomops), tribe Cheiromelini, and tribe Mormopterini. Based on five nuclear sequence markers, Napier (2013) found strong support for the monophyly of the combined genera Chaerephon and Mops (with the exception of C. jobimena). There was no support for monophyly of either of these genera, suggesting that both should be combined in a single genus. Her study also suggests that a new species of Otomops should be names from northeast Africa and Arabia. COMMON NAMES: Czech: praví morousi. English: Free-tailed Bats, Wrinkled-lipped Bats, Bulldog Bats, Mastiff Bats, Bonneted Bats, Wolf-dog Bats. French: Molosses. 466 ISSN 1990-6471 Genus Mops Lesson, 1842 *1842. Mops Lesson, Nouv. Tabl. Regn. Anim. Mammifères, 18. Publication date: 1842. Comments: Type species: Mops indicus Lesson, 1842 (=Molossus mops de Blainville, 1840; Genotype Mops indicus Lesson = Dysopes mops F. Cuvier; listed in synonymy of Nyctinomus by Miller, 1907, Bull. U.S. Nat. Mus., 57, p. 251; but listed as a genus with distinct characters by Thomas, 1913, J. Bombay Nat. Hist. Soc., 22, p. 91). - Etymology: From the masculine German substantive Mops, meaning "pug", a small companion dog resembling a bulldog, corresponding to the French "carlin" and the Italian "carlino"; the name refers to the wrinkled lips characteristic of the genus Mops and the family Molossidae. According to one seemingly erroneous interpretation, from the Malaysian substantive mòps, meaning "bat" (see Lanza et al., 2015: 293). (Current Combination) 1907. Chaerephon: Andersen. (Name Combination) 1951. Tadarida (Chaerephon): Ellerman and Morrison-Scott. (Name Combination) TAXONOMY: Formerly included in Tadarida; for synonyms see Freeman (1981b: 158). See also Legendre (1984). Harrison and Bates (1991: 64), Ansell and Dowsett (1988: 45), Corbet and Hill (1992: 157), Grubb et al. (1998: 93) and Horácek et al. (2000: 137), who include Mops as subgenus in Tadarida. A key for the species traditionally assigned to Mops is provided by Dunlop (1999: 1, Mammalian Species, 615), whereas a key for the species traditionally assigned to Chaerephon is provided by Bouchard (1998: 1, Mammalian Species, 574). In their molecular phylogeny study, Lamb et al. (2010: 204; 2011: 9) found strong support for the monophyly of the Chaerephon + Mops taxa (about 17.2 Mya old), but not for either of the genera separated. They also found some indications that certain Mops species are ancestral and Chaerephon species more derived. Their results suggest that Chaerephon should not be considered a subgenus of Tadarida, nor a genus on itself. They suggest combining Chaerephon (except Chaerephon jobimena) together with Mops into a single genus pending a more complete study. Ammerman et al. (2012: 20) also found support for a Mops - Chaerephon clade, which resulted in paraphyly of Chaerephon, and confirm again that Chaerephon is distinct from Tadarida. A further morphological study by Gregorin and Cirranello (2015: 10) confirmed the paraphyly of Chaerephon, but was unable to resolve the relationships within the genus. Simmons and Cirranello (2020) included "Chaerephon" in the list of synonyms for Mops, but did not assign it to as subgenus, as they do for the species traditionally assigned to the genus Mops. The morphological study by Gregorin and Cirranello (2015: 10) was unable to resolve the relations within the genus Mops, but it supports the monophyly of the genus. Currently (Simmons and Cirranello, 2020) recognized subgenera and species of the genus Mops: Mops Lesson, 1842: condylurus (A. Smith, 1833); congicus J. A. Allen, 1917; demonstrator (Thomas, 1903); leucostigma G. M. Allen, 1918; midas (Sundevall, 1843); mops (de Blainville, 1840) – western Malaysia, Sumatra, Borneo perhaps Java (Simmons, 2005: 443); niangarae J. A. Allen, 1917; niveiventer Cabrera and Ruxton, 1926; sarasinorum (A. Meyer, 1899) – Sulawesi (Indonesia) and adjacent small islands, Philippines (Simmons, 2005: 443); trevori J. A. Allen, 1917. Xiphonycteris Dollman, 1911: brachypterus (Peters, 1852); nanulus J. A. Allen, 1917; petersoni (El Rayah, 1981); spurrelli (Dollman, 1911); thersites (Thomas, 1903). No subgenus: aloysiisabaudiae (Festa, 1907); ansorgei Thomas, 1913; atsinanana Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, 2010; bemmeleni (Jentink, 1879); bivittatus (Heuglin, 1861); bregullae (Felten, 1964) – Vanuatu, Fiji Isls (Simmons, 2005: 433); chapini J. A. Allen, 1917; gallagheri (Harrison,1975); jobensis (Miller, 1902) – Seram (Moluccas), Yapen Isl (Indonesia), New Guinea, northern and central Australia (Simmons, 2005: 434); jobimena Goodman and Cardiff, 2004; johorensis (Dobson, 1873) – western Malaysia, Sumatra (Indonesia) (Simmons, 2005: 433); leucogaster A. Grandidier, 1870; major (Trouessart, 1897); nigeriae Thomas, 1913; plicatus (Buchannan, 1800) – India and Sri Lanka to southern China, Hong Kong, Cambodiaand Vietnam, southeast through Malyasia to the Philippines, Borneo and Lesser Sunda Isls, Hainan (China), Cocos Keeling Isl (Indian Ocean) African Chiroptera Report 2020 (Simmons, 2005: 434); pumilus (Cretzschmar, 1830-1831); pusillus (Miller, 1902); russatus J. A. Allen, 1917; solomonis (Troughton, 1931) – Solomon Isls (Simmons, 2005: 435); tomensis (Juste and Ibañez, 1993). Additionally, there is the extinct African †rusingae Arroyo-Cabrales, Gregorin, Schlitter and Walker, 2002. Simmons and Cirranello (2020) included "Chaerephon" in the list of synonyms for Mops, but did not assign it to as subgenus, as they do for the species traditionally assigned to the genus Mops. For convenience reasons, however, we tentatively retain Chaerephon as a subgenus. 467 COMMON NAMES: Czech: velcí morousi. English: Mops Bats, Greater Free-tailed Bats, Mops Free-tailed Bats. French: Tadarides. Italian: Pipistrèllo carlìno. SEXUAL DIMORPHISM: El-Rayah (1980: 163) confirmed the sexual dimorphism in the premolar size reported earlier: in males the first lower premolar is larger at the base than the second lower premolar, whereas in females this is the opposite. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Burundi, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Ghana, Kenya, Madagascar, Malawi, Namibia, South Africa, South Sudan, Tanzania, The Gambia, Uganda, Zambia, Zimbabwe. †Mops kerio Gunnell and Manthi, 2018 *2018. Mops kerio Gunnell and Manthi, J. Hum. Evol., p. "6", fig. 6Bj. Publication date: 6 April 2018. Type locality: Kenya: Kanapoi Bat Site [02 18 00 N 36 04 00 E] [Goto Description]. Holotype: NMK KP-69821: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. The holotype consists of a M2. - Etymology: Referring to the Kerio river that flows through the Kanapoi area (Gunnell and Manthi, 2018: "6"). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pliocene (Brown et al., 2019: Suppl.) †Mops rusingae (Arroyo-Cabrales, Gregorin, Schlitter and Walker, 2002) *2002. Tadarida rusingae Arroyo-Cabrales, Gregorin, Schlitter and Walker, J. Vert. Paleont., 22 (2): 382. Type locality: Kenya: Rusinga Island, west; KNM Locality 114 [00 20 S 34 06 E] [Goto Description]. - Comments: Hiwegi formation; late Early Miocene, ca. 17.5 - 18.0 Ma. - Etymology: The species name is derived from the specific locality where the holotype was collected, Rusinga Island (see Arroyo-Cabrales et al., 2002). 2003. Tadarida (Mops) rusingae: Czaplewski, Morgan and Naeher, Acta Chiropt., 5 (1): 65. (Name Combination) 2008. Mops rusingae: Van Cakenberghe and Seamark, African Chiroptera Report. Publication date: July 2008. (Name Combination, Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: The holotype was found in the Hiwegi formation; late Early Miocene (Burdigalian - Brown et al., 2019: Suppl.) deposits, dating between 17.5 to 18.0 Mya (Arroyo-Cabrales et al., 2002). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". †Mops turkwellensis Gunnell and Manthi, 2018 *2018. Mops turkwellensis Gunnell and Manthi, J. Hum. Evol., p."5", "6", fig. 6C. Publication date: 6 April 2018. Type locality: Kenya: Kanapoi Bat Site [02 18 00 N 36 04 00 E] [Goto Description]. Holotype: NMK KP-69820: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. The holotype consists of a left M 2. - Etymology: Referring to the Turkwell river, which flows into Lake Turkana north of Kanapoi (Gunnell and Manthi, 2018: "6"). 468 ISSN 1990-6471 PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pliocene (Brown et al., 2019: Suppl.) Subgenus Mops (Chaerephon) Dobson, 1874 Chœrephon: Dobson, J. Asiatic Soc. Bengal, 431. - Comments: "Incorrect subsequent spelling", see Jackson and Groves (2015: 259), who indicated that "this spelling does not appear to have [been] subsequently used", although Bræstrup (1935b: 94) and Roberts (1932a: 17; 1935: 245) a.o. used Choerephon. Most authors (including Jackson and Groves, 2015: 259) assign this spelling to another publication by Dobson (Dobson, 1878), but this was actually the original spelling. *1874. Nyctinomus (Chaerephon) Dobson, J. Asiatic Soc. Bengal, 43 (2): 144. Publication date: 17 October 1874. - Comments: Type species: Molossus (Nyctinomus) johorensis Dobson, 1873. Not Chaerephon Godman in Godman and Salvin, 1900 (Insecta, Lepidoptera, Hesperiidae) (see Jackson and Groves, 2015: 258). - Etymology: From the Greek "chaerephon" meaning joyful-sounding. Flannery (1990: 370; 1995a: 404) refers to it as "a proper name, significance unknown". Dobson (1874) in Bouchard (1998: 4) and Lanza et al. (2015: 268) indicate that it refers to the friend of Socrates who is depicted as "Χαιρεφῶν" ("Chaerephon the bat") in Aristophanes' comedy, Aves, maybe alluding to his nocturnal habits or bony appearance or as suggested by Plato in Apology, to his sudden, excitable nature. 1917. Chærephon (Lophomops) J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 460, text-fig. 9 11. Publication date: 29 September 1917 [Goto Description]. - Comments: Type species: Chærephon (Lophomops) chapini J.A. Allen, 1917, by original designation (see Jackson and Groves, 2015: 259). 1938. Lephomops: Frechkop, Exploration du Parc National Albert, 48. - Comments: Reported as subgenus of Chaerephon by Frechkop (1938: 48), but clearly a lapsus, as furtheron (p. 53) the correct Lophomops is used. (Lapsus) 1962. Tadarida (Chaerophoron): Taufflieb, Bull. Inst. Rech. Sci. Congo, 1: 112. (Lapsus) 1965. Nyctinimus: Koopman, Am. Mus. Novit., 2219: 21. Publication date: 22 June1965. (Lapsus) 2004. Chaeraphon: Okafor, Igbinosa and Ezenwaji, Anim. Res. Int., 1 (1): 64. Publication date: April 2004. (Lapsus) 2017. Chaereophon: Weiss, Dabrowski, Kurth, Leendertz and Leendertz, Virology J., 14 (181): 1. Publication date: 18 September 2017. (Lapsus) ? Chaerephon sp.: ? Chaerophon: (Lapsus) 1874. TAXONOMY: Formerly included in Tadarida; see Freeman (1981b: 60, 133, 150); see also Legendre (1984). Harrison and Bates (1991: 62) and Ansell and Dowsett (1988: 45), Corbet and Hill (1992: 157), Grubb et al. (1998: 93) and Horácek et al. (2000: 137) include Chaerephon in Tadarida. A key is provided by Bouchard (1998: 1, Mammalian Species, 574). Nyctinomus is considered a valid genus by McFarland (1998: 34), but included in Tadarida by Koopman (1993a: 240), Grubb et al. (1998: 93), or included in Chaerephon by Simmons (2005), or in Mormopterus by Jacobs and Fenton (2002, Mammalian Species, 703: 1). Grubb et al. (1998: 93) also indicate that Mahoney and Walton (1988a: 148 - 149) considered that Nyctinomus was dated from 1813 and was therefore the prior name for the genus Tadarida Rafinesque, 1814. Their case is unconvincing, and Grubb et al. (1998) do not think it desirable to change the long established usage of Tadarida. However, since Chaerephon Dobson, 1874 is clearly more recent it might be that Nyctinomus should be used instead of Chaerephon. Simmons and Cirranello (2020) include "Chaerophon" Dobson, 1874 in the list of synonyms of "Mops" Lesson, 1842, but they do not mention any of the Chaerephon species in their listing. In their molecular phylogeny study, Lamb et al. (2010: 204; 2011: 9) found strong support for the monophyly of the Chaerephon + Mops taxa (about 17.2 Mya old), but not for either of the genera separated. They also found some indications that certain Mops species are ancestral and Chaerephon species more derived. Their results suggest that Chaerephon should not be considered a subgenus of Tadarida, nor a genus African Chiroptera Report 2020 on itself. They suggest to combine Chaerephon (except Chaerephon jobimena) together with Mops into a single genus pending a more complete study. Ammerman et al. (2012: 20) also found support for a Mops - Chaerephon clade, which resulted in paraphyly of Chaerephon, and confirm again that Chaerephon is distinct from Tadarida. A further morphological study by Gregorin and Cirranello (2015: 10) confirmed the paraphyly of Chaerephon, but was unable to resolve the relationships within the genus. Simmons and Cirranello (2020) include all the "Chaerephon" species in the genus Mops. Currently recognized species of the subgenus Chaerephon: aloysiisabaudiae (Festa, 1907); ansorgei Thomas, 1913; atsinanana Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, 2010; bemmeleni (Jentink, 1879); bivittatus (Heuglin, 1861); bregullae (Felten, 1964) – Vanuatu, Fiji Isls (Simmons, 2005: 433); chapini J. A. Allen, 1917; gallagheri (Harrison,1975); jobensis (Miller, 1902) – Seram (Moluccas), Yapen Isl (Indonesia), New Guinea, northern and central Australia (Simmons, 2005: 434); jobimena Goodman and Cardiff, 2004; johorensis (Dobson, 1873) – western Malaysia, Sumatra (Indonesia) (Simmons, 2005: 433); leucogaster A. Grandidier, 1870; major (Trouessart, 1897); nigeriae Thomas, 1913; plicatus (Buchannan, 1800) – India and Sri Lanka to southern China, Hong Kong, Cambodiaand Vietnam, southeast through Malyasia to the Philippines, Borneo and Lesser Sunda Isls, Hainan (China), Cocos Keeling Isl (Indian Ocean) (Simmons, 2005: 434); pumilus (Cretzschmar, 1830-1831); pusillus (Miller, 1902); russatus J. A. Allen, 1917; solomonis (Troughton, 1931) – Solomon Isls (Simmons, 2005: 435); tomensis (Juste and Ibañez, 1993). For convenience reasons, however, we tentatively retain Chaerephon as a subgenus. COMMON NAMES: Czech: malí morousi, příšerecové. English: Wrinkle-lipped Bats, Lesser Free-tailed Bats, Lesser Mastiff Bats. French: Tadarides. Italian: Cherefónti. CONSERVATION STATUS: Assessment History IUCN (2004) - assessed C. shortridgei as NT ver 3.1 (2001). 469 BIOGEOGRAPHY: Taylor et al. (2009) investigated patterns in mitochondrial DNA (cytochrome b and D-loop) variation in Chaerephon spp. specimens from southern Africa and Madagascar, including individuals from Madagascar with distinct phenotypic characters and identified as C. leucogaster and C. pumilus. Taylor et al. (2009) found southern African populations of C. pumilus to be paraphyletic, Malagasy C. leucogaster to be nested within C. pumilus sensu lato, and Malagasy C. pumilus forming a sister group to a complex of African C. pumilus and Malagasy C. leucogaster clades. Goodman et al. (2010a) including C. pusillus samples from the Comoros (all four islands) and Aladabra, found that on Mayotte and Madagascar individuals that can phenotypically be assigned to C. leucogaster and C. pumilus sensu lato, are known to occupy the same day roost sites. Morphological and molecular genetics data indicate they act as a good biological species. The individuals from the Comoros and Aldabra formed a monophyletic group, which shows morphological similarity. Comorian individuals, classically assigned to C. pumilus, are best considered distinct from African and Madagascan populations, and represent the same taxon occuring in the western Seychelles under the name C. pusillus. PARASITES: BACTERIA: Di Cataldo et al. (2020: 2) examined 11 Nigerian Chaerephon sp. bats and found 4 (36 %) to be infected by hemoplasma bacteria. NEMATODA: Durette-Desset and Chabaud (1975: 317) described Molinostrongylus bauchoti from a Madagascan bat belonging to this genus. ACARI Trombiculidae: Stekolnikov (2018a: 154) mentioned Microtrombicula intranasalis Vercammen-Grandjean, 1965 from Chaerephon sp. VIRUSES: Of the 13 Chaerephon sp. specimens from Kisumu, Kenya, one tested positive for Adenovirus, one for Rhabdovirus and five for Polyoma virus (Conrardy et al., 2014: 259). Of the 16 specimens from Moi University, one tested positive for Adenovirus, one for Paramyxovirus and two for Polyomavirus. Adenoviridae Conrardy et al. (2014) detected an adenovirus in two individuals out of 35 sampled in Kenya. This virus groups within the Mastadenovirus genus. 470 ISSN 1990-6471 Coronaviridae - Coronaviruses SARS-CoV - During February 2008 in Ghana, six fecal samples were tested negative for coronavirus (CoV) RNA by Pfefferle et al. (2009) [as Chaerephon spp.]. Tong et al. (2009) tested several bats collected in Kenya in 2006 positive for the presence of coronavirus RNA in fecal samples (genus Betacoronavirus, subgenus Sarbecovirus; see Markotter et al., 2020: 5). Hu et al. (2015: 5) indicate that in Ghana, Kenya and Nigeria novel betacoronaviruses related to SARS-CoV have been detected. 12 of the 113 Kenyan specimens tested by Tao et al. (2017: Suppl.) were positive for CoV (10.6 %). Flaviviridae Pegivirus (BPgV) - 1 out of 3 Kenyan Chaerephon sp specimens was infected by clade K type Pegivirus (Quan et al., 2013: Table S5). Conrardy et al. (2014) detected a paramyxovirus in one out of 35 individuals sampled in Kenya. This virus groups with Morbillivirus-related viruses. Rhabdoviridae Conrardy et al. (2014) detected a rhabdovirus in one out of 35 individuals sampled in Kenya. This virus groups with the Tupavirus genus, with a near identical sequence detected in Miniopterus africanus. Polyomaviruses Orthopolyomavirus (BPyV) – DNA of two novel polyomaviruses were detected from fecal/oral swabs from Chaerephon sp specimens from Kenya (Tao et al., 2012) DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Cameroon, Congo (Democratic Republic of the), Kenya, Sierra Leone, South Sudan, Tanzania, Uganda. Paramyxoviridae Mops (Chaerephon) aloysiisabaudiae (Festa, 1907) *1907. Nyctinomus aloysii-sabaudiae Festa, Boll. Mus. Zool. Anat. Comp. Univ. Torino, 22 (546): 1. Publication date: 21 January 1907. Type locality: Uganda: Western province: Toro distrirct: East of Ruwenzori: "Toro distrirct" [ca. 00 30 N 30 00 E] [Goto Description]. Comments: Coordinates mentioned as 01 00 N 30 22 E by Thorn et al. (2009: 71). Allen (1939a: 107) mentioned the type locality as "Ruwenzoni". Grubb et al. (1998: 84) mention "East of Ruwenzori, Toro district". 1917. Nyctinomus aloysii-sabaudiæ: J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 460. (Name Combination) 1922. Chaerephon angolensis sabaudiae: de Beaux, Ann. Mus. civ. Stor. nat. Genova, ser. 3, (O.S. 49) 9: 371. Publication date: 28 August 1922. (Lapsus) 1966. Tadarida (Chaerephon) cyclotis Brosset, Biol. gabon., 2 (1): 80. Type locality: Gabon: Bélinga [01 13 N 13 10 E] [Goto Description]. 2020. Mops aloysiisabaudiae: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon aloysiisabaudiae: (Name Combination) ? Tadarida aloysiisabaudiae: (Name Combination) TAXONOMY: See Simmons (2005). threatened category (Mickleburgh et al., 2008bn; IUCN, 2009; Monadjem et al., 2017ax). COMMON NAMES: Czech: morous vévodský. English: Duke of Abruzzi's Wrinkle-lipped Bat, Abruzzi's Wrinklelipped Bat, Duke of Abruzzi's Free-tailed Bat. French: Tadaride du Duc des Abruzzes, Molosse du Duc des Abruzzes. German: Fürst von Abruzzen Bulldoggfledermaus. Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon aloysiisabaudiae] (Monadjem et al., 2017ax). 2008: LC ver 3.1 (2001) assessed as Tadarida aloysiisabaudiae (Mickleburgh et al., 2008bn; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bi; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more Regional None known. African Chiroptera Report 2020 MAJOR THREATS: This species is threatened by deforestation of its habitat by logging activities (Mickleburgh et al., 2008bn; IUCN, 2009; Monadjem et al., 2017ax). CONSERVATION ACTIONS: Mickleburgh et al. (2008bn) [in IUCN (2009)] and Monadjem et al. (2017ax) report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further studies are needed into the distribution, natural history and threats to this species. GENERAL DISTRIBUTION: Mops aloysiisabaudiae has been patchily recorded in West and Central Africa. It ranges from Côte d'Ivoire and Ghana in the west, through Cameroon and northern Central Africa, to southern Sudan and western Uganda. It is known from around 50 localities (mostly single specimens). 471 groups (Mickleburgh et al., 2008bn; IUCN, 2009; Monadjem et al., 2017ax). Trend:- 2016: Decreasing (Monadjem et al., 2017ax). 2008: Decreasing (Mickleburgh et al., 2008bn; IUCN, 2009). VIRUSES: Reoviridae Weiss et al. (2017: 1) isolated a new virus belonging to the subfamily Spinareovirinae: Taï Forest reovirus (TFRV) (tentatively assigned to the genus Coltivirus) from a bat in Côte d'Ivoire. This was the first bat-borne reovirus in Africa (Weiß, 2013: 65). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Uganda. Native: Cameroon; Central African Republic; Congo (The Democratic Republic of the); Côte d'Ivoire (Beaucournu and Fahr, 2003: 175); Gabon; Ghana; Nigeria (Tanshi et al., 2019; 14788); Sudan; Uganda. Presence uncertain: Ethiopia (Simmons, 2005). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) reported 2n = 48, FN = 66, BA = 20, a submetacentric X chromosome and an acrocentric Y chromosome. Protein / allozyme - Unknown. POPULATION: Structure and Density:- This high-flying species is rarely captured. It is found in relatively small Figure 158. Distribution of Mops (Chaerephon) aloysiisabaudiae Mops (Chaerephon) ansorgei (Thomas, 1913) *1913. Nyctinomus ansorgei Thomas, Ann. Mag. nat. Hist., ser. 8, 11 (63): 318. Publication date: 1 March 1913. Type locality: Angola: Malange [=Malanje] [09 32 S 16 20 E, 1 150 m] [Goto Description]. Holotype: BMNH 1910.4.8.4: ad ♂. Collected by: Dr. William John Ansorge; collection date: 17 February 1909; original number: 5. - Etymology: In honour of Dr. William John Ansorge (1850 - 1913), a collector of zoological specimens, who collected the type specimen of this species in Malange, northern Angola (see Taylor, 2005; Lanza et al., 2015: 271). 1946. Nyctinomus rhodesiae Roberts, Ann. Transv. Mus., 20 (4): 307. Type locality: Zimbabwe: Bindura district: Masembura Native Reserve: Chikupo caves [17 24 S 31 20 E] [Goto Description]. Holotype: TM 9977: ad ♂, skin and skull. Collected by: H.B. Masterson; collection date: 10 July 1945. 1967. Tadarida bivittata: Ansell, Arnoldia Rhod., 2 (38): 25. - Comments: Not of Heuglin, 1861. 2020. Mops ansorgei: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon ansorgei: (Name Combination) ? Tadarida (Chaerephon) ansorgei: (Name Combination) ? Tadarida ansorgei: (Name Combination) 472 ISSN 1990-6471 TAXONOMY: Distinct from bivittata; see Eger and Peterson (1979: 1889), who revised it. COMMON NAMES: Afrikaans: Ansorge se losstertvlermuis, Ansorgelosstertvlermuis. Chinese: 安氏犬吻蝠. Czech: morous Ansorgeův. English: Ansorge's Wrinklelipped Bat, Ansorge's Free-tailed Bat. French: Tadaride d'Ansorge, Molosse d'Ansorge, Molosse d'Afrique du Sud. German: Ansorges Bulldoggfledermaus. Italian: Cherefónte di Ansòrge. Portuguese: Morcego de cauda livre de Ansorge. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Hildebrand et al. (2010: 281) recorded this species from the Koka site in southern Ethiopia. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it is likely to occur in a number of protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ac; IUCN, 2009; Monadjem et al., 2017v). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon ansorgei (Monadjem et al., 2017v). 2008: LC ver 3.1 (2001) [assessed as Tadarida ansorgei] (Mickleburgh et al., 2008ac; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004al; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016aa). 2008: LC ver 3.1 (2001) (Friedmann and Daly, 2004). 1986 Indeterminate (Smithers, 1986). MAJOR THREATS: There appear to be no major threats to this species. In parts of West Africa, some populations may be threatened by overharvesting for subsistence food (Mickleburgh et al., 2008ac; IUCN, 2009; Monadjem et al., 2017v). CONSERVATION ACTIONS: Mickleburgh et al. (2008ac) [in IUCN (2009)] and Monadjem et al. (2017v) report that in view of the species wide range it is presumably present in many protected areas (including Comoe National Park, Côte d'Ivoire). No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Mops ansorgei is widely distributed over much of sub-Saharan Africa. It ranges from northeastern Côte d'Ivoire in the west, through much of the northern part of West and Central Africa (north of the Congo basin), to Ethiopia and East Africa, from here southwards as far as northeastern South Africa. It has been recorded between 400 and 2,000 m asl. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 673,792 km2. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 541; Taylor et al., 2018b: 62); Cameroon (Eisentraut, 1968: 171 - 1st record from West Africa); Congo (The Democratic Republic of the) (Hayman et al., 1966; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 541); Côte d'Ivoire; Ethiopia; Mozambique (Monadjem et al., 2010d: 541: Monadjem et al., 2010c: 382); South Africa (Monadjem et al., 2010d: 541); Sudan; Uganda (Kityo and Kerbis, 1996: 63); Zambia (Ansell, 1978: Monadjem et al., 2010d: 541); Zimbawe (Cotterill, 2004a: 261: Monadjem et al., 2010d: 541). Presence uncertain: Namibia. ECHOLOCATION: Jacobs (1996) and Fenton and Bell (1981) reported a highest frequency of 28 kHz for individuals at Sengwa in Zimbabwe. Taylor (2000: 67) reported call details as: long duration (15 ms), multi-harmonic (up to two), narrow bandwidth (16 - 28 kHz) shallow-FM calls, with low dominant frequency of 18 kHz, in a field guide to bats of Southern Africa. For "Chaerephon ansorgei" from Mozambique, Taylor et al. (2013b: 18) reported the following parameters for 3 calls: Fmax: 23.1 ± 3.5 (20.1 - 27.0) kHz, Fmin: 186. ± 1.5 (16.9 - 19.8) kHz, Fknee: 21.5 ± 1.4 (19.9 - 22.6) kHz, Fchar: 19.8 ± 1.1 (18.5 20.5) kHz, duration: 10.6 ± 1.2 (9.4 - 11.9) msec. And for 12 calls from Waterberg, South Africa: Fmax: 22.1 ± 1.53 (19.5 - 243) kHz, Fmin: 18.5 ± 0.92 (17.5 - 20.0) kHz, Fknee: 20.1 ± 1.05 (18.9 - 21.8) kHz, Fchar: 19.0 ± 0.94 (17.8 - 20.5) kHz, duration: 6.4 ± 1.51 (4.5 - 9.3) msec. At Farm Welgevonden (RSA), Taylor et al. (2013a: 556) report a Fknee value of 20 (19 - 22) kHz. In the Soutpansberg area (RSA), Linden et al. (2014: 40) mentioned the following characteristics for C. cf. ansorgei: Fmin: 17 - 20 kHz, Fchar: 18 - 21 African Chiroptera Report 2020 kHz, Fknee: 19 - 23, Slope: 24 - 212 OPS, duration: 4 - 12 msec. 20 calls from the Mapungubwe National Park (RSA) were recorded by Parker and Bernard (2018: 57): Fchar: 18.21 ± 1.01 kHz, Fmax: 20.92 ± 3.39 kHz, Fmin: 17.59 ± 1.03 kHz, Fknee: 19.38 ± 1.49 kHz, duration: 10.07 ± 3.03 msec with 2.47 ± 1.43 calls/sec. Weier et al. (2020: Suppl.) reported on 12 calls from the Okavango River Basin with the following characteristics: Fmax: 19.68 ± 0.53 kHz, Fmin: 18.35 ± 3.33 kHz, Fknee: 19.49 ± 0.45 kHz, Fchar: 19.19 ± 0.26 kHz, slope: 6.26 ± 9.90 Sc, duration: 5.99 ± 3.52 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) and Rautenbach et al. (1993) agreed on 2n = 48, and the Y chromosome being acrocentric. However, Smith et al. (1986) reported FN = 66, BA = 20, and a subtelocentric X chromosome, whereas Rautenbach et al. (1993) reported for specimens from South African, FN = 68, BA = 22, and a submetacentric X chromosome. Protein / allozyme - Unknown. 473 Trend:- 2016: Stable (Monadjem et al., 2017v). 2008: Stable (Mickleburgh et al., 2008ac; IUCN, 2009). REPRODUCTION AND ONTOGENY: Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] mentioned lactating females in NovemberJanuary and April. PARASITES: SIPHONAPTERA Ischnopsyllidae: Lagaropsylla anciauxi Smit, 1957 from Katanga, Congo (Haeselbarth et al., 1966: 190). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Cameroon, Congo (Democratic Republic of the), Ethiopia, Kenya, Malawi, Mozambique, Nigeria, Rwanda, South Africa, South Sudan, Uganda, Zimbabwe. PREDATORS: Mikula et al. (2016: Supplemental data) mention the Southern yellow-billed hornbill (Tockus leucomelas Lichtenstein, 1842) as diurnal avian predator. POPULATION: Structure and Density:- It appears to be generally uncommon, although colonies may contain hundreds of bats (Mickleburgh et al., 2008ac; IUCN, 2009; Monadjem et al., 2017v). Figure 159. Distribution of Mops (Chaerephon) ansorgei Mops (Chaerephon) atsinanana (Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, 2010) *2010. Chaerephon atsinanana Goodman, Buccas, Naidoo, Ratrimomanarivo, Taylor and Lamb, Zootaxa, 2551: 21, figs 34, 6A, 7. Publication date: 28 July 2010. Type locality: Madagascar: Fianarantsoa: Farafangana: Collèges d'Enseignement Général (CEG) Fenoarivo [22 49.275 S 47 49.860 E, 10 m] [Goto Description]. Holotype: FMNH 185259: ad ♀, skull and alcoholic. Collected by: Fania H. Ratrimomanarivo; collection date: 26 April 2005. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 185260: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185261: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185262: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185263: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185264: Collected by: ?: 474 ISSN 1990-6471 2020. ? Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185265: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185266: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185267: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). Paratype: FMNH 185268: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Same locality as the type specimen (see Goodman et al., 2010a: 21). - Etymology: The name atsinanana is derived from the Malagasy word meaning "from the east" (see Goodman et al., 2010a: 23). Mops atsinanana: Simmons and Cirranello, BatNames.org. (Current Combination) Chaerephon pumilus: CONSERVATION STATUS: Global Justification This species is known from numerous sites in the eastern portion of the island, particularly in synanthropic settings, where it can be very common (Goodman et al., 2010a; Goodman and Ramasindrazana, 2013). Assessment History Global 2016: LC ver. 3.1 (2001) [assessed Chaerephon atsinanana] (Goodman, 2017e). as Regional None known. MAJOR THREATS: Goodman (2017e) report that none are known. CONSERVATION ACTIONS: On Madagascar, it has not been recorded from any protected areas, but it has been recorded from houses close to protected areas as "Chaerephon pumilus" (Mickleburgh et al. (2008dr) [in IUCN (2009)]). While Goodman (2017e) make no suggestions for conservation action. GENERAL DISTRIBUTION: Mops atsinanana is known from numerous localities across the eastern half of Madagascar from near sea level to over 1,100 m (see Goodman et al., 2010a: 26). In Madagascar, the species is restricted to the humid areas of the island (in the north and east) and can be found up to 1,300 m asl. Its southernmost range in Madagascar is not well understood and it has not been recorded south of Moramanga (Goodman and Cardiff, 2004). Native: Madagascar (Goodman et al., 2010a: 26). BIOGEOGRAPHY: Mops atsinanana is common in the mesic eastern portion of Madagascar, across an elevational range from near sea level to 1,100 m a.s.l. (Lamb et al., 2012: 19). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 60) found that the triangular shaped baculum of this species could be subdivided into three morphotypes: morphotype A is more rounded than the flattened morphotype B, and morphotype C is similar to B, but with an indentation in the proximal base. The shaft curvature is more pronounced in types B and C as compared to type A. In all morphotypes, the distal portions are similar, gradually tapering from the midshaft towards a blunt tip. Length: 0.90 ± 0.066 (0.78 - 1.00) mm, width: 0.34 ± 0.056 (0.26 - 0.46) mm. MOLECULAR BIOLOGY: DNA - See Goodman et al. (2010a). Karyotype - Unknown. Protein / allozyme - Unknown. Napier (2013) hypothesized that this species should show relatively low levels of intraspecific genetic structure as bats are strong fliers and are able to traverse the riverine and mountain barriers. However, she needed to reject this hypothesis as genetic distances of up to 8.14 % were found. She furthermore suggests that these differences are consistent with the existence of female philopatry. HABITAT: Goodman et al. (2010a: 26) report that all of the sites where this species is found, are in urban or at least rural areas and outside of natural forest. They were also unable to find a natural day-roost of this species. A further analysis by Lamb et al. (2012: 29) indicated that the predicted distribution pattern was rather patchy and limited to the Madagascar Lowland Forests ecoregion and furthermore strongly associated with human population centres and transport routes. African Chiroptera Report 2020 ROOST: Wilkinson et al. (2012: 160) indicate that synanthropic structures are the typical roosts for this species on Madagascar. 475 Madagascar (Goodman, 2017e). On the basis of inference from genetic data, this species has witnessed a recent population expansion (Lamb et al., 2012). Trend:- 2016: Increasing (Goodman, 2017e). DIET: The faeces of five specimens examined by Rasoanoro et al. (2015: 64) contained the following volume percentages of prey: Coleoptera: 44.0 ± 5.08, Diptera: 8.3 ± 4.15, Hemiptera: 3.1 ± 1.36, Homoptera: 19.0 ± 4.42, Hymenoptera: 0.3 ± 0.31, and Lepidoptera: 25.4 ± 1.86. Kemp et al. (2018: Suppl.) used DNA metacarcoding to detect insect pest species in the diet of these bats and found the following prey orders (in descending order): Coleoptera, Ephemeroptera, Lepidoptera (Meyrickiella homosema (Meyrick, 1887), Leguminivora sp., Mythimna sp., Cydia choleropa (Meyrick, 1913)), Diptera (Simulium lineatum (Meigen, 1804), Culex MBI-27, Anopheles squamosus Theobald, 1901, Culex annulioris Theobald, 1901), Orthoptera, Trichoptera, Blattodea, Hemiptera, Sarcoptiformes, Trombidiformes, Odonata, Siphonaptera, Hymenoptera. POPULATION: Structure and Density:- There are no estimates of its population in Madagascar, but it is thought to be locally common in the eastern zone (Mickleburgh et al., 2008dr; IUCN, 2009, as C. pumilus). Not estimated, but can be common as a synanthropic species in villages and towns of eastern VIRUSES: Wilkinson et al. (2012: 160) tested 20 individuals from the Madagascar using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Figure 160. Distribution of Mops (Chaerephon) atsinanana Mops (Chaerephon) bemmeleni (Jentink, 1879) *1879. Nyctinomus Bemmeleni Jentink, Notes Leyden Mus., 1 (1): 125. Publication date: April 1879. Type locality: "Africa": "Liberia ??" [Goto Description]. Holotype: RMNH MAM.21649: ad ♀, skull and alcoholic. Collection date: 1875. - Comments: The type locality was originally mentioned as "Liberia", but see Kuhn (1965: 327), who doubts this, since the type specimen had come from a zoo. - Etymology: In honour of Mr. A.A. van Bemmelen, director of the Zoological Gardens at Rotterdam, who presented the type specimen to the museum (see Jentink, 1879b: 127). 1897. [Nyctinomus (Nyctinomus)] Vemmeleni: Trouessart, Catalogus Mammalium tam viventium quam fossilium, nov. ed. 1, pt. 1: 146. Publication date: 1897. (Lapsus) 1903. Nyctinomus cisturus Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (71): 502. Publication date: 1 November 1903. Type locality: Sudan: Equatoria province: 25 mi (37.5 km) N Gondokoro: Mangala (=Mongalla) [05 10 N 31 50 E, 500 m] [Goto Description]. Holotype: BMNH 1902.7.4.4: ad ♂. Collected by: W.L.S. Loat Esq. Original number: 2805. Presented/Donated by: W.L.S. Loat Esq. 2020. Mops bemmeleni Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon bemmeleni: (Name Combination) ? Tadarida bemmeleni cistura: (Name Combination) ? Tadarida bemmeleni: (Name Combination) 476 ISSN 1990-6471 TAXONOMY: Includes cisturus; see Koopman (1975: 425). The data used by Herkt et al. (2017: Appendix S9) support the idea that bemmeleni includes more than one species. However, genetic tests to confirm this are not yet available, but material from the small overlapping area can clearly be allocated to the ssp. cisturus, rendering a large geographic gap between occurrence records of either population: one in West and western Central Africa (bemmeleni) and one in East Africa (cisturus). COMMON NAMES: Chinese:腺尾犬吻蝠. Czech: morous liberijský. English: Gland-tailed Wrinkle-lipped Bat, Van Bemmelen's Wrinkle-lipped Bat, Gland-tailed Free-tailed Bat. French: Tadaride à glande caudale, Molosse à glandes caudales. German: Schwanzdrüsen-Bulldoggfledermaus, Bemmelens Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in a number of protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008am; IUCN, 2009; Monadjem et al., 2017ai). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon bemmeleni (Monadjem et al., 2017ai). 2008: LC ver 3.1 (2001) [assessed as "Tadarida bemmeleni"] (Mickleburgh et al., 2008am; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ba; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is threatened by habitat loss, largely through logging activities and the conversion of land to agricultural use (Mickleburgh et al., 2008am; IUCN, 2009; Monadjem et al., 2017ai). CONSERVATION ACTIONS: Mickleburgh et al. (2008am) [in IUCN (2009)] and Monadjem et al. (2017ai) report that it is present in Meru National Park, Kenya, and is presumably present in several more protected areas. Further studies are needed into the taxonomy, natural history, threats and distribution of this species. GENERAL DISTRIBUTION: Mops bemmeleni is widely distributed in Africa from Sierra Leone in the west, then patchily through West and Central Africa to southern Sudan, Uganda, Kenya and Tanzania in the east. It has been recorded up to 1,700 m asl on Mount Nimba along the border of Côte d'Ivoire and Guinea (J. Fahr pers. Comm., 2004 in Mickleburgh et al., 2008am in IUCN, 2009). Native: Cameroon; Congo (The Democratic Republic of the); Côte d'Ivoire; Guinea; Kenya (Start, 1969; as 'Tadarida cistura"); Liberia; Rwanda; Sierra Leone; Sudan; Tanzania; Uganda. Presence uncertain: Burkina Faso; Central African Republic; Chad; Ethiopia; Guinea-Bissau; Mali; Nigeria; Senegal. MOLECULAR BIOLOGY: DNA - unknown. Karyotype - Nagorsen et al. (1976) reported 2n = 48, Fna = 54, BA = 8, a submetacentric X chromosome and an acrocentric Y chromosome. POPULATION: Structure and Density:- The abundance of Mops bemmeleni is unclear, in parts of the range it seems to be a rare species that is found in small colonies consisting of a few individuals. In East Africa this species exhibits a clumped distribution, yet is relatively abundant locally (Webala et al., 2004). Trend:- 2016: Unknown (Monadjem et al., 2017ai). 2008: Unknown (Mickleburgh et al., 2008am; IUCN, 2009). PARASITES: Lainson and Naiff (2000: 128) and Duszynski (2002: 21) indicate that M. bemmeleni is the type host for Eimeria levinei Bray, 1958 (Apicomplexa). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Congo (Democratic Republic of the), Kenya, Liberia, Sierra Leone, South Sudan, Tanzania, Uganda. African Chiroptera Report 2020 477 Figure 161. Distribution of Mops (Chaerephon) bemmeleni Mops (Chaerephon) bivittatus (Heuglin, 1861) *1861. Nyctinomus bivittatus Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 13. Publication date: 1861. Type locality: Eritrea: Kérén [ca. 15 45 N 38 20 E] [Goto Description]. Holotype: RMNH ??:. See Jentink (1888b: 203). Syntype: ZMB 2792a: skull and alcoholic. Collected by: Theodor Von Heuglin. Formerly SMNS, from "Abyssinia, Bogos[land]". See Turni and Kock (2008: 72). Syntype: ZMB 2792b: skull and alcoholic. Collected by: Theodor Von Heuglin. Formerly SMNS, from "Abyssinia, Bogos[land]". See Turni and Kock (2008: 72). Lectotype: SMNS 981: ad ♂, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. Dieterlen et al. (2013: 294) indicated this specimen to be the lectotype, referring to ICZN article 74.6, which would indicate that this status was established by Dobson (1878), who called specimen this "'type". However, Dobson (1878: 426) presents the measurements of an adult male, which he called the "type", but since there are two adult males (981 and 981b), it is not clear which of these two specimens was referred to by Dobson (1878). Paralectotype: SMNS 981a: ad ♀, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. See Dieterlen et al. (2013: 294). Paralectotype: SMNS 981b: ad ♂, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. See Dieterlen et al. (2013: 294), but see also note above. - Etymology: From the masculine scientific Latin adjective bivittàtus, meaning "bivittate", as the species is provided with two narrow longitudinal stripes on and behind the head (see Lanza et al., 2015: 273). 1967. Tadarida bivittata: Ansell, Arnoldia Rhod., 2 (38): 29. (Name Combination) 1999. Chaerephon bivitattus: Taylor, Acta Chiropt., 1 (2): 199. (Lapsus) 2002. Chaerephon bivittata: Arroyo-Cabrales, Gregorin, Schlitter and Walker, 22 (2): 386. 2016. Chaerophon bivittattus: Simmons, Seiffert and Gunnell, Am. Mus. Novit., 3857: 43. Publication date: 9 May 2016. (Lapsus) 2020. Mops bivittatus: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon bivittatus: (Name Combination) ? Tadarida bivittata bivittata: (Name Combination) TAXONOMY: Revised by Eger and Peterson (1979). COMMON NAMES: Afrikaans: Gevlekte losstertvlermuis. Chinese: 斑 犬 吻 蝠 . Czech: morous skvrnitý. English: Spotted Wrinkle-lipped Bat, Spotted Free-tailed Bat, Spotted Gland-tailed Bat. French: Tadaride tachetée, Molosse tacheté. German: Gefleckte Bulldoggfledermaus, zweistreifige Doggengrämler. Italian: Cherefónte bivittàto. Portuguese: Morcego de cauda livre malhado. 478 ISSN 1990-6471 ETYMOLOGY OF COMMON NAME: So named because of the tiny white, and sometimes barely discernible, flecks on the upper parts of the body (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001) ) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ad; IUCN, 2009; Monadjem et al., 2017x). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon bivittatus] (Monadjem et al., 2017x). 2008: LC ver 3.1 (2001) [assessed as "Tadarida bivittata"] (Mickleburgh et al., 2008ad; IUCN, 2009). 2004: LC ver 3.1 (2001) [assessed as "Chaerephon bivittata"] (Mickleburgh et al., 2004an; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There appear to be no major threats to this species as a whole (Mickleburgh et al., 2008ad; IUCN, 2009; Monadjem et al., 2017x). CONSERVATION ACTIONS: Mickleburgh et al. (2008ad) [in IUCN (2009)] and Monadjem et al. (2017x) report that there appear to be no direct conservation measures in place. It is presumably present in some East African protected areas. Further studies are needed into the abundance, distribution and natural history of this bat. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Peterson and Nagorsen (1975) reported 2n = 48, FN = 54, BA = 8, a metacentric X chromosome, and an acrocentric Y chromosome for specimens from Kenya and Zimbabwe. Protein / allozyme - Unknown. POPULATION: Structure and Density:- Although there is little information on the abundance of this species, it is common in collections suggesting that it is easily found (Mickleburgh et al., 2008ad; IUCN, 2009; Monadjem et al., 2017x). Trend:- 2016: Stable (Monadjem et al., 2017x). 2008: Stable (Mickleburgh et al., 2008ad; IUCN, 2009). PARASITES: Beaucournu and Kock (1996) mentioned the first record for the flea Lagaropsylla anciauxi Smit, 1957 on this bat. Hastriter (2016: 15) described the flea Araeopsylla smiti from a M. bivittatus collected at Maji Moto, Kenya. Copeland et al. (2011: 363) reported the rediscovery of the "terrible hairy fly" (Mormotomyia hirsuta (after 62 years) in guano produced by "Chaerephon cf. bivittatus" and Tadarida aegyptiaca at Ukasi Hill, Kenya. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Eritrea, Ethiopia, Kenya, Malawi, Mozambique, South Sudan, Tanzania, Uganda, Zambia, Zimbabwe. GENERAL DISTRIBUTION: Mops bivittatus is an East African species ranging from Eritrea and Ethiopia in the north, through southern Sudan, Uganda, Kenya and eastern Tanzania, to Zambia, Zimbabwe and Mozambique in the south. Native: Eritrea; Ethiopia; Kenya; Malawi (Monadjem et al., 2010d: 541); Mozambique (Crawford-Cabral, 1989; Simmons, 2005); Sudan; Tanzania; Uganda; Zambia (Ansell, 1978; Monadjem et al., 2010d: 541); Zimbabwe (Ansell, 1986; Monadjem et al., 2010d: 541). Figure 162. Distribution of Mops (Chaerephon) bivittatus African Chiroptera Report 2020 479 Mops (Chaerephon) chapini (J.A. Allen, 1917) *1917. Chærephon (Lophomops) chapini J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 460, 461, text-fig. 9. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Faradje [03 44 N 29 43 E] [Goto Description]. Holotype: AMNH 48841: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 11 November 1912; original number: 1971. 1926. Chaerephon (Lophomops) shortridgei Thomas, Proc. zool. Soc. Lond., 1926, I: 289. Publication date: 29 April 1926. Type locality: Namibia: NW Ovamboland: 32 mi NW of Rehoboth Mission Station: Ukualukasi [17 32 S 14 37 E, 3 400 ft] [Goto Description]. Holotype: BMNH 1925.12.4.24: skin and skull. Collected by: Captain Guy Chester Shortridge; original number: 1389. - Comments: Considered a valid species by Peterson et al. (1995: 159), Goodman and Cardiff (2004: 236). - Etymology: I have named this striking species in honour of Capt. Shortridge, to whose energy and enthusiasm the great success of the South West African collecting expeditions has been almost wholly due, and who was greatly struck by its remarkable appearance (see Thomas, 1926). 1938. Chærephon lancasteri Hayman, Ann. Mag. nat. Hist., ser. 11, 1 (4): 383. Publication date: 1 April 1938. Type locality: Zambia: Loangwa Valley: Lundazi district: Lunzi River [ca. 11 S 33 E] [Goto Description]. Holotype: BMNH 1937.12.8.25: ad ♂, skin and skull. Collected by: D. Gordon Lancaster; collection date: August 1936; original number: 724. Notes: caught in hole in tree. - Comments: Kock (1969a: 140) situates the Lunzi river in Zambia, rather than in Zimbabwe. 2009. Choerephon (Lopomops) langi: ACR [Goto Description]. - Comments: nomen nudum. 2020. Mops chapini: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon chapini chapini: (Name Combination) ? Chaerephon chapini lancasteri: (Name Combination) ? Chaerephon chapini: (Name Combination) ? Chaerephon shortridgei: ? Tadarida chapini lancasteri: (Name Combination) ? Tadarida chapini: (Name Combination) TAXONOMY: Included in pumilus by Kock (1969a: 139). See Fenton and Eger (2002). COMMON NAMES: Afrikaans: Bleek losstertvlermuis. Chinese: 查平 犬 吻 蝠 . Czech: morous chocholatý. English: Long-crested Free-tailed Bat, Chapin's Wrinklelipped Bat, Long-crested Wrinkle-lipped Bat, Crested Wrinkle-lipped Bat, Chapin's Free-tailed Bat, Pale Free-tailed Bat, Long-crested Glandtailed Bat, Crested Free-tail Bat. French: Tadaride de Chapin, Molosse à crinière, Molosse pâle. German: Chapins Bulldoggfledermaus. ETYMOLOGY OF COMMON NAME: Named after Mr. James P. Chapin, co-organiser of the American Museum expedition to the former Belgian Congo that resulted in the discovery of the species (Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: No fossils are known (Fenton and Eger, 2002). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, it occurs in a number of protected areas, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ae; IUCN, 2009; Monadjem et al., 2017y). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon chapini] (Monadjem et al., 2017y). 2008: LC ver 3.1 (2001) [assessed as "Tadarida chapini"] (Mickleburgh et al., 2008ae; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004ag; IUCN, 2004). NT ver 3.1 (2001) [assessed as "Chaerephon shortridgei"] (Mickleburgh et al., 2004ak; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. 480 ISSN 1990-6471 MAJOR THREATS: There appear to be no major threats to this widespread species as a whole. It may be threatened in parts of its range through habitat loss and degradation (e.g. in northern Zimbabwe and parts of Zambia) (Mickleburgh et al., 2008ae; IUCN, 2009; Monadjem et al., 2017y). SEXUAL DIMORPHISM: Sexual dimorphism in the structure of interaural crest, with females having a small crest (3 - 5 mm) and males a long, bicolored one (12 - 15 mm) (Fenton and Eger, 2002). Krutzsch (2000: 128) pointed out that this crest is also odoriferous in the breeding male. CONSERVATION ACTIONS: Mickleburgh et al. (2008ae) [in IUCN (2009)] report that it has been recorded from Kruger National Park in South Africa, and from Mole National Park in Ghana and Comoe. Monadjem et al. (2017y) makes no recommendations for conservation actions. ECHOLOCATION: Jacobs (1996) and Fenton and Bell (1981) reported a highest frequency of 27 kHz for individuals at Sengwa in Zimbabwe. GENERAL DISTRIBUTION: Mops chapini has been widely recorded over much of sub-Saharan Africa. It has been collected from Côte d'Ivoire and Ghana in West Africa, southern Sudan, northeastern Democratic Republic of the Congo, Ethiopia, Uganda and Kenya, ranging south into southern Democratic Republic of the Congo and Congo, Angola, Zimbabwe, Zambia, Botswana and Namibia. Native: Angola (Monard, 1935; Crawford-Cabral, 1989; Hill and Carter, 1941; Monadjem et al., 2010d: 541); Botswana (Monadjem et al., 2010d: 541); Cameroon (Bol A Anong et al., 2011: 46; Bakwo Fils et al., 2014: 3); Congo (The Democratic Republic of the) (Hayman et al., 1966; Fenton and Eger, 2002: 2; Monadjem et al., 2010d: 541); Ethiopia; Ghana; Kenya (Fenton and Eger, 2002: 2); Namibia (Monadjem et al., 2010d: 541); Sudan (Fenton and Eger, 2002: 2); Uganda; Zambia (Ansell, 1969; Monadjem et al., 2010d: 541); Zimbabwe (Monadjem et al., 2010d: 541). Bates et al. (2013: 338) reject the occurrence of M. chapini in the Republic of Congo as the "Marine Park Nature Reserve" was erroneously located in that country in ACR (2012) rather than in the DRC. The record from Cameroon is probably doubtful (Jakob Fahr, pers. comm.). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Fenton and Eger (2002). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Fenton and Eger (2002). DETAILED MORPHOLOGY: Baculum - Unknown For a description of the crania, teeth, ears and tragus see Fenton and Eger (2002). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported 2n = 48, FN = 64, BA = 18, a subtelocentric X chromosome, and an acrocentric Y chromosome, for specimens from Namibia and Zimbabwe. Protein / allozyme - Unknown. HABITAT: See Fenton and Eger (2002). HABITS: See Fenton and Eger, 2002). Fenton and Griffin (1997: 248) recorded echolocation calls from "Tadarida chapini" at altitudes between 0 and 450 m above a Brachystegia woodland in Zimbabwe. ROOST: See Fenton and Eger (2002). PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- In general there is little information on the abundance of this species over its range (Mickleburgh et al., 2008ae; IUCN, 2009; Monadjem et al., 2017y). It can be common in northwestern Zimbabwe, but it is not regularly recorded. The species is found in small colonies, with a single observation of a small group of breeding females. Trend:- 2016: Unknown (Monadjem et al., 2017y). 2008: Unknown (Mickleburgh et al., 2008ae; IUCN, 2009). ACTIVITY AND BEHAVIOUR: See Fenton and Eger, 2002). REPRODUCTION AND ONTOGENY: See Fenton and Eger (2002). African Chiroptera Report 2020 481 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Botswana, Cameroon, Congo (Democratic Republic of the), Ethiopia, Ghana, Kenya, Namibia, Rwanda, South Sudan, Uganda, Zambia, Zimbabwe. Figure 163. Distribution (Chaerephon) chapini of Mops Mops (Chaerephon) gallagheri (Harrison, 1975) *1975. Tadarida (Chaerephon) gallagheri Harrison, Mammalia, 39 (2): 313. Type locality: Congo (Democratic Republic of the): Kivu province: 30 km SW Kindu: Scierie Forest [03 10 S 25 46 E] [Goto Description]. Holotype: BMNH 1976.207: ad ♂. Collected by: Major Michael D. Gallagher; collection date: 14 November 1974; original number: MDG-3256. Presented/Donated by: ?: Collector Unknown. Formerly HZM 1.7797. Holotype: HZM 1.7797: ♂. Collected by: Major Michael D. Gallagher; collection date: 14 November 1974; original number: MDB-3256. Presented/Donated by: ?: Collector Unknown. - Etymology: In honour of Major M.D. Gallagher, the collector of the type specimen. 2020. Mops gallagheri: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon gallagheri: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: morous kivuský. English: Gallagher's Wrinkle-lipped Bat, Gallagher's Free-tailed Bat, Zaïre Gland-tailed Bat. French: Tadaride de Gallagher, Molosse du Zaïre. German: Gallaghers Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of recent information on its extent of occurrence, ecological requirements, threats and conservation status (Mickleburgh et al., 2008ag; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) [assessed as "Tadarida gallagheri"] (Mickleburgh et al., 2008ag; IUCN, 2009). 2004: CR D ver 3.1 (2001) (Mickleburgh et al., 2004ai; IUCN, 2004). 2002: CR: B1+2c (Mickleburgh et al., 2002a: 22). 1996: CE (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are unclear. It may be threatened by habitat loss, through the conversion of forest to agricultural use and timber extraction, however, this needs to be confirmed (Mickleburgh et al., 2008ag; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008ag) [in IUCN (2009)] report that there are no direct conservation measures in place for this species. It is not known if the species is present within any protected areas. Additional studies are needed into the distribution, abundance, natural history and threats to this poorly known species. GENERAL DISTRIBUTION: Mops gallagheri has only been recorded from the type locality of 'Scierie Forest, 30 km of southwest Kindu, Kivu' in the Democratic Republic of the Congo. It may range more widely distributed, 482 ISSN 1990-6471 however, other surveys in the general area have not recorded this species. Native: Congo (The Democratic Republic of the) (Gallagher and Harrison, 1977; Harrison, 1975a). POPULATION: Structure and Density:- There is little information on the abundance of this species (Mickleburgh et al., 2008ag; IUCN, 2009). Trend:- 2008: Unknown (Mickleburgh et al., 2008ag; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the). Figure 164. Distribution of Mops (Chaerephon) gallagheri Mops (Chaerephon) jobimena (Goodman and Cardiff, 2004) *2004. Chaerephon jobimena Goodman and Cardiff, Acta Chiropt., 6 (2): 230, fig 2, 3, 4. Type locality: Madagascar: Antsiranana province: Ankarana special reserve, 2.6 km E Andrafiabe: Andrafiaba cave, in forest near [12 55.9 S 49 03.4 E, ca. 50 m] [Goto Description]. Holotype: FMNH 169716: ad ♂, skin and skull. Collected by: Steven M. Goodman; collection date: 23 January 2001; original number: SMG 11932. See Goodman and Cardiff (2004: 230). - Etymology: From the northern dialects of Malagasy, with "joby" meaning dark or black and "mena" red. The combination refers to the two color morphs of this species (see Goodman and Cardiff, 2004: 230). 2020. Mops jobimena: Simmons and Cirranello, BatNames.org. (Current Combination) TAXONOMY: Lamb et al. (2011: 9) indicate that, although morphologically similar to other "Chaerephon" taxa, jobimena is genetically more similar to Tadarida. COMMON NAMES: Czech: morous malgašský. English: Black and Red Free-tailed Bat. French: Tadaride rouge ou noir. German: Variable Bulldoggfledermaus. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) because although it appears to be rather rare in the few areas where it is known to occur, it is widespread and its population status may not be related to the amount or state of native forest. Additional information are needed on its propensity for roosting in large colonies in synanthropic settings as this will be crucial to a better understanding of its conservation status (Andriafidison et al., 2008l; IUCN, 2009; Monadjem et al., 2017bz). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon jobimena] (Monadjem et al., 2017bz). 2008: LC ver 3.1 (2001) [assessed as "Tadarida jobimena"] (Andriafidison et al., 2008l; IUCN, 2009). Regional None known. MAJOR THREATS: The threats to this species are not well known, but it is probably hunted in the south of Madagascar for food (Andriafidison et al., 2008l; IUCN, 2009; Monadjem et al., 2017bz). CONSERVATION ACTIONS: Andriafidison et al. (2008l) [in IUCN (2009)] and Monadjem et al. (2017bz) report that this species is known from in or near to four protected areas, but is never as common or as abundant as other sympatric molossid species (Goodman and Cardiff, 2004; Goodman et al., 2005a) and its conservation status should be reviewed in the future when new data are available. In particular, information on its propensity to dwell in buildings is needed. African Chiroptera Report 2020 GENERAL DISTRIBUTION: Mops jobimena is endemic to Madagascar where it is found from the north-west to the south-west of the island. It is known to occur in areas with dry deciduous or spiny forest from 50 to 870 m above sea level (Goodman and Cardiff, 2004; Goodman et al., 2005a). Although further surveys may reveal that this species is more widespread than currently known, it has yet to be recorded from some sites with extensive limestone caves, from where it would be expected based on habitat preference and distribution (Goodman and Cardiff, 2004; Goodman et al., 2005a; Cardiff, 2006). Native: Madagascar (Goodman and Cardiff, 2004; Goodman et al., 2005a; Cardiff, 2006). 483 2008l [in IUCN, 2009]; Monadjem et al., 2017bz). Forty individuals were recorded from a house roughly 10 km from Parc National de Namoroka (A. F. Kofoky pers. obs., in Andriafidison et al., 2008l [in IUCN, 2009]; Monadjem et al., 2017bz). Trend:- 2016: Unknown (Monadjem et al., 2017bz). 2008: Unknown (Andriafidison et al., 2008l; IUCN, 2009). PARASITES: Hastriter (2016: 9) described the flea Araeopsylla goodmani from a male M. jobimena collected at Isalo, 3.8 km NW Ranohira, along Namaza River, Madagascar. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 61) report the following shape for the baculum of this species: the proximal portion is notably expanded. Its size decreases slightly in the mid-shaft portion, then more notably distally. The distal tip tapers to a rounded point. Length: 1.16 mm, width: 0.37 mm. HABITAT: Dammhahn and Goodman (2013: 108) indicate that this species' foraging habitat consists of open areas and the area above the forest canopy. POPULATION: Structure and Density:- The population and local abundance of this species are not known, but at Ankarana the recorded roost is 1,200 individuals (S. G. Cardiff pers. obs., in Andriafidison et al., Figure 165. Distribution of Mops (Chaerephon) jobimena Mops (Chaerephon) leucogaster (A. Grandidier, 1869) *1869. Nyctinomus leucogaster A. Grandidier, Rev. Mag. Zool., ser. 2, 21: 337. Publication date: September 1869. Type locality: Madagascar: Mahab (?Mahabo) and Ménabé (E of Morondava) [20 23 S 44 40 E, 50 m] [Goto Description]. Neotype: FMNH 176137: ad ♂, skull and alcoholic. Collected by: Vola Razakarivony; collection date: 11 November 2002. - Comments: Ratrimomanarivo et al. (2009a: 28) mention "The only specimen possibly fitting the details of the Grandidier type is MCZ 45094, which is in poor condition and the original specimen label bears the following information: 'Cote Ouest' [= west coast], 'Nyctinomus pumelus [sic]', 'collection Grandidier', and '1901'. Based on numerous characters, including aspects of external, cranial, and dental measurements as well as pelage and soft part coloration, this animal is referable to C. leucogaster. However, the collection details are too ambiguous to clearly conclude that this specimen can be considered the type of C. leucogaster described by Grandidier (1869) and, further, the skull is considerably damaged." They also select a neotype (FMNH 176137): "collected in the Menabe Region of western coastal Madagascar, specifically at Belo sur Mer (20°44.139’S, 44°00.266’E) at 10 m above sea level on 11 November 2002 by Vola Razakarivony. This site is approximately 80 km southwest of Mahabo. The animal is an adult male, the body was preserved in formalin and subsequently transferred to 70% ethanol, the skull has been extracted, cleaned, and is in excellent condition.". - Etymology: From the Greek "leugos" and "gaster" meaning white-bellied (see Kozhurina, 2002: 16). 484 ISSN 1990-6471 1908. 1917. 1917. 1928. 1956. 2020. ? ? ? ? ? ? ? ? ? ? ? Chœrephon websteri Dollman, Ann. Mag. nat. Hist., ser. 8, 2 (12): 546. Publication date: 1 December 1908. Type locality: Nigeria: N Nigeria, on Bénoué river: Yola [09 12 N 12 29 E]. Holotype: BMNH 1908.10.6.8: ad ♀, skin and skull. Collected by: G.W. Webster; collection date: 27 July 1908; original number: 11. Chærephon (Lophomops) cristatus J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 463, textfig. 10 A, B. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Kinshasa province: near mouth of Congo River: Boma [05 50 S 13 03 E] [Goto Description]. Holotype: AMNH 48844: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 26 January 1915; original number: 2628. Chærephon frater J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 456. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Kinshasa province: near Boma: Malela [05 57 S 12 38 E, 30 m] [Goto Description]. Holotype: AMNH 49275: ad ♀, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 8 July 1915; original number: 2613. Topotype: MCZ 17219: ♀, alcoholic (skull not removed). Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 8 July 1915. Presented/Donated by: ?: Collector Unknown. Topotype: MCZ 17221: ♀, alcoholic (skull not removed). Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 8 July 1915. Presented/Donated by: ?: Collector Unknown. Chærophon (Lophomops) nigri Hatt, Bull. Soc. zool. France, 53: 374, text-fig. B. Publication date: 10 December 1928. Type locality: Mali: Timbuctu district: on Niger River: Bourem [16 59 N 00 20 W] [Goto Description]. Holotype: MNHN 736a: ad ♂, alcoholic (skull not removed). Collected by: Th. Monod; collection date: 6 January 1928. Paratype: AMNH 90390: ad ♂, skin and skull. Collected by: Th. Monod; collection date: 6 January 1928; original number: M.S.A.D. 736b. See AMNH website. Chaerephon nigri: Aellen, Mém. IFAN, 48A: 32. (Name Combination) Mops leucogaster: Simmons and Cirranello, BatNames.org. (Current Combination) Chaerephon frater: (Alternate Spelling) Chaerephon leucogaster: (Name Combination) Chaerephon pumila cristata: (Name Combination) Chaerephon pumila websteri: (Name Combination) Chaerephon websteri: (Alternate Spelling) Tadarida frater: (Name Combination) Tadarida leucogaster leucogaster: (Name Combination) Tadarida leucogaster websteri: (Name Combination) Tadarida leucogaster: (Name Combination) Tadarida pumila frater: Tadarida websteri: (Name Combination) TAXONOMY: Included in pumilus by Koopman (1993a: 233), Bouchard (1998: 1). Considered a valid species in the genus Tadarida (Chaerephon) by Peterson et al. (1995: 152) and Russ et al. (2001). Considered a valid species in the genus Chaerephon by Hutson et al. (2001: 33), Goodman and Cardiff (2004: 227), Lavrenchenko et al. (2004b: 141), and Simmons (2005). Russ et al. (2001) distinguish it from M. pumilus on smaller size and different echolocation calls. Ammerman et al. (2012: 23) indicate that the separation between leucogaster and pumilus took place less than 2 million years ago. COMMON NAMES: Czech: morous bělobřichý. English: Grandidier's Free-tailed Bat, Madagascan white-bellied Free- tailed Bat. French: Tadaride de Madagascar à ventre blanc, Tadaride à ventre blanc, Petite tadaride de Madagascar. German: Grandidiers Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Assessed by Mickleburgh et al. (2014b) as part of Chaerephon pumilus. Assessment History Global 2014: LC ver. 3.1 (2001) as a synonym of Chaerephon pumilus (Mickleburgh et al., 2014b). DD ver 2.3 (1994). Regional None known. African Chiroptera Report 2020 GENERAL DISTRIBUTION: Ethiopia to Ghana, Nigeria, Cameroon (Bol A Anong et al., 2011: 46; Bakwo Fils et al., 2014: 4; Waghiiwimbom et al., 2019b: 6 [as "Chaerephon nigri"]), Congo (Bates et al., 2013: 336), Congo (Democratic Republic of the Congo?), Mali, Madagascar (Russ et al., 2001), Nosy Be and Nosy Komba (Rakotonandrasana and Goodman, 2007: 6). O'Brien (2011: 288) reports it further from the Comoros, and the Tanzanian islands of Pemba, Unguja [=Zanzibar], and Mafia. BIOGEOGRAPHY: Mops leucogaster is found across an elevational range of 0 - 920 m a.s.l., and is confined almost exclusively to the more extensive and drier western portion of Madagascar (Lamb et al., 2012: 19). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 61) describe the baculum of M. leucogaster as "Eiffel tower shaped", with a bifurcated proximal tip that tapers abruptly to a distal point; length: 0.78 ± 0.072 (0.65 - 0.89) mm, width: 0.47 ± 0.055 (0.37 - 0.58) mm. HABITAT: Lamb et al. (2012: 29) indicate that the highly suitable habitat for this species was associated with low annual rainfall, high mean annual temperature, and high human influence. 485 PARASITES: BACTERIA Gomard et al. (2016: 5) found 1 out of 94 tested bats to be positive for Leptospira. SIPHONAPTERA Ischnopsyllidae: Lagaropsylla consularis Smit, 1957 from Ethiopia, Kenya, Uganda, Congo and Congo (Democratic Republic of), Angola, Zimbabwe and Mozambique (Haeselbarth et al., 1966: 190, host referred to as "Tadarida frater"). VIRUSES: Paramyxoviridae Specimens from this species tested by Wilkinson et al. (2014: 8270) were found to be positive for paramyxoviruses. Mélade et al. (2016b: 4) tested 94 Madagascan specimens, of which 6 were found positive for paramyxoviruses. Rhabdoviridae Mélade et al. (2016a: 6) tested 28 bats for Duvenhage lyssavirus and foud seroreactivlty in nine of them. UTILISATION: See Goodman et al. (2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Ghana, Madagascar, Mali, Mayotte, Nigeria, Sierra Leone, Tanzania, Zambia. Dammhahn and Goodman (2013: 108) indicate that this species' foraging habitat consists of open areas and the area above the forest canopy. ROOST: Ravelomanantsoa et al. (2019: 111) indicated that Mops leucogaster uses human constructions for roosting. PREDATORS: Goodman et al. (2015c: 78) found the remains of four individuals in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. Figure 166. Distribution of Mops (Chaerephon) leucogaster Mops (Chaerephon) major (Trouessart, 1897) *1897. [Nyctinomus (Nyctinomus) pumilus] Var. Major Trouessart, Catalogus Mammalium tam viventium quam fossilium, nov. ed. 1, pt. 1: 146. Publication date: 1897. Type locality: Sudan: N Sudan: 5th Cataract of the Nile [18 32 N 33 41 E] [Goto Description]. 486 ISSN 1990-6471 1901. 1917. 1967. 2020. ? ? ? ? ? Holotype: BMNH 1849.2.8.36: Collected by: Sir Francis Galton. - Comments: Kock (1969a: 148) mentions "Nilus super = 5. Nil-Katarakt, nördlich Berber, Sudan" as original type locality. He also states that de Winton (1901a: 40) changed the type locality in "erster Kakarakt der Nil".). - Etymology: From the Latin "maior" (see Kozhurina, 2002: 16). Nyctinomus Emini de Winton, Ann. Mag. nat. Hist., ser. 7, 7 (37): 40. Publication date: 1 January 1901. Type locality: Tanzania: "Mosambiro" [=Usambiro] [03 00 S 32 34 E] [Goto Description]. Holotype: BMNH 1890.6.8.15: ad ♂, skin and skull. Collection date: 9 September 1889. Notes: Local name - 'Katunke'. - Comments: See Thorn et al. (2009: 72). Chærephon (Lophomops) abæ J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 464, text-fig. 11. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Aba [03 53 N 30 17 E] [Goto Description]. Holotype: AMNH 48887: ad ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 17 December 1911; original number: 1823. Tadarida (Chaerephon) pumila: Happold, Sudan Notes & Records, 48: 122. - Comments: Preoccupied by pumilus Cretzschmar, 1830 (see Simmons, 2005). Attribution to major seems strange. Mops major: Simmons and Cirranello, BatNames.org. (Current Combination) Chaerephon abae: (Alternate Spelling) Chaerephon major: (Name Combination) Tadarida (Chaerephon) major: (Name Combination) Tadarida abae: (Name Combination) Tadarida major: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Chinese: 垂耳犬吻蝠. Czech: morous nubijský. English: Large Wrinkle-lipped Bat, Lappet-eared Wrinkle-lipped Bat, Lappet-eared Free-tailed Bat, Giant gland-tailed bat. French: Tadaride à oreillettes, Grand molosse à glandes caudales. German: Stirnlappen-Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ah; IUCN, 2009; Monadjem et al., 2017z). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon major] (Monadjem et al., 2017z). 2008: LC ver 3.1 (2001) [assessed as "Tadarida major"] (Mickleburgh et al., 2008ah; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004am; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There are no major threats to this species. Some populations are threatened by general habitat loss and disturbance of old buildings (Mickleburgh et al., 2008ah; IUCN, 2009; Monadjem et al., 2017z). CONSERVATION ACTIONS: Mickleburgh et al. (2008ah) [in IUCN (2009)] and Monadjem et al. (2017z) report that the species has a wide range, and likely occurs in some protected areas, although this needs to be confirmed. No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Mops major is a lowland species is found throughout much of West Africa (including Senegal, Liberia, Côte d'Ivoire, Mali, Burkina, Ghana, Togo, Benin, Niger, and Nigeria), as well as a narrow distribution following the Nile from northern Sudan to Uganda. A second population is known from the east of Lake Victoria in eastern Uganda, western Kenya, and north-western Tanzania. A third, apparently disjunct, population is present in southeastern Kenya and northeastern Tanzania. The records from Liberia should be re-examined as they may represent a different species, while the record from Malawi is doubtful (J. Fahr pers. comm. 2004 in Mickleburgh et al., 2008ah in IUCN, 2009). African Chiroptera Report 2020 Native: Benin (Capo-Chichi et al., 2004: 163); Burkina Faso (Kangoyé et al., 2015a: 617); Cameroon (Bakwo Fils et al., 2014: 4); Congo (The Democratic Republic of the); Côte d'Ivoire; Ghana; Guinea; Kenya; Liberia; Mali; Niger; Nigeria; Senegal; Sudan; Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 63); Zambia (Monadjem et al., 2010d: 541). ECHOLOCATION: From Maroua, Cameroon, Manga Mongombe (2012: 79) and Bakwo Fils et al. (2018: 4) reported for 16 calls a call type of FM and the following parameters: Fmax: 39.3 ± 6.9 (31.9 - 53.1) kHz, Fmin: 37.4 ± 6.7 (25.9 - 49.0) kHz, Fmean: 36.4 ± 6.7 (28.7 - 51.0) kHz, Fknee: 38.2 ± 7.3 (29.9 - 53.1) kHZ, Fchar: 33.7 ± 7.3 (26.2 - 49.0) kHz, and duration: 1.66 ± 0.40 (0.85 - 2.73) msec. 487 PARASITES: Beaucournu and Kock (1996) reported the first record for the flea Lagaropsylla anciauxi Smit, 1957 on "Tadarida major" from Burkina Faso. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Burkina Faso, Cameroon, Central African Republic, Congo (Democratic Republic of the), Ghana, Liberia, Mali, Niger, Nigeria, South Sudan, Sudan, Tanzania, Togo, Uganda, Zambia, Zimbabwe. POPULATION: Structure and Density:- This is a relatively common species, found in fairly small colonies (Mickleburgh et al., 2008ah; IUCN, 2009; Monadjem et al., 2017z). Trend:-2016: Stable (Monadjem et al., 2017z). 2008: Stable (Mickleburgh et al., 2008ah; IUCN, 2009). Figure 167. Distribution of Mops (Chaerephon) major Mops (Chaerephon) nigeriae (Thomas, 1913) *1913. Chærephon nigeriæ Thomas, Ann. Mag. nat. Hist., ser. 8, 11 (63): 319. Publication date: 1 March 1913. Type locality: Nigeria: Northern region: Zaria province [ca. 11 01 N 07 44 E] [Goto Description]. Holotype: BMNH 1911.3.22.1: ad ♀. Collected by: A.C. Francis Esq. Presented/Donated by: A.C. Francis Esq. - Etymology: From the genitive scientific Latin feminine substantive nigèriae, meaning "from Nigeria", as the species was described from a Nigerian specimen (see Lanza et al., 2015: 277). 1957. Tadarida (Chearephon) spillmani: Ansell, Ann. Mag. nat. Hist., ser. 12 (10): 539. (Lapsus) 1988. Tadarida chaeraphon nigeriae: Okafor, Misc. Zool., 12: 11. (Lapsus) 2004. Tadarida (Chaeraphon) nigeriae: Okafor, Igbinosa and Ezenwaji, Anim. Res. Int., 1 (1): 64. Publication date: April 2004. (Name Combination, Lapsus) 2020. Mops nigeriae: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon nigerae: (Lapsus) ? Chaerephon nigeriae: (Current Spelling) ? Tadarida (Chaerephon) nigeriae nigeriae: (Name Combination) ? Tadarida nigeriae nigeriae: (Name Combination) ? Tadarida nigeriae: (Name Combination) 488 ISSN 1990-6471 TAXONOMY: LC ver 3.1 (2001) (Mickleburgh et al., 2004af; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2004: considered a vagrant not assessed, not recorded in SA (Friedmann and Daly, 2004). Figure 168. Chaerephon nigeriae (ECJS-77/2009) caught in the Okavango, Botswana. See Willis et al. (2002, Mammalian Species, 710). This may be a complex of two distinct species, with Mops nigeriae in the north of the known distribution and M. spillmanni in the south. We follow Simmons (2005) in currently recognizing spillmanni Monard, 1933 as a subspecies of M. nigeriae. Herkt et al. (2017: Appendix S9) found that the localities in the southern part of the distribution area suggest rather different ecological niches than those for the northern part, which would suggest that more than one species might be included. COMMON NAMES: Afrikaans: Nigeriese losstertvlermuis. Azande (DRC): Fulo. Czech: morous nigerijský. English: Nigerian Free-tailed Bat, Nigerian Wrinkle-lipped Bat. French: Tadaride du Nigeria, Molosse du Nigéria. German: NigeriaBulldoggfledermaus. Italian: Cherefónte nigeriàno. ETYMOLOGY OF COMMON NAME: The species was originally described from a specimen from Nigeria (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008ai; IUCN, 2009; Monadjem et al., 2017d). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Chaerephon nigeriae] (Monadjem et al., 2017d). 2008: LC ver 3.1 (2001) [assessed as "Tadarida nigeriae"] (Mickleburgh et al., 2008ai; IUCN, 2009). 2004: MAJOR THREATS: There appear to be no major threats to this species (Mickleburgh et al., 2008ai; IUCN, 2009; Monadjem et al., 2017d). The ability of the species to exploit buildings as roosts, and its association with cleared rainforest in Nigeria, suggests that it is relatively resistant to anthropogenic impacts (Willis et al., 2002). CONSERVATION ACTIONS: Mickleburgh et al. (2008ai) [in IUCN (2009)] and Monadjem et al. (2017d) report that in view of the species wide range, it is presumably present in some protected areas. No direct conservation measures are currently needed for this species as a whole. GENERAL DISTRIBUTION: Mops nigeriae is widely distributed over much of sub-Saharan Africa. It ranges from Sierra Leone in the west to Ethiopia in the east, being recorded as far south as Angola, Botswana and Zimbabwe. There are some additional records from the Arabian Peninsula in Yemen and western Saudi Arabia. The species has been found at elevations of up to 1,000 m asl, and possibly occur higher. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,133,308 km 2. Native: Angola (Crawford-Cabral, 1989; AlJumaily, 2002: 55; Monadjem et al., 2010d: 541; Taylor et al., 2018b: 60); Botswana (Archer, 1977; Al-Jumaily, 2002: 55; Monadjem et al., 2010d: 541); Burkina Faso (Kangoyé et al., 2012: 6024; 2015a: 618); Burundi; Cameroon; Central African Republic (Al-Jumaily, 2002: 55); Chad (Al-Jumaily, 2002: 55); Congo; Congo (The Democratic Republic of the) (Hayman et al., 1966; Al-Jumaily, 2002: 55; Monadjem et al., 2010d: 541); Côte d'Ivoire (Beaucournu and Fahr, 2003: 157; but see Al-Jumaily, 2002); Ethiopia (Al-Jumaily, 2002: 55); Ghana (Al-Jumaily, 2002: 55); Guinea (Al-Jumaily, 2002: 55); Malawi (Happold and Happold, 1997b: 823; Al-Jumaily, 2002: 55); Mali (Al-Jumaily, 2002: 55); Mozambique; Namibia (Al-Jumaily, 2002: 55; Monadjem et al., 2010d: 541); Niger (Al-Jumaily, 2002: 55); Nigeria (Al-Jumaily, 2002: 55); Saudi Arabia (Nader and Kock, 1980); Sierra Leone (AlJumaily, 2002: 55); Sudan; Tanzania (Al- African Chiroptera Report 2020 Jumaily, 2002: 55); Uganda (Al-Jumaily, 2002: 55); Yemen (Al-Jumaily, 2002: 55; Benda et al., 2011b: 46); Zambia (Ansell, 1957; Ansell and Ansell, 1973; Cotterill, 2004a: 261; Al-Jumaily, 2002: 55; Monadjem et al., 2010d: 541); Zimbabwe (AlJumaily, 2002: 55; Monadjem et al., 2010d: 542). Presence uncertain: Togo (Al-Jumaily, 2002: 55). ECHOLOCATION: Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 17 kHz and duration: 10 msec. Eight calls from the Okavango River Basin reported by Weier et al. (2020: Suppl.) had the following characteristics: Fmax: 16.51 ± 0.57 kHz, Fmin: 14.11 ± 4.77 kHz, Fknee: 16.29 ± 0.54 kHz, Fchar: 16.12 ± 0.43 kHz, slope: 2.99 ± 9.56 Sc, duration: 6.83 ± 4.45 msec. See also Taylor (1999b). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported 2n = 48, FN = 62, BA = 16, a submetacentric X chromosome, and an acrocentric Y chromosome, for specimens from Namibia and Zimbabwe 489 PARASITES: BACTERIA Bartonellae Kamani et al. (2014: 628) tested 16 Nigerian C. nigeriae animals, and found 2 of them testing positive for Bartonella DNA. Of the two Nigerian bats examined by Di Cataldo et al. (2020: 2), one was infected by hemoplasma bacteria. HELMINTHS Okafor et al. (2004: 64) investigated the helminth parasite load of 857 M. nigeriae specimens from Nigeria and found 658 (76.78 %) of them to be infected by 9 species: Nematodes: Rictularia chaerephoni, Histostrongylus coronatus, Capillaria annulosa Dujardin, 1845, Cheiropteronema globocephala Sandground, 1929, Litosoma pujoli Bain, 1966; Trematodes: Posthodendrium panouterus, Castroia nyctali Gvosdev, 1954; Cestodes: Hymenolepis kerivoulae (Prudhoe and Manger, 1969), Oochoristica agamae Bayliss, 1919. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- This species is typically found in small groups of 10 to 15 animals, with maternity roosts being formed of up to 25 bats (Mickleburgh et al., 2008ai; IUCN, 2009; Monadjem et al., 2017d). Trend:- 2016: Unknown (Monadjem et al., 2017d). 2008: Unknown (Mickleburgh et al., 2008ai; IUCN, 2009). Figure 169. Distribution of Mops (Chaerephon) nigeriae Mops (Chaerephon) nigeriae nigeriae (Thomas, 1913) *1913. Chærephon nigeriæ Thomas, Ann. Mag. nat. Hist., ser. 8, 11 (63): 319. Publication date: 1 March 1913. Type locality: Nigeria: Northern region: Zaria province [ca. 11 01 N 07 44 E] [Goto Description]. - Etymology: From the genitive scientific Latin feminine substantive nigèriae, meaning "from Nigeria", as the species was described from a Nigerian specimen (see Lanza et al., 2015: 277). ? Chaerephon nigeriae nigeriae: (Name Combination) ? Tadarida (Chaerephon) nigeriae nigeriae: (Name Combination) ? Tadarida nigeriae nigeriae: (Name Combination) 490 ISSN 1990-6471 GENERAL DISTRIBUTION: E Côte d'Ivoire, Ghana, Benin, Togo, Nigeria, Cameroon, Central African Republic to NE Democratic Republic of the Congo; Niger; Chad; Ethiopia; Saudi Arabia (Harrison and Bates, 1991: 64). Subsaharan Africa to NE Zaire: see Horácek et al. (2000: 137). E Côte d'Ivoire, Ghana, Benin, Togo, Nigeria, Cameroon, Central African Republic, N Zaire, Niger, Chad, Ethiopia: see Willis et al. (2002: 2). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Ghana, Mali, Niger, Nigeria, Sierra Leone, South Sudan. Mops (Chaerephon) nigeriae spillmanni (Monard, 1933) *1933. Nyctinomus spillmanni Monard, Bull. Soc. Neuchât. Sci. nat., 57: 51 (for 1932?). Publication date: 1933. Type locality: Angola: South-Central Angola: Vila da Ponte [14 25 S 16 15 E]. 1963. Tadarida (Chaerephon) spillmanni: Hayman. (Name Combination, Current Spelling) 1965. Nyctinimus spillmani: Koopman, Am. Mus. Novit., 2219: 21. Publication date: 22 June 1965. (Lapsus) 1967. Tadarida nigeriae spillmanni: Ansell, Arnoldia Rhod., 2 (38): 25. (Name Combination) ? Chaerephon nigeriae spillmani: (Lapsus) ? Chaerephon nigeriae spillmanni: (Name Combination) ? Tadarida (Chærephon) spillmani: (Name Combination, Alternate Spelling) GENERAL DISTRIBUTION: Southern Africa (Angola (Horácek et al., 2000: 137), Namibia (Willis et al., 2002: 2), Botswana (Willis et al., 2002: 2), Zimbabwe (Wilson and La Val, 1974: 2), Zambia (Willis et al., 2002: 2), S Democratic Republic of the Congo (Willis et al., 2002: 2), Tanzania (Horácek et al., 2000: 137). PARASITES: SIPHONAPTERA Ischnopsyllidae: Lagaropsylla lipsi Smit,1957 known only from the vicinity of Elisabethville, Katanga, Congo (Haeselbarth et al., 1966: 191). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Botswana, Congo (Democratic Republic of the), Kenya, Malawi, Namibia, South Africa, Tanzania, Uganda, Zambia, Zimbabwe. Mops (Chaerephon) pumilus (Cretzschmar, 1826) *1826. Dysopes pumilus Cretzschmar, in: Rüppell, Atlas Reise Nördlichen Afrika, Zoologie Säugethiere, 1: 69, pl. 27a. Type locality: Eritrea: Massawa [15 37 N 39 26 E] [Goto Description]. Lectotype: SMF 4311: ♀, skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell; collection date: 1825. Old catalog II.K.3.a: see Mertens (1925: 20). - Comments: The publication date of Cretzschmar's work is sometimes indicated as "1826" (as mentioned on the title page), but also as "1827", "1828", "1830-1831". Etymology: From the masculine Latin adjective pùmilus, meaning "of short stature, dwarf or pygmy" as the species is a particularly small African molossid (see Lanza et al., 2015: 282). 1838. Nyctinomus pusilus: Lesson, Compléments de Buffon, ed. 2, Paris, 1: 328. Publication date: 1838. (Lapsus) 1852. Dysopes dubius Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 60, pl. 15, fig. 2. Publication date: 1852. Type locality: Mozambique: South bank of Zambezi River: Sena [17 28 S 35 01 E] [Goto Description]. Holotype: ZMB 85539: juv, skull only. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. Pullus specimen; see Turni and Kock (2008: 72). - Comments: Preoccupied by dubius A. Smith, 1833 (see Simmons, 2005). Allen (1939a: 111) also mentioned Nyctinomus dubius A. Smith, 1833 as a separate species. Turni and Kock (2008: 72) state that no other African molossid collected was ever identified with D. dubius until Ellerman et al. (1953: 68) synonymized it without any comment. Peters´ species description on the pullus specimen ZMB 85539 is very questionable, so they conclude that dubius is a nomen dubium. African Chiroptera Report 2020 1852. 1877. 1877. 1878. 1901. 1904. 1916. 491 Dysopes limbatus Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 56, pl. 14. Publication date: 1852. Type locality: Mozambique: Mozambique Island [15 03 S 40 46 E] [Goto Description]. Syntype: BMNH 1907.1.1.704:. Syntype: ZMB 537/85520: ad ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008) [skin: ZMB 537 / skull: ZMB 85520]. Syntype: ZMB 538/85519: ad ♀, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008) [skin: ZMB 538 + skull: ZMB 85519]. Syntype: ZMB 539: juv, skin only. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008). - Comments: Peters (1852: 56) mentions "Africa orientalis, Insula Mozambique, Sena at 15° ad 17° Let. Aust.", and Roberts (1951: 98) "Mozambique Island and Sena"). Possibly valid as a subspecies. Considered a valid species by Peterson et al. (1995: 159). Nyctinomus [(Nyctinomus)] limbatus: Dobson, Proc. zool. Soc. Lond., 1876, IV: 724. Publication date: April 1877. (Name Combination) Nyctinomus [(Nyctinomus)] pumilus: Dobson, Proc. zool. Soc. Lond., 1876, IV: 723. Publication date: April 1877. (Name Combination) Nyctinomus limbatus: Dobson, Catalogue of the Chiroptera of the collection of the British Museum, 428. - Comments: . (Name Combination) Nyctinomus gambianus de Winton, Ann. Mag. nat. Hist., ser. 7, 7 (37): 39. Publication date: 1 January 1901. Type locality: The Gambia: "Gambia" [Goto Description]. Holotype: BMNH 1889.10.7.3:. Nyctinomus hindei Thomas, Ann. Mag. nat. Hist., ser. 7, 13 (75): 210. Publication date: 1 March 1904. Type locality: Kenya: Fort Hall [=Muranga] [00 43 S 37 09 E, 4 000 ft] [Goto Description]. Holotype: BMNH 1903.3.2.4: ad ♂. Collected by: Mrs. Hildegarde Hinde; collection date: 1 January 1903; original number: 134. Presented/Donated by: Mrs. Hildegarde Hinde. - Comments: Possibly valid as a subspecies. Considered a valid species by Peterson et al. (1995: 159). Thomas (1904a: 210) mentions 2 specimens. Chaerephon pumilus naivashæ Hollister, Smiths. Misc. Coll., 66 (1) (2406): 4. Publication date: 10 February 1916. Type locality: Kenya: Naivasha Station [00 43 S 36 26 E] [Goto Description]. Holotype: USNM 166658: ad ♂, skull and alcoholic. Collected by: John Alden Loring; collection date: 7 August 1909; original number: 6955. Topotype: ROM 36466: ad ♂, skin and skull. Collected by: John George Williams; collection date: 10 July 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 36467: ad ♀, skin and skull. Collected by: John George Williams; collection date: 17 July 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 36468: ad ♀, skin and skull. Collected by: John George Williams; collection date: 17 July 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 36469: ad ♀, skin and skull. Collected by: John George Williams; collection date: 17 July 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 36470: ad ♂, skin and skull. Collected by: John George Williams; collection date: 17 July 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 36471: ad ♀, skin and skull. Collected by: John George Williams; collection date: 10 August 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 36472: ad ♀, skin and skull. Collected by: John George Williams; collection date: 10 August 1965. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41842: juv ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41843: juv ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41844: juv ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41845: juv ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41846: juv ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41847: juv ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and 492 ISSN 1990-6471 Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41848: juv ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41849: juv ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41850: ad ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41851: ad ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41852: ad ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41853: ad ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41854: ad ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41855: ad ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41856: ad ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41857: ad ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41858: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41859: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41860: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41861: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41862: juv ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41863: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41864: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41865: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41866: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41867: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41868: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41869: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41870: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41871: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; African Chiroptera Report 2020 493 collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41872: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41873: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41874: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41875: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41876: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41877: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41878: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41879: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41880: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41881: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41882: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41883: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41884: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41885: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41889: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41890: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41891: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41892: ad ♂, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41893: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41894: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41895: ad ♀, skull and alcoholic. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41896: ad ♂, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41897: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41898: ad ♀, alcoholic (skull not removed). Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41899: ad ♀, skin and skull. Collected by: Randolph Lee 494 ISSN 1990-6471 Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41900: ad ♂, complete skeleton. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41901: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41902: ad ♀, complete skeleton. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41903: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41904: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41905: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41906: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41907: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 41908: ad ♀, skin and skull. Collected by: Randolph Lee Peterson, John George Williams and Robert M. Glen; collection date: 9 August 1967. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47000: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47001: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47002: ad ♂, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47003: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47004: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47005: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47006: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47007: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47008: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47009: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47010: ad ♂, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47011: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47012: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47013: ad ♂, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47014: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47015: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47016: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47017: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47018: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. African Chiroptera Report 2020 1926. 1932. 1951. 1975. 1999. 2017. 2019. 2020. ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 495 Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47019: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47020: ad ♂, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. Topotype: ROM 47021: ad ♀, skin and skull. Collected by: John George Williams; collection date: 30 June 1968. Presented/Donated by: ?: Collector Unknown. - Comments: Considered a valid species by Peterson et al. (1995: 157). Chaerophon pumilus elphicki Roberts, Ann. Transv. Mus., 11: 245. Publication date: 14 September 1926. Type locality: South Africa: SE Transvaal province: Barberton district: Malelane Estate [25 29 S 31 31 E] [Goto Description]. Holotype: TM 2488: ad ♂, skin and skull. Collected by: Captain G.J. Elphick; collection date: 30 March 1920. Choerephon (Lophomops) langi Roberts, Ann. Transv. Mus., 15 (1): 17. Publication date: 1 October 1932. Type locality: Botswana: N Bechuanaland, Ngamiland,: Tsotsoroga Pan [18 40 S 24 25 E] [Goto Description]. Holotype: TM 6554: ad ♂, skin and skull. Collected by: The Vernay-Lang Kalahari Expedition 1930; collection date: 29 June 1930; original number: 870. - Comments: Possibly valid as a subspecies. Tadarida (Chaerephon) faini Hayman, Rev. Zool. Bot. afr., 45 (1-2): 82. Publication date: 22 December 1951. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Upper Congo, between Blukwa and Lake Albert: Wago Forest [ca. 01 44 N 30 39 E] [Goto Description]. Holotype: RMCA 20509: ad ♂, skin and skull. Collected by: Dr. Alexandre Fain; collection date: 1 June 1951; original number: 1. Paratype: BMNH 1951.629: ad ♀, skin and skull. Collected by: Dr. Alexandre Fain; original number: 2. Presented/Donated by: ?: Collector Unknown. - Comments: Possibly valid as a subspecies. Chaerophon limbatus: Durette-Desset and Chabaud, Ann. Parasit., 50 (3): 315. (Lapsus) Chaerephon pumilla: Dumont, Etzel and Hempel, Mammalia, 63 (2): 159. (Lapsus) Chaerophon pumiluss: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 5. Publication date: 6 April 2017. (Lapsus) Chaerephon pumilas: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) Mops pumilus: Simmons and Cirranello, BatNames.org. (Current Combination) Chaerephon cf. pumilus: Chaerephon limbatus: (Name Combination) Chaerephon pumila hindei: (Name Combination) Chaerephon pumila langi: (Name Combination) Chaerephon pumila limbata: (Alternate Spelling) Chaerephon pumila limbatus: (Name Combination) Chaerephon pumilus elphicki: (Name Combination) Chaerephon pumilus hindei: (Name Combination) Chaerephon pumilus langi: (Name Combination) Chaerephon pumilus limbatus: (Name Combination) Chaerephon pumilus naivashae: (Alternate Spelling) Chaerephon pumilus pumilus: (Name Combination) Chaerephon pumilus sensu lato: Chaerephon pumilus: (Name Combination) Chaerephon pusillus: Chaerophon pumilus: (Name Combination) Tadarida (Chaerephon) pumila frater: (Name Combination) Tadarida (Chaerephon) pumila: (Name Combination) Tadarida faini: (Name Combination) Tadarida limbata: (Name Combination) Tadarida naivashae: (Name Combination) Tadarida pumila elphicki: Tadarida pumila faini: (Name Combination) Tadarida pumila frater: (Name Combination) Tadarida pumila hindei: (Name Combination) Tadarida pumila langi: (Name Combination) Tadarida pumila limbata: (Name Combination) Tadarida pumila naivashae: (Name Combination) 496 ISSN 1990-6471 ? ? ? ? ? Tadarida pumila pumila: (Name Combination) Tadarida pumila pumila-limbata: Tadarida pumila pumila-naivashae: Tadarida pumila websteri: (Name Combination) Tadarida pumila: (Name Combination) TAXONOMY: Figure 170. Chaerephon pumilus. See Bouchard (1998, Mammalian Species, 574). Formerly included pusillus; see Hayman and Hill (1971: 64), but see Goodman and Ratrimomanarivo (2007: 391). The description date is sometimes mentioned as "1830 or 1831" (see Happold and Happold (1997b: 823). Peterson et al. (1995: 162) consider pumilus not to include most of the forms occurring on Madagascar. Horácek et al. (2000: 136) include pumilus in the genus Tadarida, subgenus Chaerephon. Taylor (1999b), Aspetsberger et al. (2003: 250; 252) indicate that cryptic species might exist within the genus Chaerephon (more specifically in pumilus), but find it premature to identify them already. Jacobs et al. (2004ar: 17) identified five haplotypes, but indicate they represent only one species. Naidoo et al. (2013c: 134) indicate that southeast African M. pumilus s.l. contains several wellsuported sympatric genetic lineages (they identified three clades), which exibit paraphyly with the Malagasy M. leucogaster. They were not able to to distinguish these clades based on echolocation characteristics (p. 137), but suggest that these stable sympatric lineages might be the result of female philopatry or roost faithfulness. Naidoo et al. (2016) used cyt b, control region, and Rag2 data on a large number of representatives from the Mops pumilus species complex from islands in the western Indian Ocean, and eastern and southeastern Africa. They found that Mops pumilus s.s. (Eritrea and Yemen) is specifically distinct from the following taxa/samples: M. pumilus from southeastern Africa, the syntype of limbatus, the holotypes of elphicki and langi, and the topotype of naivashae. They suggest (p. 642) that additional evidence from more variable nuclear markers is necessary to resolve the issue of the number of species present in this complex. The genetic differentiation is probably the result of habitat fragmentation during glacial maxima when the ancestral population was separated over isolated subtropical forests (Schoeman and Monadjem, 2018: 4). COMMON NAMES: Afrikaans: Klein losstertvlermuis. Chinese: 小犬 吻 蝠 . Czech: morous malý, tadarida malá. English: Little Free-tailed Bat, Lesser Free-tailed Bat, Little Wrinkle-lipped Bat, White-bellied Freetailed Bat. French: Petite tadaride, Petit molosse à glandes caudales, Tadaride bornée. German: Kleine Bulldoggfledermaus, ZwergDoggengrämler, Sena-Doggengrämler, Gesäumte Doggengrämler. Italian: Cherefónte pigmèo. Kiluba (DRC): Kasusu. Nyungwe (Malawi): Kalemawalema (not specific). Portuguese: Morcego pequeno de cauda livre. ETYMOLOGY OF COMMON NAME: From the Latin "pumilus" meaning dwarf, because it is the smallest of the molossids occuring in the (southern African sub)region (see Taylor, 2005). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a broad range of habitats, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008dr; IUCN, 2009; Mickleburgh et al., 2014b). Assessment History Global 2014: LC ver 3.1 (2001) assessed as "Chaerephon pumilus" (Mickleburgh et al., 2014b). 2008: LC ver 3.1 (2001) assessed as "Tadarida pumila" (Mickleburgh et al., 2008dr; IUCN, 2009). 2004: LC ver 3.1 (2001) assessed as "Chaerephon pumila" (Mickleburgh et al., 2004ah; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016a). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). African Chiroptera Report 2020 MAJOR THREATS: There are no major threats to the species. In some parts of its range, it is threatened from persecution as a pest, especially since it roosts in buildings (Mickleburgh et al., 2008dr; IUCN, 2009; Mickleburgh et al., 2014b). CONSERVATION ACTIONS: Mickleburgh et al. (2014b) reported that due to its wide range it seems probable that this species is present in some protected areas. On Madagascar, it has not been recorded from any protected areas, but it has been recorded from houses close to protected areas. The status of the population on Aldabra need to be clarified with complementary research needed into the conservation status of this population. GENERAL DISTRIBUTION: This widespread species is found from Senegal in the west of its range, eastwards to Yemen and southwestern Saudi Arabia, and as far south as South Africa, where Babiker Salata (2012: 49) found that its distribution is stongly associated with the mean temperature of the coldest quarter. It has been recorded from the island of Bioko and the Annobon Islands (Equatorial Guinea), Pemba and Zanzibar (Tanzania). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 809,867 km2. Records from Western Cape Province are rejected by Jacobs and Fenton (2001), since these represent Sauromys petrophilus. Records from the Seychelles [Aldabra] (Hutson, 2004a) and the Comoros (Louette, 2004; Goodman, 2007; ACR, 2006, 2007, 2008, 2009, 2010) are rejected by Goodman et al. (2010a) since these represent Mops pusillus. Records from Madagascar (Bouchard, 1998; Russ et al., 2001; Goodman and Cardiff, 2004; ACR, 2006, 2007, 2008, 2009,2010) are now assigned to Mops atsinanana (Goodman et al., 2010a). Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 542; Taylor et al., 2018b: 62); Benin (CapoChichi et al., 2004: 164); Botswana (Archer, 1977; Monadjem et al., 2010d: 542); Burkina Faso (Kangoyé et al., 2015a: 618); Burundi; Cameroon (Bakwo Fils et al., 2014: 4): Chad; Congo (Bates et al., 2013: 336); Congo (The Democratic Republic of the) (Allen, 1917; Schouteden, 1944; Hayman et al., 1966; Dowsett et al., 1991: 259; Monadjem et al., 2010d: 542); Côte d'Ivoire; Equatorial Guinea [Annobón, Bioko]; Eritrea; Ethiopia (Lavrenchenko et al., 2004b: 132; Benda et al., 2020: 2586); Gambia (Emms and Barnett, 497 2005: 50); Ghana; Guinea; Guinea-Bissau (Monard, 1939; Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 73); Kenya; Liberia (Fahr, 2007a: 104); Malawi (Happold et al., 1988; Ansell and Dowsett, 1988: 46; Monadjem et al., 2010d: 542); Mali (Meinig, 2000: 105); Mozambique (Smithers and Lobão Tello, 1976; Lopes and Crawford-Cabral, 1992; Monadjem et al., 2010d: 542; Monadjem et al., 2010c: 382); Namibia (Monadjem et al., 2010d: 542); Niger; Nigeria; Rwanda; Sao Tomé and Principe [Sâo Tomé] (Juste and Ibáñez, 1993b: 906; Rainho et al., 2010a: 37); Saudi Arabia; Senegal; Sierra Leone; Somalia; South Africa (De Graaf, 1960; Monadjem et al., 2010d: 542); Sudan; Swaziland (Monadjem et al., 2010d: 543); Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 62); Yemen (Benda et al., 2011b: 47); Zambia (Ansell, 1969; Ansell, 1973; Ansell, 1978; Monadjem et al., 2010d: 543); Zimbabwe (Cotterill, 2004a: 261; Monadjem et al., 2010d: 543). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: From western Uganda, Monadjem et al. (2011: 30) reported the following data: Fa: 38.95 mm, mass: 10.5 g, wing loading: 12.4 N/m 2, aspect ratio: 7.3. DETAILED MORPHOLOGY: Baculum - Unknown Brain - Amrein et al. (2007) from specimens collected from Bénin reported moderate proliferating cell activity, detected with Ki-67 and MCM2, in the subgranular layer of the dentate gyrus, with doublecortin (DCX) they detected young migrating neuron in the subgrannular layer of the hippocampus. Amrein et al. (2007) also reported moderate to ample proliferating cells (Ki67 positive cells) and migrating young neurons (DCX positive cells) in the rostral migratory stream. No NeuroD was detected in the hippocampal granule cells (Amrein et al., 2007). Kruger et al. (2010a) describe, based on three brains from Kenya, using immunohistochemical methods, the nuclear organization of the cholinergic, catecholaminergic and serotonergic systems. Kruger et al. (2010b) describes, based on three brains from Kenya, the distribution of Orexin-A immunoreactive cell bodies and terminal networks. Gastro-intestinal tract (GIT): Aylward et al. (2019: 1108) reported for six specimens an average weight of 9.17 ± 0.85 g, and a GIT of 0.44 ± 0.05 g (4.85 ± 0.40 %). SEXUAL DIMORPHISM: Chaverri et al. (2018: 1944) [referring to Bouchard, 2001b] indicates that the sexes can be distinguished by scent produced from the interaural and muzzle glandular areas. 498 ISSN 1990-6471 ECHOLOCATION: Frequency modulated call. In Kenya, Taylor et al. (2005) reported a peak frequency of 25.6 (1.7) kHz from Kivoko. In Tanzania, Aspetsberger et al. (2003) reported a maximum frequency of 22.7 (3.3) kHz from Amani Nature Reserve. In Swaziland, Taylor (1999b) reported a maximum frequency of 32.9 (4.5) kHz from Mlawula, and maximum frequencies at three different localities around Durban in South Africa of 43.0 (1.0) kHz, 28.7 (1.8) kHz and 28.7 (2.5) kHz. Fenton et al. (2004) reported a highest frequency of 29 (4.6) kHz for a locality in Durban, South Africa. Rainho et al. (2010a: 21) reports the calls [as Tadarida pumila] of 32 individuals from Sao Tomé. Bohmann et al. (2011: 1) [referring to Monadjem et al. (2010d)] mention peak frequencies between 25 - 40 kHz. Schoeman and Waddington (2011: 291) mention a peak frequency of 28.2 ± 1.5 kHz and a duration of 10.8 ± 0.2 msec for specimens from Durban, South Africa. From the Durban area, Naidoo et al. (2011: 23) report Fpeak = 24.7 ± 1.0 kHz and a duration of 12.4 ± 0.9 msec. Taylor et al. (2013b: 18) recorded the following parameters for 8 calls from specimens from Swaziland: Fmax: 27.7 ± 4.40 (23.0 - 32.8) kHz, Fmin: 23.3 ± 2.38 (20.9 - 27.4) kHz, Fknee: 26.2 ± 3.34 (22.6 - 30.8) kHz, Fchar: 24.3 ± 2.91 (21.5 - 28.1) kHz, duration: 8.3 ± 2.10 (5.3 - 11.9) msec, and for 6 calls from KwaZulu-Natal, South Africa: Fmax: 31.6 ± 4.05 (24.6 - 36.7) kHz, Fmin: 24.5 ± 1.55 (21.7 - 26.5) kHz, Fknee: 28.9 ± 2.51 (24.4 - 31.3) kHz, Fchar: 25.6 ± 1.57 (22.7 - 27.3) kHz, duration: 8.4 ± 1.71 (6.6 - 11.0) msec. From Swaziland also, Monadjem et al. (2017c: 179) reported the following values: Fmin: 23.3 ± 1.41 (20.8 - 26.5) kHz, Fknee: 26.1 ± 2.09 (22.5 - 29.6) kHz, Fc: 24.0 ± 1.53 (21.3 - 27.2) kHz and duration: 9.0 ± 2.10 (3.9 - 12.6) msec. They were able to detect these calls up to a distance of 15 m. At Farm Welgevonden, South Africa, Taylor et al. (2013a: 556) report a Fknee value of 29 (24 - 31) kHz. Linden et al. (2014: 40) mentioned the following parameters for the Soutpansberg area (RSA): Fmin: 21 - 27 kHz, Fchar: 22 - 28 kHz, Fknee: 23 - 31 Khz, Slope:13 - 124 OPS, duration: 5 - 12 msec. 20 calls from Mapungubwe National Park (RSA) were recorded by Parker and Bernard (2018: 57) with the following characteristics: Fchar: 26.65 ± 2.53 kHz, Fmax: 34.87 ± 5.75 kHz, Fmin: 24.67 ± 2.68 kHz, Fknee: 29.72 ± 3.05 kHz, duration: 8.80 ± 2.65 msec, with 4.24 ± 1.89 calls/sec. Monadjem et al. (2011: 32) reported values for two calls from a "Chaerephon cf. pumilus" from western Uganda: Fmin: 24.95 kHz, Fmax: 27.78 kHz, Fchar: 25.22 kHz, Fknee: 75.61 kHz, and duration 8.4 msec. Two calls from specimens from the Aberdares Range in Kenya were reported by Eisenring et al. (2016: SI 2) with PF: 26.3 ± 2.8 (24.4 - 28.3), HF: 32.6 ± 1.2 (31.8 - 33.5), LF: 20.9 ± 0.4 (20.6 - 21.1), DT: 0.1 ± 0.1 (0.0 - 0.1), DF: 11.8 ± 1.5 (10.7 12.9). From Maroua, Cameroon, Manga Mongombe (2012: 79) reported for 3 calls a call type of FM and the following parameters: Fmax: 44.2 ± 3.4 (40.4 47.0) kHz, Fmin: 38.2 ± 2.6 (35.3 - 40.4) kHz, Fmean: 41.2 ± 3.0 (37.9 - 43.7) kHz, Fknee: 44.1 ± 3.4 (40.4 - 47.0) kHZ, Fchar: 39.1 ± 2.4 (36.5 - 41.2) kHz, and duration: 1.32 ± 0.11 (1.26 - 1.45) msec. Two calls by bats from the Kalahari Desert had the following characteristics: Fchar: 22.9 ± 0.3 kHz, Fmax: 25.0 ± 0.1 kHz, Fmin: 22.4 ± 0.4 kHz and duration: 5.6 ± 1.5 msec (Adams and Kwiecinski, 2018: 4). Weier et al. (2020: Suppl.) reported on 11 calls from the Okavango River Basin with the following characteristics: Fmax: 29.69 ± 4.89 kHz, Fmin: 22.96 ± 1.91 kHz, Fknee: 25.46 ± 1.84 kHz, Fchar: 23.65 ± 1.71 kHz, slope: 35.10 ± 33.29 Sc, duration: 8.54 ± 4.71 msec. Luo et al. (2019a: Supp.) reported the following data (bats leaving their roost): Fpeak: 23.9 kHz, Fstart: NA kHz, Fend: NA kHz, Band width: NA kHz, and duration: 13.6 msec. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Ðulic and Mutere (1973a), Smith et al. (1986), Rautenbach et al. (1993) and Sreepada et al. (2008) all reported 2n = 48, and an acrocentric Y chromosome. However, while Ðulic and Mutere (1973a) and Smith et al. (1986) reported FN = 58, BA = 12, Rautenbach et al. (1993) reported FN = 66, BA = 20. And, while Ðulic and Mutere (1973a) and Rautenbach et al. (1993) reported a metacentric X chromosome, Smith et al. (1986) reported a submetacentric X chromosome. The work of Rautenbach et al. (1993) included specimens from the Western Cape that have subsequently been reidentified as Sauromys petrophilus (Jacobs and Fenton, 2001: 134), hence these karyotypic variations should be viewed with caution. Reddy (2008: ii) used the cytochrome b (845 nucleotides) and D-loop (314 nucleotides) regions of the mitochondrial DNA to assess the phylogenetic relationships within M. pumilus (southern Africa) and in relation to "Chaerephon" African Chiroptera Report 2020 species from Madagascar (Mops pumilus, M. leucogaster). The result indicates the possibility of cryptic species in southern African M. pumilus, which are closer related to the Malagasy M. leucogaster than to the Malagasy M. pumilus. In South Africa, Naidoo et al. (2015b) were able to retrieve a strong mitochondrial structure, but only little nuclear genetic structure, which suggests a single interbreeding population. They found these data to be consistent with female philopatry, secondary contact between diverged genetic lineages and introgression. Protein / allozyme - Unknown. HABITAT: Noer et al. (2012: 1) found that M. pumilus prefered to forage over sugarcane fields rather than over savanna, riparian forest and urban areas. To reach their foraging grounds, they traveled up to 4.2 km from their roost, and their mean activity area encompases 976 to 1319 ha. HABITS: Bouchard (2001b: 109) found that, in scent-choiseexperiments, only males were able to distinguish the gender of conspecifics as well as the difference between roostmates and strangers. Females did not show any preference during these experiments. ROOST: Taylor et al. (1999: 67) found these bats to be roosting in a variety of roofs in the Durban area: from corrugated iron to tiles, between the tiles and the plastic lining underneath, or in any convenient crevices between rafters and brickwork. These roofs were located at a height of 3 to 12 m. In the Mount Nimba area, Monadjem et al. (2016y: 367) found a roost in the roof of a school at 540 m. DIET: Based on DNA analysis of 59 fecal pellets collected between April and May (austral summer) in the Simunye region of Swaziland, Bohmann et al. (2011) reported insects from 24 families (belonging to seven orders) being taken as prey. The highest frequency of occurrence was for Lepidoptera (52.5 %), Diptera (40.7 %), and Hemiptera (18.6 %). Within the Lepidoptera, Noctuidae (Owlet moths - 25.4 %), Nymphalidae (Brush-footed butterflies 30.5 %) and Crambidae (Grass moths - 10.2 %) were found the most. Within the Diptera, it were representatives of the Culicidae (Mosquitoes - 27.1 %), which occurred most frequently, followed by the Tephritidae (Fruit flies and kin - 10.2 %). In the Hemiptera, both Lygaeidae (Chinch bugs and seed bugs) and 499 Pentatomidae (Stink bugs) were found in 10.2 % of the fecal pellets. Naidoo et al. (2011: 26) reported the following volume percentages in the bat's diet at Umbilo River (KwaZulu Natal, RSA): in summer: Lepidoptera (9.9 ± 9.5), Coleoptera (47.7 ± 30.5), Hemiptera (32.8 ± 6.9), Diptera (5.4 ± 8.3), and Trichoptera (4.2 ± 7.2); in winter: Lepidoptera (30.3 ± 16.0), Coleoptera (19.5 ± 16.5), Hemiptera (8.0 ± 7.4), Diptera (24 ± 7.7), Trichoptera (6.5 ± 10.4), and Hymenoptera (11.8 ± 11.1). In the Limpopo province (RSA), Taylor et al. (2017b: 246, 252 - 254) found the diet to contain 21 prey items, covering five insect orders: Coleoptera (unmatched), Hemiptera (Macrorhaphis acuta + Nezara viridula + unmatched), Lepidoptera (Toxocampinae tathorhynchus + Isturgia roraria + Sena sp. + Sena prompta + Cortyta canescens + Enmonodia capensis + Leucania imperfecta + Pericyma atrifusa + Rhesala moestalis + Spodoptera exigua + Maurilia arcuata + Stenoptilia sp. + unmatched), Orthoptera (Tettigoniinae + unmatched) and Blattodea (unmatched). They also remarked that the diet of C. pumilus varies geographically. From Maroua (Cameroon), and based on fecal analyses, Bol A Anong (2013: 31) reported the diet to include Hemiptera (35.60%), Coleoptera (34.81%), Lepidoptera (17.00%), Diptera (10.22%), Hymenoptera (1.55%) and Orthoptera (0.02%). PREDATORS: At the Kruger Park, Rautenbach et al. (1990: 17) reported that "Tadarida pumila" was regularly taken as prey by a pair of African goshawks (Accipiter tachiro (Daudin, 1800)). On some occasions, 1 to 5 bats were taken per evening (see Kemp and Rautenbach, 1987). Mikula et al. (2016: Supplemental data) mention the following avian predators for "Tadarida pumila": African goshawk (Accipiter tachiro (Daudin, 1800)), Fox kestrel (Falco alopex (Heuglin, 1861)), Lanner falcon (Falco biarmicus Temminck, 1825), Eurasian hobby (Falco subbuteo Linnaeus, 1758), Common kestrel (Falco tinnunculus Linnaeus, 1758), Wahlberg's eagle (Hieraaetus wahlbergi (Sundevall, 1851)), Bat hawk (Macheiramphus alcinus Bonaparte, 1850), Black kite (Milvus migrans (Boddaert, 1783)). POPULATION: Structure and Density:- The colonies of this species range from a few animals (between 5 and 20) to hundreds of individuals (Mickleburgh et al., 2008dr; IUCN, 2009; Mickleburgh et al., 2014b). There are no estimates of its population in 500 ISSN 1990-6471 Madagascar but it is thought to be locally common in the eastern zone. On Aldabra, the population might be fewer than 250 animals. Trend:- 2014: Unknown (Mickleburgh et al., 2014b). 2008: Unknown (Mickleburgh et al., 2008dr; IUCN, 2009). REPRODUCTION AND ONTOGENY: van der Merwe et al. (1986: 355) indicate that in Transvaal M. pumilus has an extended breeding of 8 months. They found females to be polyoestrous and could have up to three pregnancies (one young) per breeding season. Implantation only occurs in the right uttering horn and the gestation period was about 60 days. The young are weaned before 21 days of age and females become sexually mature at the age of 5 to 12 months. Happold and Happold (1989b: 133) found that in Malawi, Mops pumila can have up to 5 births per year, all followed by a post-partum oestrus. Happold and Happold (1990b: 568) mention two to four consecutive litters, with births at the end of November, January, March, and May. Females giving birth in November, all had a second litter in January, and 44 % had a third litter in March. Krutzsch (2000: 95) refers to Marshall and Corbet (1959), who pointed out that in Uganda spermatogenesis was not continuous in all males (as "Tadarida (Chaerephon) hindei"). They also indicated that it is one of the best known virtually continuous breeding molossids. le Grange et al. (2011: 173 - 174) indicate that M. pumilus is a seasonal polyoestrous bat, which reduces the number of reproductive cycles per season with increasing latitute. In Ethiopia, Kruskop et al. (2016: 61) found a maternity colony with both pregnant and lactating females (one even in both stages) on 9 November 2012. Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] mentioned births in early November, late January and April. MATING: McWilliam (1988a) and McCracken and Wilkinson (2000: 331) indicate that there is a correlation between the male body size and the harem size, where larger males have larger harems, consisting of up to 21 females). McWilliam (1988a) also suggests that the harem might be kin groups. PARASITES: BACTERIUM Gram-negative bacterium - Cassel-Béraud and Richard (1988), Cassel-Beraud et al. (1989: 234), and Sara (2002: 41) reported Alkalescens dispar, Aeromonas hydrophila, Citrobacter freundii, Enterbacter aerogens, Enterobacter agglomerans, Enterobacter cloacae, Enterobacter intermedius, Enterobacter sp., Escherichia coli, Hafnia alvei, Klebsiella oxytoca, Klebsiella pneumoniae, Kluyvera sp., Koserella trabulsii, Proteus mirabilis, Proteus morganii, Proteus rettgeri, Proteus vulgaris, Providencia sp., Pseudomonas spp., Salmonella enterica subsp. enterica, Salmonella enteritidis, and Serratia marcescens. APICOMPLEXA Lainson and Naiff (2000: 128) and Duszynski (2002: 20) indicate that "Nyctinomus limbatus / C. pumila" is the type host of Eimeria dukei Lavier, 1927. ACARI Laelapidae: Taufflieb (1962: 112) reported Chelanyssus aethiopious Hirst, 1921 from "Tadarida limbata" from Brazzaville, Congo. Sarcoptidae: Fain (1959d: 149) described Notoedres (Notoedres) tadaridae from "Tadarida (Chaerephon) faini" from Mount Wago, near Blukwa, Haut-Ituri, DRC. Trombiculidae: Stekolnikov (2018a: 163) reported Microtrombicula nyctinomi (Taufflieb, 1960) from a bat from Brazzaville, Congo. HEMIPTERA Cimicidae: Haeselbarth et al. (1966: 9) found a great number of Loxaspis barbara (Roubaud, 1913) in a roost, near Mopti, Former French Sudan (Haute Senegal-Niger). Loxaspis miranda Rothschild, 1912 recorded in Uganda (Haeselbarth et al., 1966: 10, host referred to as "Tadarida limbata"). Polyctenidae: Hypoctenes clarus Jordan, 1922, from Congo (Haeselbarth et al., 1966: 18, host referred to as "Tadarida limbata"). DIPTERA Streblidae: Raymondia seminuda Jobling, 1954 in Nairobi, Kenya (Haeselbarth et al., 1966: 103, host referred to as "Tadarida faini"). Nycteribiidae: Basilia ansifera Theodor, 1956 mainly a West African species, reaching Lake Victoria in the east and Lake Nyasa in the south, nearly all localities lie between 15° latitude north and south (Haeselbarth et al., 1966: 110). Dipseliopoda biannulate (Oldroyd, 1953) distributed over the tropical parts of Africa and recorded from localities in Nigeria, Cameroon, the Congo and the Sudan and Kenya (Haeselbarth et al., 1966: 116, host referred to as "Tadarida faini"). African Chiroptera Report 2020 SIPHONAPTERA Ischnopsyllidae: Lagaropsylla consularis Smit, 1957 from Ethiopia, Kenya, Uganda, Congo and Congo (Democratic Republic of), Angola, Zimbabwe and Mozambique (Haeselbarth et al., 1966: 1990, host referred to as "Tadarida limbata"), Malawi (Beaucournu and Kock, 1996), Madagascar (Hastriter, 2016: 16). Lagaropsylla idae Smit ,1957 widespread in tropical Africa (Haeselbarth et al., 1966: 190, host referred to both "Chaerephon pumilus" as well as "Tadarida limbata"). Lagaropsylla oblique Smit, 1957 from Sierra Leone, Cameroon and the Congo (Haeselbarth et al., 1966: 191, host referred to as "Tadarida limbata"). NEMATODA Durette-Desset and Chabaud (1975: 315) described Molinostrongylus benexae from a "Chaerophon limbatus" from Périnet, Madagascar. TREMATODA Prosthodendrium (Prosthodendrium) chilostomum madagascariense was described by Richard (1966: 423) from a "Chaerophon limbatus" from Périnet, Madagascar. VIRUSES: Conrardy et al. (2014) tested 6 bats sampled in Kenya for the presence of adenoviruses, rhabdoviruses and paramyxoviruses, all samples tested negative. In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Mops pumilus: Astroviruses and Coronaviruses. In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against dengue 2 virus (DENV-2) and non-specific Flaviviruses. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 to 1999, for antibody to SARS-CoV in sera in 35 individuals from Limpopo Province, South Africa, none were tested positive (0/35) and 18 individuals from Mpumalanga Province, South Africa, none tested positive (0/18) and one individual from Bandundu Province, DRC, not tested positive (0/1). Tong et al. (2009) report the detection of coronavirus RNA from fecal material of this species in Kenya and was found to be distantly related to the human coronavirus 229E. Anthony et al. (2017b: Suppl.) mention the alphacoronavirus: Charephon_CoV_KY22. Nziza et al. (2019: 156) reported this virus in a rectal swap from a bat from Rwanda. 501 Filoviridae Ebolavirus: EFSA AHAW Panel (EFSA Panel on Animal Health and W (2017: 5) refer to Swanepoel et al. (1996), who experimentally inoculated the virus in this bat. Goldstein et al. (2018: 1084) described a new Ebolavirus from three Sierra Leonan Mops pumilus and one M. condylurus: Bomali virus (BOMV). They were sampled in human dwellings in small villages in the Bomali district. This virus might infect humans, but it is not known (yet) whether it is virulent in humans too. Flaviviridae Flavivirus Bukalasa bat virus - Williams et al. (1964) report the isolation of the virus in Uganda, 1964, from the salivary glands of Mops pumilus. Reported by de Jong et al. (2011: 10) and Luis et al. (2013: suppl). Dakar bat virus - Kemp et al. (1974) report the isolation of the virus from Nigeria, 1974, from the salivary glands and organ pools of Mops pumilus. Reported by de Jong et al. (2011: 10) and Luis et al. (2013: suppl). Entebbe bat virus. - Peat and Bell (1970) report the isolation of the virus in Uganda, 1957, from the salivary glands of M. pumilus. Reported by de Jong et al. (2011: 10). Zika virus (ZIKV) - Malmlov et al. (2019: 2) refer to Shepherd and Williams (1964) who examined 44 pumilus specimens in Uganda, of which 16 had antibodies against ZIKV. Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 25 Tanzanian Mops pumilus specimens but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). Kia et al. (2014: 1) tested 33 specimens from Kanke (Nigeria), and found one of them to test positive for Lagos bat virus and one for Shimoni bat virus. Togaviridae Alphavirus de Jong et al. (2011: 10) reported Chikungunya virus occurring on M. pumilus. Luis et al. (2013: suppl.) also reported this virus, but used "Tadarida pumila" as name for the bat. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Egypt, Equatorial Guinea, Eritrea, Eswatini, Ethiopia, Gabon, Ghana, Guinea, Kenya, Madagascar, Malawi, Mali, Mozambique, Namibia, Niger, Nigeria, Rwanda, 502 ISSN 1990-6471 Senegal, Sierra Leone, Somalia, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. Figure 171. Distribution of Mops (Chaerephon) pumilus Mops (Chaerephon) pusillus (Miller, 1902) 1838. Nyctinomus pusilus: Lesson, Compléments de Buffon, ed. 2, Paris, 1: 328. Publication date: 1838. (Lapsus) 1879. Nyctinomus pumilus: Gunther. Publication date: 1879. *1902. Nyctinomus pusillus Miller, Proc. Biol. Soc. Wash., 15: 245. Publication date: 16 December 1902. Type locality: Seychelles: Aldabra Island [Goto Description]. Holotype: USNM 20991: ♀, skull and alcoholic. Collected by: Dr. William Louis Abbott. Presented/Donated by: ?: Collector Unknown. Holotype: USNM 37852: ad ♀, skull and alcoholic. Collected by: Dr. William Louis Abbott. Alcoholic catalogued: 30 June 1893, skull catalogued: 12 February 1900. Poole and Schantz (1942: 120) also mention the number as 20991/37852, and furthermore indicate "Type designated by No. 37852/20997, an error for 20991/37852". 1971. Tadarida pumila: Hayman and Hill. Publication date: 1971. 1981. Tadarida (Chaerephon) pumila: Freeman. Publication date: 1981. 2004. Chaerephon pumila: Louette. Publication date: 2004. 2004. Tadarida pumilus: Louette. Publication date: 2004. 2007. Chaerephon pusillus: Goodman and Ratrimomanarivo. (Name Combination) 2020. Mops pusillus: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon pumila: ? Chaerephon pumilus: TAXONOMY: Included in pumilus by most other authors, but its status has been revised by Goodman and Ratrimomanarivo (2007: 391) and Goodman et al. (2010a: 21). COMMON NAMES: Czech: morous seychelský. English: Seychelles Free-tailed bat. French: Tadaride des Seychelles. German: Seychellen Bulldoggfledermaus. CONSERVATION STATUS: Assessment History Global 2014: LC ver. 3.1 (2001) as a synonym of Chaerephon pumilus (Mickleburgh et al., 2014b). GENERAL DISTRIBUTION: M. pusillus was restricted to the Seychelles, but recent molecular work (Goodman et al., 2010a) indictes that the populations on the Comore Islands should assigned to M. pusillus and not M. pumilus. Louette (2004) recorded as "Tadarida pumilus" occurs on at least three of the Comore Islands (Grande Comore, Anjouan, and Mayotte. Goodman et al. (2010c: 126) also recorded "Charephon pusillus" from Moheli. Its known elevational range is 15 to 250 m on Grande Comore, 5 to 410 m on Anjouan, 5 to 10 m on Moheli, and 10 to 90 m on Mayotte (Goodman et al., 2010c: 126). African Chiroptera Report 2020 Native: Comoros (Louette, 2004; Goodman et al., 2010c: 126); Seychelles (Goodman et al., 2010c: 126). ROOST: On the Comoros, Goodman et al. (2010c: 127) did not find a single roost in a natural location, but rather all were synanthropic (see also Wilkinson et al., 2012: 160). The roosts sites were often in the attics of public buildings (schools, civic centers, and hospitals). MIGRATION: Goodman et al. (2010c: 128) interviewing a director of a primary school on Mitsamiouli mentioned that the presence of bats in the school attic is very seasonal, and as the cold season approaches the site is abanodoned. POPULATION: Structure and Density:- On Aldabra, the population might be fewer than 250 animals (Mickleburgh et al., 2008dr; IUCN, 2009, as "Chaerephon pumilus"). Trend:- Unknown. REPRODUCTION AND ONTOGENY: Goodman et al. (2010c: 126) provide data that indicates possible inter island seasonal variation in breeding in the Comoros. On 14 November 2006, all of the 10 individuals caught in a colony in central Moroni were female and all reaching term in their pregnancy with single embryos measuring 17 - 20 mm. The same colony was visited on the 28 March 2007, an additional 10 individuals were captured, all of which were females of which five had placental scars, four assumed to be subadults without placental scars, and one had a tiny embryo. Juste (unpublished data in Goodman et al. (2010c: 127)) report that the site was visited in December 2005, and all of the 20 individuals caught were female. As no males were caught on the two visits with a 5 month difference, Goodman et al. (2010c: 127) presumes that this site is a maternity colony. Between 20 November and 26 November 2006, Goodman et al. (2010c: 127) visited five different M. pusillus colonies on Anjouan and in contrast to the Moroni site, these colonies contained both males and females. At three of the sites on Anjouan, females out numbered males with a consistent sex ratio in each case of 0.5, and at one site the sex ratio was 1.0, and the fifth site it was 1.6. Goodman et al. (2010c: 127) found at all the colonies, the breeding females were reaching term, with embryos ranging in crown-rump length from 17 - 23 mm, and at one site neonates were present. Of the 25 males captured at the Anjoun 503 sites, all were adult, only seven (28 %) had scrotal testes and convoluted epididymides. The only evidence of non-synchronized breeding in M. pusillus on Anjouan was a free-flying animal captured at Patsi on 20 November with a single embryo measuring 1 mm in crown rump length. Goodman et al. (2010c: 127) report on a visit between 28 November and 2 December 2006 that two separate colonies on Moheli were visited and both male and females were captured and the sex ratios were 0.57 and 0.66. At the first colony visited 28 November, of the seven females captured, four had recently given birth and were lactating and three had single embryos measuring 24 mm in crown-rump length. At the second site visited on the 2 December, all of the females were still pregnant with single embryos and with crownrump length of 19 - 22 mm. Of the males handled from these two colonies, three of 12 (25 %) were in breeding condition with largely scrotal testes with enlarged epididymides. On Mayotte, between 27 February and 1 March 2007, two colonies were visited and both had males and females, not one of the females showed signs of active reproduction, in serveral cases the females had remaining lactation tissue and placental scars or large mammae and placental scars, indicating that they had recently terminated a breeding period, also none of the males caugth were in reproductive condition (Goodman et al., 2010c). Four different colonies were visted on Grande Comore between 29 March and 1 April 2007, and in all cases, both sexes were found in each and the sex ratio varied from 0.15 to 2.3, with one exeption none of the individuals showed active signs of reproduction, although many of the females had large mammae and placental scars. The exeption was a female captued on the 29 March in estrous (Goodman et al., 2010c: 127). PARASITES: Lagadec et al. (2012: 1696) found this species to be infected by Leptospira bacteria. Hutson (2004a: 130) mentions an endemic ectoparasitic polyctenid bug, Hypoctenes hutsoni Maa, 1970. VIRUSES: Astroviridae Hoarau et al. (2018: 1) tested 58 bats from the Comoros, found none of them to test positive for this group of viruses. Coronaviridae Joffrin et al. (2020: 5) reported Alphacoronavirus from a bat from Mayotte. an 504 ISSN 1990-6471 Paramyxoviridae Wilkinson et al. (2012: 160) tested 20 individuals from the Comoros using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Rhabdoviridae Of the 19 bats from Anjouan that were tested by Mélade et al. (2016a: 6), two showed seroreactivity to Lagos bat lyssavirus, and two of seven other bats showed the same reaction to Duvenhage lyssavirus. On Mayotte, one out of three were positive for these two viruses. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, Mayotte, Seychelles. Figure 172. Distribution of Mops (Chaerephon) pusillus Mops (Chaerephon) russatus (J.A. Allen, 1917) *1917. Chærephon russatus J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 458, text fig. 25. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Medje [02 25 N 27 18 E, 800 m] [Goto Description]. Holotype: AMNH 48925: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 8 September 1910; original number: 993. Topotype: MCZ 17386: ♀, skin and skull. Collected by: ?: Collector Unknown; collection date: 8 September 1910. Presented/Donated by: ?: Collector Unknown. Topotype: MCZ 17387: ♀, skin and skull. Collected by: ?: Collector Unknown; collection date: 8 September 1910. Presented/Donated by: ?: Collector Unknown. 2020. Mops russatus Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon russata: (Alternate Spelling) ? Chaerephon russatus: (Name Combination) ? Tadarida russata: (Name Combination) TAXONOMY: See Simmons (2005). Gregorin and Cirranello (2015: 10) suggest that "Chaerephon russatus" is the basalmost branch of the Chaerephon + Mops clade within the Molossidae. COMMON NAMES: Chinese: 赤 犬 吻 蝠 . Czech: morous guinejský. English: Russet Wrinkle-lipped bat, Russet Freetailed Bat. French: Tadaride rousse. German: Rostfarbene Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Mickleburgh et al., 2008bu; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) assessed as "Tadarida russata" (Mickleburgh et al., 2008bu; IUCN, 2009). 2004: NT ver 3.1 (2001) assessed as "Chaerephon russata" (Mickleburgh et al., 2004aj; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: It is presumably threatened by habitat loss in parts of its West African range, although further studies are needed to confirm this (Mickleburgh et al., 2008bu; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008bu) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. The species has been recorded from the Hell's Gate National Park in Kenya and Tai National Park in Côte d'Ivoire. African Chiroptera Report 2020 Further studies are needed into the distribution, natural history and threats to this species. 505 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Guinea, Kenya. GENERAL DISTRIBUTION: Mops russatus has only been recorded from a dozen or so, widely dispersed, localities. It has been recorded from the Tai National Park in Côte d'Ivoire in the west to the Hell's Gate Canyon National Park in Kenya. Native: Cameroon; Congo (The Democratic Republic of the); Côte d'Ivoire (Beaucournu and Fahr, 2003: 157); Ghana; Guinea (1st record: Decher et al., 2016: 275); Kenya. POPULATION: Structure and Density:- It is believed to be a quite rare species that occurs in small colonies (Mickleburgh et al., 2008bu; IUCN, 2009). Trend:- 2008: Unknown (Mickleburgh et al., 2008bu; IUCN, 2009). Figure 173. Distribution of Mops (Chaerephon) russatus Mops (Chaerephon) tomensis (Juste and Ibáñez, 1993) *1993. Tadarida (Chaerephon) tomensis Juste and Ibáñez, J. Mamm., 74 (4): 901, figs 1 - 2. Publication date: November 1993. Type locality: São Tomé and Principé: São Tomé Island: 3 km NW Guadalupe: Praia das Conchas [00 24 N 06 37 E] [Goto Description]. Holotype: EBD MAM 18256: ♀, complete skeleton. Collected by: Javier Juste and Antonio Ayong; collection date: 22 April 1988; original number: B7085. Paratype: EBD MAM 17371: ♀. Collected by: Javier Juste and Carlos J. Ibáñez Uargui; collection date: 11 April 1988. From Agua Izé. Paratype: EBD MAM 17734: ♀. Collected by: Javier Juste and Antonio Ayong; collection date: 22 April 1988. - Etymology: Referring to the island of São Tomé, where the species occurs. 2020. Mops tomensis: Simmons and Cirranello, BatNames.org. (Current Combination) ? Chaerephon tomensis: (Name Combination) ? Tadarida tomensis: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: morous tomášský. English: São Tomé Free-tailed Bat. French: Tadaride de São Tomé. German: São Tomé-Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Endangered (EN B1ab(iii) ver 3.1 (2001)) because its extent of occurrence is less than 1000 km2, all individuals are in a single location (the island of São Tomé), and there is continuing decline in the extent and quality of its lowland habitat (Mickleburgh et al., 2008dl; IUCN, 2009). Assessment History Global 2008: EN B1ab(iii) ver 3.1 (2001) [assessed as "Tadarida tomensis"] (Mickleburgh et al., 2008dl; IUCN, 2009). 2004: CR B1ab(I,iii) ver 3.1 (2001) (Mickleburgh et al., 2004cy; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is believed to be threatened by general habitat loss resulting from coastal development, including tourism activities, conversion of land to agricultural use and possible port construction. The species may be competing with the much more abundant species Mops pumilus (J. Juste pers. comm., 2008 in Mickleburgh et al., 2008dl in IUCN, 2009), with 506 ISSN 1990-6471 further studies into this relationship needed (Mickleburgh et al., 2008dl; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008dl) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further research is needed into the distribution, natural history and threats to this species. Trend:- 2008: Decreasing (Mickleburgh et al., 2008dl; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: São Tomé and Principé. GENERAL DISTRIBUTION: Mops tomensis has been recorded from two lowland localities, Praia das Conchas (3 km NW of Guadalupe) and Agua Izé, in the northern part of São Tomé Island, of São Tomé and Príncipe. Extensive surveys have not located the species elsewhere on the island. Native: São Tomé and Principe [São Tomé] (Rainho et al., 2010a: 40). POPULATION: Structure and Density:- It seems to be a very rare species that, despite considerable trapping effort, is currently only known from the type series of three specimens (J. Juste pers. comm., 2008 in Mickleburgh et al., 2008dl [in IUCN, 2009]). Figure 174. Distribution of Mops (Chaerephon) tomensis Subgenus Mops (Mops) Lesson, 1842 *1842. Mops Lesson, Nouv. Tabl. Regn. Anim. Mammifères, 18. Publication date: 1842. Comments: Type species: Mops indicus Lesson, 1842 (=Molossus mops de Blainville, 1840; Genotype Mops indicus Lesson = Dysopes mops F. Cuvier; listed in synonymy of Nyctinomus by Miller, 1907, Bull. U.S. Nat. Mus., 57, p. 251; but listed as a genus with distinct characters by Thomas, 1913, J. Bombay Nat. Hist. Soc., 22, p. 91). - Etymology: From the masculine German substantive Mops, meaning "pug", a small companion dog resembling a bulldog, corresponding to the French "carlin" and the Italian "carlino"; the name refers to the wrinkled lips characteristic of the genus Mops and the family Molossidae. According to one seemingly erroneous interpretation, from the Malaysian substantive mòps, meaning "bat" (see Lanza et al., 2015: 293). (Current Combination) 1917. Allomops J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 470, text-fig. 12 - 15. Publication date: 29 September 1917 [Goto Description]. - Comments: Type species: Chærephon (Allomops) osborni Allen, 1917 (=Nyctinomus condylurus A. Smith, 1833). 1951. Tadarida (Mops): Ellerman and Morrison-Scott. (Name Combination) ? Mops (Mops): (Name Combination) DISTRIBUTION BASED ON VOUCHER SPECIMENS: Benin, Chad, Kenya, South Sudan, Tanzania. Mops (Mops) condylurus (A. Smith, 1833) *1833. Nyctinomus Condylurus A. Smith, S. Afr. Quart. J., ser. 2, 1 (2): 54. Publication date: November 1833. Type locality: South Africa: KwaZulu-Natal: Port Natal [=Durban], near [29 52 S 31 00 E] [Goto Description]. - Etymology: From the masculine scientific Latin adjective made up of the masculine Greek substantive "κονδύλς" (kóndylos), meaning "knob, knuckIe" and the feminine Green substantive "ούρἀ" (urá), meaning "tail" as, according to Smith (1833: 54) the tail is "[…] enlarged and wrinkled at the point" (see Lanza et al. (2015: 298). African Chiroptera Report 2020 1864. 1870. 1911. 1917. 1917. 1936. 1939. 1939. 507 Dysopes hepaticus Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 31 (7): 14. Type locality: Sudan: Bahr-el-Ghazal province: Country of the Req natives: Djur River [Goto Description]. - Comments: Meester et al. (1986: 74): "Allen (1939a), who regards it as a synony of bivittata. Hayman and Hill (1971) regard it as unidentifiable, but point out that Kock (in litt) believes that it probably represents condylurus. The latter view is expressed also by Kock (1969a) and Koopman (1975)". Nyctinomus angolensis Peters, J. Sci. mat. phys. nat., ser. 1, 3 (10): 124. Publication date: December 1870. Type locality: Angola: Quenza river [= Quanza River] [09 20 S 13 00 E] [Goto Description]. Holotype: ZMB 4904: ad ♂, skull and alcoholic. Collected by: Toulson (Museum Bocage). Formerly Mus. Bocage; see Turni and Kock (2008: 70). Comments: Considered a synonym of leucostigma by Peterson et al. (1995: 168). Roberts (1951: 97), and Ansell and Dowsett (1988: 47) assigned it to Gunther, 1870, 1871. According to Meester et al. (1986: 74) "Günther 1871, Zoological Record (1870): 8. Quenza River [="?Cuanza River], Angola. Usually wrongly attributed to Peters, 1870, but see Cabrera and Ruxton, Ann. Mag. Nat. Hist., (9) 17: 595, 1926." According to Peterson et al. (1995)? "Kaudern, 1915: Ark. Zool., 9 (18): 77", but Peters, 1870 according to Simmons (2005). Chærephon emini Wroughton, Ann. Mag. nat. Hist., ser. 8, 8 (46): 458. Publication date: 1 October 1911. Type locality: Sudan: Rosieres [11 52 N 34 23 E]. - Comments: Not of de Winton, 1901: see Meester et al. (1986: 74). Chærephon (Allomops) osborni: J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 470. Publication date: 29 September 1917. (Name Combination, Lapsus) Mops (Allomops) osborni J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 470, 473, text-fig. 12 - 14, 26. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Kinshasa (near Leopoldville) [04 18 S 15 18 E, 350 m] [Goto Description]. Holotype: AMNH 49230: ad ♂, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 22 December 1914; original number: 2570. Paratype: AMNH 49244: ad ♀. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition. Presented/Donated by: ?: Collector Unknown. Mops angolensis wonderi Sanborn, Field Mus. Nat. Hist., Zool. Ser., (362) 20 (14): 114. Publication date: 15 August 1936. Type locality: Mali: 7 km E Bamako: Sotuba [12 36 N 07 55 W] [Goto Description]. Holotype: FMNH 42138: ad ♀, skin and skull. Collected by: Frank C. Wonder; collection date: 13 April 1934; original number: 814. - Comments: Considered a valid subspecies by Grubb et al. (1998: 93). Mops osborni occidentalis Monard, Arq. Mus. Bocage, 10: 78. Publication date: March 1939. Type locality: Guinea-Bissau: Mansoa [12 04 N 15 19 W] [Goto Description]. Syntype: NMBA 5268: ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 11-12 December 1937; original number: 226. See Oakeley (1998: 2). From Mansoa (see Monard, 1939: 78). Syntype: NMBA 5269: ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 11-12 December 1937; original number: 275. See Oakeley (1998: 2). From Mansoa (see Monard, 1939: 78). Syntype: RMCA 15307: ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 1937; original number: 223. From Mansoa (see Monard, 1939: 78). Syntype: RMCA 15308: ♀, alcoholic (skull not removed). Collected by: Dr. Albert Monard; original number: 498. From Contubo-el (see Monard, 1939: 78). - Comments: The type locality is either Mansoa [12 04 N 15 19 W], Contubo-el [12 22 N 14 34 W] or Pitche [12 20 N 13 57 W]. Monard (1939: 78) does not mention a holotype, but a series of specimens from Mansoa (14), Cotubo-el (5) and Pitche (9). Mops osborni occidentalis f. fulva Monard, Arq. Mus. Bocage, 10: 80. Publication date: March 1939. Type locality: Guinea-Bissau: Mansoa [12 04 N 15 19 W] [Goto Description]. Syntype: NMBA 5270: ♀, alcoholic (skull not removed). Collected by: Dr. Albert Monard; collection date: 11-12 December 1937; original number: 232. Monard (1939: 80) mentions a series of 10 specimens (nrs 228 to 237, 5 MM + 5 FF) from Mansoa and 1 M (nr 781) from Pitche. Syntype: RMCA 15309: ♂, alcoholic (skull not removed). Collected by: Dr. Albert Monard; original number: 229. Monard (1939: 78) mentions a larger series: nrs 215 to 227, 275 (13 MM + 1 F from Mansoa); 494 to 498 (1 M + 4 FF) from Contubo-el; 77. - Comments: The type locality is either Mansoa [12 04 N 15 19 W] or Pitche [12 20 N 13 57 W]. Grubb et al. (1998: 93) indicate that Monard's quadrinominal name was validated, in accordance with International Code of Zoological Nomenclature 508 ISSN 1990-6471 1942. 1962. 1979. 1981. 2018. ? ? ? ? ? ? ? ? ? ? ? ? ? Articke 45 (g)(ii)(1), by its use as a trinomial (Freeman (1981b: 159). Monard (1939: 80) mentions a series of 10 specimens (nrs 228 to 237, 5 ♂♂ + 5 ♀♀) from Mansoa and 1 ♂ (nr 781) from Pitche. Mops angolensis orientis G.M. Allen and Loveridge, Bull. Mus. comp. Zool., 89 (4): 151, 166. Publication date: February 1942. Type locality: Tanzania: SE Tanzania, Mtwara, Rovuma river: Kitaya [300 ft] [Goto Description]. Holotype: MCZ 38829: ad ♂, skin and skull. Collected by: Professor Arthur Loveridge; collection date: 3 April 1939. See Allen and Loveridge (1942: 166). Helgen and McFadden (2001: 146) also mention 9 paratypes (MCZ 38826 - 38828, 38830 - 38835 (4 FF+ 5 MM, skin and skull): see Helgen and McFadden (2001: 145). Paratype: MCZ 38826: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38827: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38828: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38830: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38831: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38832: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38833: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38834: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. Paratype: MCZ 38835: Collected by: Professor Arthur Loveridge. Presented/Donated by: ?: Collector Unknown. - Comments: Possibly valid as a subspecies. Tadarida angolensis: Taufflieb, Bull. Inst. Rech. Sci. Congo, 1: 112. (Name Combination) Tadarida condylura: Ansell, The Puku, 3: 13. (Name Combination) Mops osborni fulva: Freeman, Field. Zool., n.s. 7: 159. (Name Combination) Mops orientis: Gunnell and Manthi, J. Hum. Evol., Suppl.. Publication date: 6 April 2018. (Name Combination) Mops (Mops) condylurus: (Name Combination, Current Combination) Mops angolensis: Mops condylura angolensis: (Name Combination, Alternate Spelling) Mops condylura condylura: (Name Combination, Alternate Spelling) Mops condylura orientis: (Name Combination, Alternate Spelling) Mops condylura osborni: (Name Combination, Alternate Spelling) Mops condylurus orientis: (Name Combination) Mops condylurus osborni: (Name Combination) Mops condylurus wonderi: (Name Combination) Mops condylurus: (Name Combination) Tadarida (Mops) condylura: (Name Combination) Tadarida condylura condylura: (Name Combination) Tadarida condylura osborni: (Name Combination) TAXONOMY: In their analyses, Ammerman et al. (2012: 20) found that Mops condylurus from clusters with M. leucostigma from Madagascar when present, and these two form a close relationship with African Chaerephon and not other Mops. They also indicate that the two species separated from one another less than 2 million years ago (Ammerman et al., 2012: 23). Figure 175. An adult female Mops condylurus caught at Mlawula-NR, Swaziland. Includes angolensis and leucostigma, but see Peterson et al. (1995: 168). COMMON NAMES: Afrikaans: Angola-losstertvlermuis, Angolase Losstervlermuis. Chinese: 安 哥 拉 犬 吻 蝠 . Czech: morous angolský. English: Angolan Mops Bat, Knob-tailed Mops Bat, Angolan Free-tailed Bat, Knob-tailed Nyctinome. French: Tadaride d'Angola, Molosse d'Angola. German: AngolaBulldoggfledermaus, Centralafrikanische African Chiroptera Report 2020 Doggengrämler. Italian: Carlìno Portuguese: Morcego de cauda livre. 509 condilùro. Africa and Swaziland. The species appears to be largely absent from the Congo Basin. ETYMOLOGY OF COMMON NAME: Originally described from the former Natal province, it might more appropriately have been called the Natal free-tailed bat, but this name is taken by Mormopterus acetabulosus. Monadjem et al. (2010d) does not record any specimens occuring in Namibia. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Gunnell et al. (2015a: 20) report on a right distal humerus from a Pleistocene deposit at Olduvai Gorge (Tanzania), which they assigned to Mops cf. M. condylurus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001) ) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008af; IUCN, 2009; Monadjem et al., 2017ab). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops condylurus] (Monadjem et al., 2017ab). 2008: LC ver 3.1 (2001) [assessed as Tadarida condylura] (Mickleburgh et al., 2008af; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004aq; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016b). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: There are no major threats to this species. Some colonies roosting within buildings are possibly threatened by general persecution (Mickleburgh et al., 2008af; IUCN, 2009; Monadjem et al., 2017ab). CONSERVATION ACTIONS: Mickleburgh et al. (2008af) [in IUCN (2009)] and Monadjem et al. (2017ab) report that this species is present in a number of protected areas. No direct conservation actions are currently needed for the species as a whole. GENERAL DISTRIBUTION: Mops condylurus is widely distributed over much of sub-Saharan Africa. It ranges from Senegal, the Gambia and Mali in the west, to the Sudan, Ethiopia and Somalia in the east; from here it ranges southwards through much of eastern and southern Africa, as far south as eastern South Besides Mops condylurus, Taylor et al. (2018b: 63) also mention Mops cf. condylurus from Angola, whose echolocation calls do not match any known species. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,057,626 km 2. In the RSA, Babiker Salata (2012: 49) found that its distribution is stongly associated with the mean temperature of the coldest quarter. Native: Angola (Lopes and Crawford-Cabral, 1992; Monadjem et al., 2010d: 543; Taylor et al., 2018b: 62); Benin (Capo-Chichi et al., 2004: 163); Botswana (Monadjem et al., 2010d: 543); Burkina Faso (Kangoyé et al. (2012: 6025; 2015a: 618); Burundi; Cameroon (Bakwo Fils et al., 2014: 4); Congo (Bates et al., 2013: 336); Congo (The Democratic Republic of the) (Allen, 1917; Hayman et al., 1966; Dowsett et al., 1991: 259; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 543); Côte d'Ivoire; Ethiopia (Lavrenchenko et al., 2004b: 141); Gambia (Emms and Barnett, 2005: 50); Ghana; Guinea (Fahr et al., 2006b: 245; Decher et al., 2016: 275); Guinea-Bissau (Seabra, 1900a; Monard, 1939; Veiga-Ferreira, 1949; Lopes and Crawford-Cabral, 1992; Rainho and Ranco, 2001: 75); Kenya; Liberia (Fahr, 2007a: 104); Malawi (Happold et al., 1988; Happold and Happold, 1997b: 823; Monadjem et al., 2010d: 543); Mali; Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 543; Monadjem et al., 2010c: 382); Namibia; Niger; Nigeria; Rwanda; Senegal; Sierra Leone; Somalia; South Africa (De Graaf, 1960; Monadjem et al., 2010d: 544); Sudan; Swaziland (Monadjem et al., 2010d: 544); Tanzania; Togo; Uganda (Kityo and Kerbis, 1996: 62); Zambia (Monadjem et al., 2010d: 544); Zimbabwe (Cotterill, 2004a: 260; Monadjem et al., 2010d: 3422). Presence uncertain: Madagascar. DENTAL FORMULA: Dorst (1949: 47-48, fig 2) discussed the milk dentition of this species (as Mops angolensis) and indicated that the incisors are quite different from both the canines and premolars, as the prior are tricuspid and the latter are peg-like. DETAILED MORPHOLOGY: Baculum - Unknown 510 ISSN 1990-6471 Brain - Amrein et al. (2007) from specimens collected from Bénin reported sparse proliferating cell activity, detected with Ki-67 and MCM2, in the subgranular layer of the dentate gyrus, with doublecortin (DCX) they detected young migrating neuron in the subgrannular layer of the hippocampus. Amrein et al. (2007) also reported moderate to ample proliferating cells (Ki-67 positive cells) and migrating young neurons (DCX positive cells) in the rostral migratory stream. No NeuroD was detected in the hippocampal granule cells (Amrein et al., 2007). FUNCTIONAL MORPHOLOGY: Maloney et al. (1999: 385) indicate that M. condylurus is well adapted to tolerated high ambivalent temperatures, which may enable them to exploit thermally challenging roost sites and allow them to survive in habitats where less stressful roosts are limited. Buffenstein et al. (1999: 15) found that urine voided prior to feeding in summer was 22% more concentrated than that in autumn, suggesting that periods of exposure to high temperatures were more stressful. The urine produced was up to eight times more concentrated than plasma. SEXUAL DIMORPHISM: Chaverri et al. (2018: 1944) [referring to Bouchard, 2001b] indicates that the sexes can be distinguished by scent produced from the interaural and muzzle glandular areas. ECHOLOCATION: 5.Bohmann et al. (2011: 1) [referring to Monadjem et al. (2010d)] mention peak frequencies between 26 - 35 kHz. Schoeman and Waddington (2011: 291) mention a peak frequency of 24.8 ± 1.5 kHz and a duration of 15.7 ± 2.1 msec for specimens from Durban, South Africa. From the same area, Naidoo et al. (2011: 23) mention a Fpeak of 25.8 ± 0.5 kHz and a duration of 6.8 ± 1.8 msec. In Swaziland, Taylor et al. (2013b: 18) recorded the following parameters for 6 calls: Fmax: 34.8 ± 2.78 (31.5 - 38.6) kHz, Fmin: 25.8 ± 1.79 (23.9 28.2) kHz, Fknee: 30.2 ± 2.16 (27.4 - 32.4) kHz, Fchar: 27.3 ± 1.98 (24.9 - 29.3) kHz, duration: 5.5 ± 1.40 (3.4 - 7.2) msec. Additional values reported by Monadjem et al. (2017c: 179) are: Fmin: 25.1 ± 1.68 (23.5 - 26.1) kHz, Fknee: 30.1 ± 2.03 (28.1 32.1) kHz, Fc: 27.3 ± 1.56 (25.9 - 28.8) kHz and duration: 4.3 ± 2.08 (2.2 - 6.4) msec. These calls could be detected up to 30 m away. At Farm Welgevonden, South Africa, Taylor et al. (2013a: 556) report a Fknee value of 30 (27 - 32) kHz. Linden et al. (2014: 40) reported the following parameters from the Soutpansberg range (RSA): Fmin: 24 - 28 kHz, Fchar: 25 - 29 kHz, Fknee: 27 - 32 kHz, Slope: 40 - 160 OPS, duration: 3 - 7 msec. From the Mapungubwe National Park (RSA), Parker and Bernard (2018: 57) recorded 6 calls with: Fchar: 29.12 ± 0.70 kHz, Fmax: 30.72 ± 1.52 kHz, Fmin: 28.45 ± 0.50 kHz, Fknee: 29.82 ± 0.97, duration: 15.44 ± 3.78 msec, at 5.57 ± 1.11 calls/sec. Five calls from the Okavango River Basin reported by Weier et al. (2020: Suppl.) had the following characteristics: Fmax: 30.74 ± 2.64 kHz, Fmin: 27.88 ± 4.69 kHz, Fknee: 30.12 ± 1.92 kHz, Fchar: 29.00 ± 1.49 kHz, slope: 11.60 ± 12.27 Sc, duration: 7.09 ± 3.46 msec. From Maroua, Cameroon, Manga Mongombe (2012: 79) and Bakwo Fils et al. (2018: 4) reported for 12 calls a call type of FM and the following parameters: Fmax: 37.8 ± 3.8 (33.9 - 41.6) kHz, Fmin: 34.2 ± 4.9 (30.1 - 40.0) kHz, Fmean: 36.4 ± 4.5 (31.0 - 41.0) kHz, Fknee: 38.2 ± 4.7 (33.9 - 41.5) kHZ, Fchar: 33.67 ± 5.1 (30.1 - 40.0) kHz, and duration: 0.78 ± 0.89 (0.57 - 0.88) msec. Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 26.7 kHz, Fstart: NA kHz, Fend: NA kHz, Band width: NA kHz, and duration: 10 msec. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Ðulic and Mutere (1973a) described the karyotype as having a diploid number of 2n = 48, aFN = 56, BA = 10, X = SM, Y = A. While Smith et al. (1986) found that the fundamental number was aFN = 66, BA = 20, this is supported by Rautenbach et al. (1993). Koubínová (2013: 7) mentioned: 2n = 48 and FN = 54. Protein / allozyme - Unknown. HABITAT: Noer et al. (2012: 1) indicate that M. condylurus prefers to forage over sugarcane fields rather than over savanna, riparian forest and urban areas. They also indicated that the bats travel up to 4.8 km from their roosts during their foraging trips. Their activity area ranges from 1,190 to 1,437 ha. HABITS: Bouchard (2001b: 109) found that, in scent-choiseexperiments, both males and females were able to distinguish the gender of conspecifics as well as the difference between roostmates and strangers. Chaverri et al. (2018: 1944) [referring to Ravel et al., 2010] further indicate that there is an inverse relation between the number of individuals that can be recognized based on olfactory cues and the African Chiroptera Report 2020 511 size of the colony. This would suggest that recognition is based on individual signatures rather than on group signatures. Trend:- 2016: Unknown (Monadjem et al., 2017ab). 2008: Unknown (Mickleburgh et al., 2008af; IUCN, 2009). ROOST: In the Mount Nimba area, Monadjem et al. (2016y: 368) found a roost in the roof of a school at 540 m. REPRODUCTION AND ONTOGENY: Mutere (1968, 1973b: 271) reported that "T. condylura" has a bimodal breeding rhythm in Uganda (pregnancies during December-March and June-September and births in March and September), which is correlated with the rainfall pattern (April-May and October-November), and that the gestation period for this species is about two months. He also indicated [in Krutzsch (2000: 95)] that the testicular position (intraabdominal or scrotal) showed no significant seasonal variation in sperm production. Bronner et al. (1999: 1) investigated the roost microclimates in three man-made structures in South Africa. The ambient temperature in these varied from below 10°C to over 40°C, but the bats preferred zones where the temperature was between 35 and 40°C, where they could maintain their basal metabolic rate. DIET: Based on DNA analysis of 30 fecal pellets collected between April and May (austral summer) in the Simunye region of Swaziland, Bohmann et al. (2011) reported insects from 24 families (belonging to seven orders) being taken as prey. The highest frequency of occurrence was for Lepidoptera (47.6 %), Diptera (26.7 %), and Hemiptera (20 %). Within the Lepidoptera, Noctuidae (Owlet moths - 23.3 %), Nymphalidae (Brush-footed butterflies - 17.6 %) and Geometridae (Geometer moths - 10 %) were found the most. Within the Diptera, it were primarly representatives of Drosophilidae (Pomace flies 6.5 %) and Muscidae (House flies and alike - 6.5 %). Within the Hemiptera, Lygaeidae (Chinch bugs and seed bugs) and Pentatomidae (Stink bugs) were found in 6.7 % of the pellets. From Maroua (Cameroon), and based on fecal analyses, Bol A Anong (2013: 32) reported the diet to include Coleoptera (63.49%), Hemiptera (27.64%), Diptera (7.42%), Lepidoptera (0.05%) and Hymenoptera (0.40%). PREDATORS: Mikula et al. (2016: Supplemental data) mention "Tadarida condylura" to be preyed upon by: African goshawk (Accipiter tachiro (Daudin, 1800)), Eurasian hobby (Falco subbuteo Linnaeus, 1758), Wahlberg's eagle (Hieraaetus wahlbergi (Sundevall, 1851)), Bat hawk (Macheiramphus alcinus Bonaparte, 1850). Bronner et al. (1999: 8) reported on the presence of an Eastern Tiger Snake (Telescopus semiannulatus semiannulatus Smith, 1849) in a colony of M. condylurus. POPULATION: Structure and Density:- This is a common species (Mickleburgh et al., 2008af; IUCN, 2009; Monadjem et al., 2017ab). Happold and Happold (1990b: 568) reported that in southern Malawi, births occurred early in the wet season and at the end of the wet season and that gestation takes about 1.5 months. Births in midMalawi occur earlier and in nothern Malawi later. Happold and Happold (1989b: 133) indicate that "Tadarida condylura" is monotocous in Malawi, where it has two births per year. They also found that the interval between births decreases with increasing latitude. Mutere (1973b) mentioned that the pattern of reproduction is not followed by a postpartum oestrous, whereas Happold and Happold (1989b) mention the opposite. Wyant and Adams (2007) present data on prenatal growth, development, and skeletal ossification, by intergrating data from changes in body size, wing area, and skeletogenesis, from specimens collected from east African countries. Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] mentioned births between early September and early May. PARASITES: Lima et al. (2012: 858) described a new species of Trypanosoma from Mozambican Mops condylurus: T. (Schizotrypanum) erneyi. This parasite was also found in two unidentified Tadarida specimens. BACTERIA: Höhne et al. (1975: 505) found Listeria monocytogenes bacteria in two out of 234 "Tadarida condylura" specimens from Lomé, Togo. Studying faecal samples from freshly captured bats, Edenborough et al. (2020: 5) found bacteria 512 ISSN 1990-6471 belonging to Proteobacteria, Bacilli, Firmicutes, Gammaproteobacteria, and Actinobacteria. HEMIPTERA Cimicidae: Loxaspis miranda Rothschild, 1912 were found in the hollow trunk of a Borassus palm containing many dead T. condylura, at the Fambani river, near mouth of Zambesi, in Mozambique (Haeselbarth et al., 1966: 10). SIPHONAPTERA Ischnopsyllidae: Lagaropsylla idea Smit, 1957 widespread in tropical Africa (Haeselbarth et al., 1966: 1990; Beaucournu and Kock, 1996). Lagaropsylla leleupi Smit, 1957 from Bugesera, Burindi (Haeselbarth et al., 1966: 191). Lagaropsylla obliqua Smit, 1957 from Sierra Leone, Cameroon and the Congo (Haeselbarth et al., 1966: 191). Lagaropsylla incerta (Rothschild, 1900) was reported from Madagascar by Beaucournu and Kock (1996). ACARI Laelapidae: Taufflieb (1962: 112) reported Chelanyssus aethiopious Hirst, 1921 from a "Tadarida angolensis [=Mops osborni]". Sarcoptidae: Notoedres tadaridae Fain, 1959 was found by Fain and Aellen (1994: 64) on a "Tadarida condylura" specimen from Tandara, DRC. Trombiculidae: Stekolnikov (2018a: 69, 172) reported Holubicula toroensis Daniel and Vercammen-Grandjean, 1985 on a bat from Uganda, and Microtrombicula tadarida Vercammen-Grandjean, 1965 on a bat from Rwanda. VIRUSES: Astroviridae Viruses belonging to this family were recorded by Waruhiu et al. (2017) from these bats in Kenya, and in one (out of 52) tested by Hoarau et al. (2018: 2) in Mozambique. Bunyaviridae Sudi et al. (2018: 3) refer to Krüger et al. (2015), who reported the presence of Lompole virus (LMPV) in Mops condylurus from the DRC (although the latter authors didn't identify the bats involved). Caliciviridae Viruses belonging to this family were recorded by Waruhiu et al. (2017) from these bats in Kenya. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in 19 individuals from Limpopo Province, South Africa, three were tested positive (3/19, 15.8 %) and 96 individuals from Mpumalanga Province, South Africa, 11 tested positive (11/96, 11.9 %) (14/115, 12.2 %). Anthony et al. (2017b: Suppl.) mention the alphacoronavirus Charephon_CoV_KY22. Maganga et al. (2020: 5) reported Alphacoronaviruses found in bats from Cameroon, Mozambique and South Africa. Filoviridae - Filo viruses Ebolavirus Muyembe-Tamfum et al. (2012: 9) refer to Pourrut et al. (2005) who found ZEBOV-specific antibodies in this species. Saéz et al. (2014) indicate that the two-year old boy, who was patient zero for the 2014 Ebola outbreak in West Africa may have been infected while playing in a hollow tree housing a colony of M. condylurus. However, Wood et al. (2015: 910) find "the suggestion that Mops condylurus was the source of infection in the index human case in Guinea noteworthy if somewhat speculative." Goldstein et al. (2018: 1084) described a new Ebolavirus from three Sierra Leonan C. pumilus and one Mops condylurus: Bomali virus (BOMV). They were sampled in human dwellings in small villages in the Bomali district. This virus might infect humans, but it is not known (yet) whether it is virulent in humans too. Forbes et al. (2019: 955) reported this virus from one M.condylurus from Kenya (out of 16 M. condylurus + 92 other bats collected), and Karan et al. (2019: 1774) reported it from three our of 109 . condylurus bats from Guinea. Flaviviridae Flavivirus de Jong et al. (2011: 10) reported Bukalasa bat virus, Dakar bat virus, Entebbe bat virus. The latter two are also reported by Luis et al. (2013: suppl.), Blitvich and Firth (2017: 3). In Uganda, Kading et al. (2018: 3) found neutralizing antibodies against Dengue 2 virus (DENV-2) and non-specific Flaviviruses. Dakar virus - Kemp (1975: 616) mentions that this virus was isolated in Nigeria on M. condylurus for the first time on 25 January 1966. Zika virus (ZIKV) - Malmlov et al. (2019: 2) refer to Shepherd and Williams (1964) who examined 36 Ugandan condylurus specimens of which 26 had antibodies against ZIKV. Pegivirus (BPgV) 1 of 10 specimens from the Democratic Republic of Congo examined by Quan et al. (2013: Table S5) (10.0 %) was infected by clade G type African Chiroptera Report 2020 Pegivirus. Of the same 10 specimens, 2 were infected by clade K type Paramxoviridae Viruses belonging to this family were recorded by Waruhiu et al. (2017) from these bats in Kenya. 513 Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Madagascar, Malawi, Mali, Mauritania, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, South Sudan, Sudan, Tanzania, The Gambia, Togo, Uganda, Zambia, Zimbabwe. Peribunyaviridae Orthobunyavirus Fagre and Kading (2019: 4) report that Bunyamwera virus (BUNV) was found either by virus isolation or molecular/serologic evidence. In their overview tables, Maganga et al. (2014a: 8) and Willoughby et al. (2017: Suppl.) reported that the following viruses were already found on M. condylurus: Bukalasa bat virus, Dakar bat virus, Entebbe bat virus, Coronavirus (SARS-CoV), Bunyamwera orthobunyavirus, West Nile virus, Yellow fever virus, Zaïre Ebola, Zika virus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Botswana, Burundi, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Eswatini, Ethiopia, Gabon, Figure 176. Distribution of Mops (Mops) condylurus Mops (Mops) congicus J.A. Allen, 1917 *1917. Mops congicus J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 467, pl. 55. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Medje [02 25 N 27 18 E, 800 m] [Goto Description]. Holotype: AMNH 48893: ad ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 8 September 1910; original number: 966. ? Mops (Mops) congicus: (Name Combination, Current Combination) ? Mops congica: (Alternate Spelling) ? Tadarida congica: (Name Combination) TAXONOMY: Does not include trevori; see Peterson (1972). COMMON NAMES: Czech: morous konžský. English: Medje Mops Bat, Medje Free-tailed Bat, Medje Greater Freetailed Bat. French: Tadaride de Medje. German: Kongo-Bulldoggfledermaus, MedjeBulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bp; IUCN, 2009; Monadjem et al., 2017ay). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops congicus] (Monadjem et al., 2017ay). 2008: LC ver 3.1 (2001) [assessed as Tadarida congica] (Mickleburgh et al., 2008bp; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004bx; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: It is threatened in parts of its range by habitat loss, largely resulting from logging activities and conversion of land to agricultural use (Mickleburgh et al., 2008bp; IUCN, 2009; Monadjem et al., 2017ay). CONSERVATION ACTIONS: Mickleburgh et al. (2008bp) [in IUCN (2009)] and Monadjem et al. (2017ay) report that it is not 514 ISSN 1990-6471 known if the species is present in any protected areas. Further studies are needed into the distribution, abundance, natural history, and threats to this poorly known species. GENERAL DISTRIBUTION: Mops congicus has been patchily recorded in Central Africa. It is found from southern Cameroon, east into the northern part of the Democratic Republic of the Congo and western Uganda. Trend:- 2016: Unknown (Monadjem et al., 2017ay). 2008: Unknown (Mickleburgh et al., 2008bp; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Central African Republic, Congo (Democratic Republic of the), Uganda. Earlier records of this species from Ghana and Nigeria are now allocated to Mops trevori (Simmons, 2005). Native: Cameroon; Congo (The Democratic Republic of the); Uganda. POPULATION: Structure and Density:- The species has been rarely collected. While a single large colony of approximately several dozen bats has been reported, it is suspected to generally occur in much smaller roosts (Mickleburgh et al., 2008bp; IUCN, 2009; Monadjem et al., 2017ay). Figure 177. Distribution of Mops (Mops) congicus Mops (Mops) demonstrator (Thomas, 1903) *1903. Nyctinomus demonstrator Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (71): 504. Publication date: 1 November 1903. Type locality: Sudan: Equatoria province: 25 mi (37 km) N of Gondokoro: Mongalla [05 10 N 31 50 E, 500 m] [Goto Description]. Holotype: BMNH 1902.7.4.3: ad ♂. Collected by: W.L.S. Loat Esq. Presented/Donated by: W.L.S. Loat Esq. 1917. Mops (Allomops) faradjius J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 476. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Faradje [03 44 N 29 43 E] [Goto Description]. Holotype: AMNH 49222: ad ♂, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 1 November 1913; original number: 3015. - Comments: One of the skull labels mentions 12 January 1913, which probably is an error. ? Mops (Mops) demonstrator: (Name Combination, Current Combination) ? Mops demonstrator: (Name Combination) ? Mops demostrator: (Lapsus) ? Tadarida (Mops) demonstrator: (Name Combination) ? Tadarida demonstrator: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: morous mongallanský. English: Mongalla Mops Bat, Mongallan Mops Bat, Mongalla Freetailed Bat. French: Tadaride de Mongalla. German: Mongalla-Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bs; IUCN, 2009; Monadjem et al., 2017u). African Chiroptera Report 2020 Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops demonstrator] (Monadjem et al., 2017u). 2008: LC ver 3.1 (2001) [assessed as Tadarida demonstrator] (Mickleburgh et al., 2008bs; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004az; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: In part of its range this species is threatened by habitat loss, mostly the cutting of larger roosting trees from savanna. It may also be threatened by encroaching desertification in the northern limits of its range (Mickleburgh et al., 2008bs; IUCN, 2009; Monadjem et al., 2017u). CONSERVATION ACTIONS: Mickleburgh et al. (2008bs) [in IUCN (2009)] and Monadjem et al. (2017u) report that this species has been recorded from the Garamba National Park in the Democratic Republic of the Congo (Freeman, 1981b), and may be present in additional protected areas. There is a need to protect suitable large roosting trees in savanna. Further research is needed to better understand the distribution, natural history and major threats to this species. 515 MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) reported 2n = 48, FN = 54, BA = 8, a submetacentric X chromosome and an acrocentric Y chromosome. Protein / allozyme - Unknown. PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- This species is rarely recorded. Animals are usually found singly or as small groups of up to 12 bats (Mickleburgh et al., 2008bs; IUCN, 2009; Monadjem et al., 2017u). Trend:- 2016: Decreasing (Monadjem et al., 2017u). 2008: Decreasing (Mickleburgh et al., 2008bs; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burkina Faso, Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Kenya, Malawi, Rwanda, South Sudan, Sudan, Uganda. GENERAL DISTRIBUTION: Mops demonstrator has been patchily recorded from West Africa (Côte d'Ivoire, Ghana and Burkina Faso) and Central to East Africa (Cameroon, eastern Democratic Republic of the Congo, western Uganda and Sudan). A record from the Gambia appears to be in error (Koopman, 1989b; Grubb et al., 1998). Native: Burkina Faso (Kangoyé et al., 2015a: 618); Cameroon; Congo (The Democratic Republic of the); Côte d'Ivoire; Ghana; Sudan; Uganda. Figure 178. Distribution of Mops (Mops) demonstrator Mops (Mops) leucostigma (G.M. Allen, 1918) *1918. Chaerephon leucostigma G.M. Allen, Bull. Mus. comp. Zool., 61 (14): 513. Publication date: 27 February 1918. Type locality: Madagascar: Tananarive [=Antananarivo] [18 55 S 47 31 E, 1 300 m]. Holotype: MCZ 16344: ad ♀, skin and skull. Collected by: Dr. Frederik Roelker Wulsin; collection date: December 1915. See Peterson et al. (1995: 168); Helgen and McFadden (2001: 145). Paratype: MCZ 16345: ♂, skin and skull. See Helgen and McFadden (2001: 145). - Comments: Ratrimomanarivo et al. (2008: 66 - 67) mentions the following: "In his description of Mops leucostigma, Allen (1918) cited two specimens - the holotype (MCZ 16344), which was sexed as a 'female' and the paratype (MCZ 16345), which was sexed as a 'male' (also see Helgen & McFadden, 2001). We examined these two specimens. In Allen’s description he specifically states, 'The skull is 516 ISSN 1990-6471 2018. ? ? ? ? ? provided with a prominent sagittal crest, which in the type, though partly broken, must have been at least a millimetre high.' This is an accurate description of the MCZ 16344. In members of the genus Mops in general (Freeman, 1981) and in M. leucostigma specifically (this study), adult males are distinctly larger than adult females and have a prominent and more developed occipital and sagittal crests. Based on size, skull shape, and the development of the occipital and sagittal crests, the holotype is a male and the paratype a female. Given that all of the collection data for the holotype (MCZ 16344) and paratype (MCZ 16345) are identical, we strongly suspect that the museum labels for these two specimens have been accidentally exchanged before the publication of Allen (1918).". Mops leucogaster: Kemp, López-Baucells, Rocha, Wangensteen, Andriatafika, Nair and Cabeza, Agric. Ecosyst. Environ., 269: 88 (for 2019). Publication date: 1 October 2018. (Lapsus) Mops (Mops) leucostigma: (Name Combination, Current Combination) Mops condylura leucostigma: (Name Combination, Alternate Spelling) Mops leucostigma: (Name Combination) Tadarida (Mops) leucostigma: (Name Combination) Tadarida leucostigma: (Name Combination) TAXONOMY: Included in condylurus by Koopman (1993a: 236), but considered a valid species in Tadarida (M.) by Peterson et al. (1995: 152) and Russ et al. (2001). Considered a valid species in the genus Mops by Hutson et al. (2001: 34), Goodman and Cardiff (2004: 227), Simmons (2005). COMMON NAMES: Czech: morous běloskvrnný. English: Madagascar White-bellied Free-tailed Bat, Malagasy White-bellied Free-tailed Bat, Whiteshouldered bat. French: Tadaride à queue libre de Madagascar. German: Gefleckte Bulldoggfledermaus. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Gunnell et al. (2014: 3) refer to fossils material from the Andrahomana cave on Madagascar. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its large distribution range, and its ability both to tolerate persecution and to persist in a variety of habitats in many parts of its range (Andriafidison et al., 2008m; IUCN, 2009; Monadjem et al., 2017ca). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops leucostigma] (Monadjem et al., 2017ca). 2008: LC ver 3.1 (2001) [assessed as Tadarida leucostigma] (Andriafidison et al., 2008m; IUCN, 2009). 2000: DD ver 2.3 (1994). Regional None known. MAJOR THREATS: In some areas it is hunted for food (Goodman et al., 2008d) or persecuted, but these are not thought to be serious threats (Andriafidison et al., 2008m; IUCN, 2009; Monadjem et al., 2017ca). CONSERVATION ACTIONS: Andriafidison et al. (2008m) [in IUCN (2009)] report that this species is known from Kirindy CFPF and Parc National de Zombitse-Vohibasia (Goodman et al., 2005a). It is also common in villages adjacent to several protected areas (Goodman et al., 2005a; Russ et al., 2003; Monadjem et al., 2017ca). GENERAL DISTRIBUTION: Mops leucostigma is known from Madagascar and the Comoros archipelago (Mohéli and Anjouan islands). On Madagascar it is found from the north to the southwest of the island (Eger and Mitchell, 2003; Peterson et al., 1995) including the islands of Nosy Be and Nosy Komba (Rakotonandrasana and Goodman, 2007: 6). Recorded at an elevation span of 5 to 1,300 m above sea level (Allen, 1918b; Eger and Mitchell, 2003). Native: Madagascar (Eger and Mitchell, 2003; Peterson et al., 1995; Rakotonandrasana and Goodman, 2007: 6); Comoros (Ratrimomanarivo et al., 2008: 57). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 62) describe the baculum as displaying a distinct "m"shape; length: 0.75 ± 0.097 (0.56 - 0.86) mm, width: 1.09 ± 0.137 (0.83 - 1.35) mm. African Chiroptera Report 2020 HABITAT: Dammhahn and Goodman (2013: 108) indicate that this species' foraging habitat consists of open areas and the area above the forest canopy. MIGRATION: Reher et al. (2019: 121) found that M. leucostigma was only present in the Vintany Cave (southwestern Madagascar) during the rainy season, whereas other species such as Triaenops menamena and Miniopterus mahafaliensis were present in both the dry and wet season. DIET: Rakotondramanana et al. (2015: 78) report the following insect orders (by volume percent, by frequency percent): Coleoptera (64.0, 100.0); Lepidoptera (30.0, 100.0); Trichoptera (6.0, 60.0). Kemp et al. (2018: Suppl.) used DNA metacarcoding to detect insect pest species in the diet of these bats and found the following prey orders (in descending order): Coleoptera, Diptera (Simulium lineatum (Meigen, 1804)), Lepidoptera (Anthozela sp., Celsumaria elongata Brown, 2017, Spodoptera mauritia (Boisduval, 1833)), Blattodea, Hemiptera (Yanga guttulata (Signoret, 1860)), Ephemeroptera, Sarcoptiformes, Orthoptera, Hymenoptera, Mesostigmata, Trombidiformes, Trichoptera. 517 Trend:- 2016: Unknown (Monadjem et al., 2017ca). 2008: Unknown (Andriafidison et al., 2008m; IUCN, 2009). VIRUSES: Paramyxoviridae Specimens from this species tested by Wilkinson et al. (2014: 8270) were found to be positive for paramyxoviruses. 11 out of 68 Madagascan specimens tested by Mélade et al. (2016b: 4) were positive for paramyxoviruses. Rhabdoviridae Mélade et al. (2016a: 6) tested 20 bats for Lagos bat lyssavirus and found positive reactions in two of them. UTILISATION: See Goodman et al. (2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, Madagascar. PREDATORS: Goodman et al. (2015c: 78) found the remains of two individuals in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. POPULATION: Structure and Density:- The population and local abundance of this species are not known (Andriafidison et al., 2008m; IUCN, 2009; Monadjem et al., 2017ca). Figure 179. Distribution of Mops (Mops) leucostigma Mops (Mops) midas (Sundevall, 1843) *1843. Dysopes midasSundevall, Kongl. Svenska Vet.-Akad. Hand., ser. 3, 30: 207, pl. 2, fig. 7 [for 1842]. Publication date: 1843. Type locality: Sudan: Blue Nile Province: E-bank White Nile: Valid Sohey [= Wad Shala]: restricted by Turni and Kock (2008: 68). [14 34 N 13 48 E]. Syntype: BMNH 1846.6.2.20: ad ♂, skin and skull. Formerly Stockholm Museum (ZSM no. 523), locality: Bahr el Abiad. See Turni and Kock (2008: 68). Syntype: ZMB 535: skin only. Collected by: Hedenborg; collection date: 1831 - 1835. From [Dar] Sennar, Sudan; see Turni and Kock (2008: 68). Syntype: ZSM. Collected by: Hedenborg; collection date: 1846. See Turni and Kock (2008: 68). Syntype: ZSM, skin only. Collected by: Hedenborg; collection date: 1836. See Turni and Kock (2008: 68). - Comments: Horácek et al. (2000: 137) mention "Bahr-el-Abiad, White Nile" as type locality. Meester et al. (1986: 73): "Bahr-el-Abiad (White Nile), Sudan. Kock (1969a: 154) suggests that the type locality is at Jebel el Funj, between the White and Blue Nile, and Koopman (1975, and in litt.) restricts it to the east (not west, as shown in 1975: 429) bank of the White Nile at 11 45 N 33 30 E, Blue Nile Province, Sudan."). El-Rayah (1980: 518 ISSN 1990-6471 ? ? ? ? 264) believes the restriction to Jebel el Funj is incorrect as Sundevall described the type locaility as Bahr el-Abiad which is the area south of Omdurman (15 19 N 32 29 E) and commonly known as Bahar Abiad and restricts the type locality to the vicinity of Jebel Aulia (15 19 N 32 31 'E). For a deeper discussion on the type locality, see Turni and Kock (2008: 68). - Etymology: According to a legend the mythological King Midas of Phrygia offended Apollo, so the god turned Midas' ears into those of a donkey, i.e. in very long ears; which in this species are relatively large (see Lanza et al., 2015: 304). Mops (Mops) midas: (Name Combination, Current Combination) Mops midas: (Name Combination) Tadarida (Mops) midas: (Name Combination) Tadarida midas: (Name Combination) TAXONOMY: Figure 180. A female Mops midas (TM 48104) caught at Makalali Game Reserve, Limpopo Province, South Africa. Dunlop (1999), following Koopman (1994), recognizes two subspecies: M. m. midas (Sundevall 18437) occuring over most of the range and M. m. miarensis (Grandidier, 1869) occuring in Madagascar (Peterson et al. (1995). Although miarensi has previously been considered a subspecies endemic to the island of Madagascar (Simmons (2005), Ratrimomanarivo et al. (2007) found no distinctive or taxonomically definitive differences between populations of Mops midas occurring on the African continent and Madagascar. COMMON NAMES: Afrikaans: Midas se losstertvlermuis, Midaslosstertvlermuis. Chinese: 米达犬吻蝠. Czech: morous oslí. English: Midas Mops Bat, Midas Bat, Midas Free-tailed Bat, Sundevall's Free-tailed Bat, Midas Groove-cheeked Bat. French: Tadaride Midas, Molosse Midas, Tadarida de Midas, Molosse de Midas. German: MidasBulldoggfledermaus, Sennaarische Doggengrämler. Italian: Carlìno Mìda. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: No known fossils (Dunlop, 1999). CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread but patchy distribution. The species is locally hunted and persecuted, but it is not thought to be declining fast enough to place it in a higher category of threat (Jenkins et al., 2008a; IUCN, 2009; Monadjem et al., 2017ag). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops midas] (Monadjem et al., 2017ag). 2008: LC ver 3.1 (2001) [assessed as Tadarida midas] (Jenkins et al., 2008a; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004ax; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Taylor et al., 2016a). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: This species is thought to be locally threatened by general persecution, collection for food and habitat loss (Jenkins et al., 2008a; IUCN, 2009; Monadjem et al., 2017ag). CONSERVATION ACTIONS: Jenkins et al. (2008a) [in IUCN (2009)] and Monadjem et al. (2017ag) report that there is a need to protect large trees and other known roosting sites for this species (this does not apply to populations on Madagascar). In Madagascar, it is known from Beza Mahafaly and ZombitseVohibasia National Parks. GENERAL DISTRIBUTION: Mops midas is a widespread lowland, savanna species ranging from West Africa eastwards to East Africa and southwards into southern Africa. It has been recorded from the Arabian Peninsula. It is present on Madagascar (including the island of Nosy Be), where it is generally distributed in the drier western and southern habitats of the island below 150 m asl (Ratrimomanarivo et al., 2007). In South Africa, the species' distribution is mostly affected by precipitation seasonality (Babiker African Chiroptera Report 2020 Salata, 2012: 49). Taylor et al. (2018b: 63) predict that the species might also occur in Angola. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 441,324 km2. Native: Benin; Botswana (Archer, 1977; Monadjem et al., 2010d: 544); Burkina Faso (Kangoyé et al., 2015a: 618); Burundi; Central African Republic; Chad; Congo (The Democratic Republic of the); Eritrea (Lavrenchenko et al., 2004b: 133); Ethiopia; Gambia; Ghana; Kenya; Madagascar (Peterson et al., 1995; Rakotonandrasana and Goodman, 2007: 6); Malawi (Ansell and Dowsett, 1988: 47; Monadjem et al., 2010d: 544); Mali; Mozambique; Namibia (Cotterill, 2004a: 260; Monadjem et al., 2010d: 544); Niger; Nigeria; Rwanda; Saudi Arabia; Senegal; South Africa (Monadjem et al., 2010d: 544); Sudan; Swaziland (Shapiro and Monadjem, 2015: "1")): Tanzania; Togo; Uganda; Zambia (Ansell, 1973; Cotterill, 2004a: 260; Monadjem et al., 2010d: 544); Zimbabwe (Monadjem et al., 2010d: 544). BIOGEOGRAPHY: Samonds et al. (2012: 5356) indicate that M. midas seems to have migrated multiple times between the African continent and Madagascar. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Dunlop (1999). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Dunlop (1999). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 62) describe the baculum as an elongated rod, which has a notable twist to the mid-shaft in a lateral view. The distal tip is rounded, and the proximal tip terminates in a blunt point length: 1.90 ± 0.091 (1.80 - 2.05) mm, width: 0.39 ± 0.058 (0.32 - 0.46) mm. For a description of the crania, teeth, ears and tragus, and wingshape and aspect ratio see Dunlop (1999). SEXUAL DIMORPHISM: Sexual dimorphism in M. midas occurs primarily in the stronger development of the sagittal and lambdoidal crests (Kingdon, 1974; Smithers, 1983; Dunlop, 1999), as well as the greater length and width of the skull (Peterson et al., 1995) in males. ECHOLOCATION: See Dunlop (1999) and Taylor (1999b). 519 In Waterberg, South Africa, Taylor et al. (2013b: 18) recorded the following parameters for 6 calls: Fmax: 14.5 ± 1.45 (12.7 - 16.2) kHz, Fmin: 12.8 ± 0.82 (11.7 - 13.7) kHz, Fknee: 13.8 ± 1.02 (12.5 15.0) kHz, Fchar: 13.1 ± 0.88 (12.0 - 14.0) kHz, duration: 10.3 ± 2.89 (7.8 - 15.8) msec. At Farm Welgevonden, South Africa, Taylor et al. (2013a: 556) report a Fknee value of 14 (12 - 15) kHz. Linden et al. (2014: 40) reported the following parameters from the Souhpansberg range (RSA): Fmin: 12 - 14 kHz, Fchar: 12 - 14 kHz, Fknee: 12 - 15 kHz, Slope: 32 - 66 OPS, duration: 8 - 16 msec. At the Mapungubwe National Park (RSA), Parker and Bernard (2018: 57) recorded 13 calls: Fchar: 13.11 ± 1.04 kHz, Fmax: 14.66 ± 1.46 kHz, Fmin: 12.33 ± 1.42 kHz, Fknee: 13.71 ± 0.98 kHz, duration: 14.79 ± 3.78 msec, with 4.29 ± 2.73 calls/sec. From Swaziland, the following values were reported by Monadjem et al. (2017c: 179): Fmin: 11.8 ± 0.50 (11.3 - 12.5) kHz, Fknee: 13.1 ± 1.08 (11.9 - 14.5) kHz, Fc: 12.3 ± 0.72 (11.6 - 13.4) kHz and duration: 14.2 ± 3.89 (9.0 - 20.7) msec. Weier et al. (2020: Suppl.) reported on four calls from the Okavango River Basin with the following characteristics: Fmax: 18.51 ± 5.52 kHz, Fmin: 14.78 ± 4.17 kHz, Fknee: 16.79 ± 4.44 kHz, Fchar: 15.27 ± 3.89 kHz, slope: 25.69 ± 20.05 Sc, duration: 10.63 ± 5.28 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) described the karyotype as having a diploid number of 2n = 48, aFN = 66, BA = 20, X = SM, Y = A, this is supported by Rautenbach et al. (1993). Protein / allozyme - Unknown. HABITAT: See Dunlop (1999). HABITS: See Dunlop, 1999). ROOST: See Dunlop (1999). MIGRATION: May make local migrations, as they only appear in parts of their range (northeastern Congo) at the end of the dry season (Allen et al., 1917; Dunlop, 1999). DIET: In the Limpopo province (RSA), Taylor et al. (2017b: 246, 252 - 254) found 93 prey items spread over seven insect orders: Lepidoptera (100 520 ISSN 1990-6471 % [found in all faecal pellets] and covering 17 families: Synemon austera + Talanga sexpunctalis + Antaeotricha sp. + Elachista sp. + Araeopteron sp. + Dyops subdifferens + Eublemma sp. + Peridrome orbicularis + Spargaloma sexpunctata + Eutelia polychorda + Lophoptera sp. + Phlegetonia callichroma + Acanthovalva inconspicuaria + Achrosis pyrrhularia + Aethalura punctulata + Cabera AH02SA + Chiasmia interrupta + Ennominae sp. + Eupithecia sp. + Isturgia arizeloides + Isturgia roraria + Isturgia pulinda + Scopula atricapilla + Hyblaea puera + Gonometa postica + Sena sp. + Sena prompta + Xenopseustis sp. + Acronicta psi + Adisura marginalis + Athetis ignava + Bastilla torrida + Bryophila AH01Pe + Chasmina lispodes + Chasmina sp. + Condica parista + Copitarsia + Cortyta canescens + Enmonodia capensis + Grammodes stolida + Harrisimemna trisignata + Heterochroma BioLep767 + Hypotacha sp. + Leucania BioLep10 + Leucania BioLep162 + Leucania multilinea + Leucania Poole05 + Leucania porphyrodes + Mythimna consanguis + Mythimna convecta + Mythimna impura + Mythimna oxygala + Mythimna nigrisparsa + Mythimna obsoleta + Mythimna sp. + Niphonyx segregata + Hadenini sp. + Ophiusa tirhaca + Pandesma robusta + Paratrachea laches + Pericyma atrifusa + Polia piniae + Rhesala moestalis + Spodoptera exigua + Tycomarptes sp. + Xanthodes malvae + Garella nilotica + Maurilia arcuata + Meganola dentata + Meganola baracoana + Pseudosomera noctuiformis + Stenoptilia sp. + Phycitinae + Copaxa lavendera + Pseudautomeris irene + Agrius convolvuli + Cochylidia implicitana + Platphalonidia felix + unmatched), Coleoptera (67 % - Cybister tripunctatus + unmatched), Hemiptera (29 % Macrorhaphis acuta + Nezar viridula + unmatched), Diptera, Orthoptera (Tettigoniinae), Blattodea (25 % - Pycnoscelus sp. + unmatched) and Neuroptera (29 % - unmatched). POPULATION: Structure Verschuren (1957) found females outnumbered males in the roost at a ratio of four to one. Density Smithers, 1983) reported M. midas roosting in groups of up to several hundred; however, Verschuren, 1957) only recorded groups of 10-20 individuals leaving roosts. In Madagascar, no large colonies have been found and it is thought to be a locally common species with a patchy distribution. The maximum recorded colony was near Amboasary of 600 individuals (Jenkins et al., 2008a; IUCN, 2009; Monadjem et al., 2017ag). Trend 2016: Decreasing (Monadjem et al., 2017ag). 2008: Decreasing (Jenkins et al., 2008a; IUCN, 2009). ACTIVITY AND BEHAVIOUR: See Dunlop (1999). REPRODUCTION AND ONTOGENY: See Dunlop (1999). Krutzsch (2000: 127) indicates that this species is polyoestrous. Happold and Happold (1990b: 568) reported that in Malawi, births occur at the end of the wet season. Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] suggest that the birth season runs from December to March. MATING: See Dunlop (1999). PARASITES: Several mites have been recorded for this species, including Chelanyssus aethiopicus (Family Macronyssidae) and Nycteriglyhus tadaridae (Family Rosensteiniidae) (Anciaux de Faveaux, 1971; 1976a). Bat fleas (Siphonaptera) have also been recorded (Dunlop, 1999). Lagaropsylla hoogstraali (Ischnopsyllidae: Ischnopsyllinae) was found amongst a batch of fleas taken from M. midas in the Kruger National Park, South Africa in the Pafuri area (Segerman and Braack, 1988). SIPHONAPTERA Ischnopsyllidae: Lagaropsylla hoogstraali Smit, 1957 from Rwanda (Haeselbarth et al., 1966: 190), and Madagascar (Hastriter, 2016: 17). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in 15 individuals from Limpopo Province, South Africa, none were tested positive (0/15). One out of two specimens tested by Geldenhuys et al. (2013: 517) in Makhado, Limpopo, RSA was found to be positive for CoV. Paramyxoviridae Mélade et al. (2016b: 4) tested 19 Madagascan specimens of which one was positive for paramyxoviruses. UTILISATION: In Madagascar, Goodman et al. (2008d) found that these bats were caught, cooked and then fed to domestic pigs. African Chiroptera Report 2020 521 Figure 181. Distribution of Mops (Mops) midas Mops (Mops) midas miarensis (A. Grandidier, 1869) *1869. Nyctinomus Miarensis A. Grandidier, Rev. Mag. Zool., ser. 2, 21: 337. Publication date: September 1869. Type locality: Madagascar: between Soua-haze and Soua-hané [sic]: Miari [probably Miary near Tuléar] [Goto Description]. Holotype: MNHN 29: ad ♀. Presented/Donated by: ?: Collector Unknown. Holotype: MNHN ZM-MO-1997-1851: ad ♀, alcoholic (skull not removed). Collected by: Alfred Grandidier. See Peterson et al. (1995: 174). Number 229 in Rode (1941). 1870. Nyctinomus unicolor A. Grandidier, Rev. Mag. Zool., ser. 2, 22: 49. Publication date: February 1870. Type locality: Madagascar: "Madagascar" [Goto Description]. ? Mops (Mops) midas miarensis: (Name Combination, Current Combination) ? Mops miarensis: (Name Combination) ? Mops midas miarensis: (Name Combination) GENERAL DISTRIBUTION: E, SW, S Madagascar. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Mops (Mops) midas midas (Sundevall, 1843) *1843. Dysopes midas Sundevall, Kongl. Svenska Vet.-Akad. Hand., ser. 3, 30: 207, pl. 2, fig. 7 [for 1842]. Publication date: 1843. Type locality: Sudan: Blue Nile Province: E-bank White Nile: Valid Sohey [= Wad Shala]: restricted by Turni and Kock (2008: 68). [14 34 N 13 48 E]. Syntype: BMNH 1846.6.2.20: ad ♂, skin and skull. Formerly Stockholm Museum (ZSM no. 523), locality: Bahr el Abiad. See Turni and Kock (2008: 68). Syntype: ZMB 535: skin only. Collected by: Hedenborg; collection date: 1831 - 1835. From [Dar] Sennar, Sudan; see Turni and Kock (2008: 68). Syntype: ZSM. Collected by: Hedenborg; collection date: 1846. See Turni and Kock (2008: 68). Syntype: ZSM, skin only. Collected by: Hedenborg; collection date: 1836. See Turni and Kock (2008: 68). - Comments: Horácek et al. (2000: 137) mention "Bahr-el-Abiad, White Nile" as type locality. Meester et al. (1986: 73): "Bahr-el-Abiad (White Nile), Sudan. Kock (1969a: 154) suggests that the type locality is at Jebel el Funj, between the White and Blue Nile, and Koopman (1975, and in litt.) restricts it to the east (not west, as shown in 1975: 429) bank of the White Nile at 11 45 N 33 30 E, Blue Nile Province, Sudan."). El-Rayah (1980: 264) believes the restriction to Jebel el Funj is incorrect as Sundevall described the type locaility as Bahr el-Abiad which is the area south of Omdurman (15 19 N 32 29 E) and commonly known as Bahar Abiad and restricts the type locality to the vicinity of Jebel Aulia (15 19 N 32 31 'E). For a deeper discussion on the type locality, see Turni and Kock (2008: 68). - Etymology: According to a legend the mythological King Midas of Phrygia 522 ISSN 1990-6471 ? ? ? offended Apollo, so the god turned Midas' ears into those of a donkey, i.e. in very long ears; which in this species are relatively large (see Lanza et al., 2015: 304). Mops (Mops) midas midas: (Name Combination, Current Combination) Mops midas midas: (Name Combination) Tadarida midas midas: (Name Combination) GENERAL DISTRIBUTION: Continental Africa; Arabia. Kenya, Malawi, Namibia, Niger, Nigeria, Senegal, South Africa, South Sudan, Sudan, Uganda, Zambia, Zimbabwe. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Botswana, Burkina Faso, Cameroon, Congo (Democratic Republic of the), Ethiopia, Mops (Mops) niangarae J.A. Allen, 1917 *1917. Mops niangaræ J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 468. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Niangara [03 24 N 27 52 E] [Goto Description]. Holotype: AMNH 48901: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 12 December 1910; original number: 1313. Etymology: Referring to the locality where the type specimen was found. ? Mops (Mops) niangarae: (Name Combination, Current Combination) ? Mops niangarae: (Current Spelling) ? Tadarida congica niangarae: (Name Combination) TAXONOMY: Peterson (1972) and Thorn et al. (2009: 68) included this species in trevori. Hayman and Hill (1971) listed it as a subspecies of Tadarida congica (=Mops congicus). Freeman (1981b: 111, 159) considered it a distinct species pending collection of additional specimens. Additional studies are needed into the systematic status of Mops niangarae, especially with regards to its relationship to M. trevori (see Hayman and Hill, 1971; Peterson, 1972; Freeman, 1981b; Simmons, 2005). COMMON NAMES: Czech: morous niangaranský. English: Niangara Mops Bat, Niangaran Mops Bat, Niangara Freetailed Bat. French: Tadaride du Niangara. German: Niangara-Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of continuing problems with its taxonomy, and lack of recent information on its extent of occurrence, ecological requirements and threats (Mickleburgh et al., 2008bq; IUCN, 2009; Mickleburgh et al., 2014). Assessment History Global 2014: DD ver 3.1 (2001) (Mickleburgh et al., 2014). 2008: DD ver 3.1 (2001) [assessed as Tadarida niangarae] (Mickleburgh et al., 2008bq; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004bj; IUCN, 2004). 2002: CR B1+2c (Mickleburgh et al., 2002a: 22). 1996: DD (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are currently unknown (Mickleburgh et al., 2008bq; IUCN, 2009; Mickleburgh et al., 2014). CONSERVATION ACTIONS: Mickleburgh et al. (2014) repeats what stated by Mickleburgh et al. (2008bq) [in IUCN (2009)] who report that it is not known if the species is present in any protected areas. Additional studies are needed into the distribution, natural history and threats to this little-known bat. GENERAL DISTRIBUTION: Mops niangarae is known only from the holotype collected from "Niangara, northeastern Belgian Congo" (Democratic Republic of the Congo near the border with Sudan) (Allen et al., 1917; Lang and Chapin, 1917a). Native: Congo (The Democratic Republic of the) (Allen et al., 1917; Lang and Chapin, 1917a). African Chiroptera Report 2020 523 POPULATION: Structure and Density:- It is known only from the holotype (Mickleburgh et al., 2008bq; IUCN, 2009; Mickleburgh et al., 2014). Trend:- 2014: Unknown (Mickleburgh et al., 2014). 2008: Unknown (Mickleburgh et al., 2008bq; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the). Figure 182. Distribution of Mops (Mops) niangarae Mops (Mops) niveiventer Cabrera and Ruxton, 1926 *1926. Mops angolensis niveiventer Cabrera and Ruxton, Ann. Mag. nat. Hist., ser. 9, 17 (103): 594. Publication date: 1 May 1926. Type locality: Congo (Democratic Republic of the): Kasai Occidental province: St. Joseph de Luluabourg [=Kananga] [ca. 05 53 S 22 26 E, 610 m] [Goto Description]. Holotype: BMNH 1926.7.6.109: ♀, skin and skull. Collection date: 16 April 1924; original number: 409. 1937. Mops chitauensis J.Eric Hill, Am. Mus. Novit., 916: 2, 3, text-fig. 1. Publication date: 17 April 1937. Type locality: Angola: Malange district: Chitau [11 15 S 17 01 E, 4 930 ft] [Goto Description]. Holotype: AMNH 88116: ad ♂, skin and skull. Collected by: Lee S. Bradley and the Phipps-Bradley Expedition; collection date: 10 February 1933; original number: 676. 1975. Tadarida nivieventer: Fenton, Life Sci. Occ. Pap., R. Ont. Mus., 104: 23. (Lapsus) ? Mops (Mops) niveiventer: (Name Combination, Current Combination) ? Mops niveiventer: (Name Combination) ? Tadarida (Mops) niveiventer: (Name Combination) ? Tadarida niveiventer: (Name Combination) TAXONOMY: Koopman (1993a) suggested that niveiventer is possibly a subspecies of demonstrator. Here considered a distinct species following Simmons (2005). COMMON NAMES: Czech: morous sněhobílý. English: White-bellied Mops Bat, White-bellied Free-tailed Bat. French: Tadaride à ventre blanc. German: WeißbauchBulldoggfledermaus. Portuguese: Morcego de cauda livre de ventre branco. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Cotterill, 2008f; IUCN, 2009; Monadjem and Cotterill, 2017c). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops niveiventer] (Monadjem and Cotterill, 2017c). 2008: LC ver 3.1 (2001) [assessed as Tadarida niveiventer] (Cotterill, 2008f; IUCN, 2009). 2004: LC ver 3.1 (2001) (Cotterill, 2004f; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: This species is locally threatened in parts of its range, often through general deforestation and the conversion of suitable habitat by shifting agricultural practices (Cotterill, 2008f; IUCN, 2009; Monadjem and Cotterill, 2017c). 524 ISSN 1990-6471 CONSERVATION ACTIONS: Cotterill (2008f) [in IUCN (2009)] and Monadjem and Cotterill (2017c) report that there appear to be no direct conservation measures in place for this species. It is not known if the species is present within any protected areas. Further studies are needed into the distribution and systematics of this species. GENERAL DISTRIBUTION: Mops niveiventer is present in Central Africa and parts of Southern Africa. It has been recorded from eastern Angola, southern Democratic Republic of the Congo, northern Zambia, Rwanda, northern Mozambique and possibly from Malawi. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 722,774 km2. Records from Botswana and Madagascar are definitely condylurus (see Hayman and Hill, 1971: 61; Meester et al., 1986: 74; Simmons, 2005). Native: Angola (Monadjem et al., 2010d: 544); Burundi; Cameroon (Bol A Anong et al., 2011: 48; Bakwo Fils et al., 2014: 4); Congo (The Democratic Republic of the) (Hayman et al., 1966; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 544); Mozambique (Monadjem et al., 2010d: 544); Rwanda; Tanzania; Zambia (Ansell, 1973; Monadjem et al., 2010d: 544). Presence uncertain: Malawi. The record from Cameroon is probably doubtful (Jakob Fahr, pers. comm.). ECHOLOCATION: From Maroua, Cameroon, Manga Mongombe (2012: 79) and Bakwo Fils et al. (2018: 4) reported a FM call type for 10 calls and the following parameters: Fmax: 37.7 ± 2.9 (32.2 - 40.8) kHz, Fmin: 34.1 ± 2.6 (29.9 - 37.6) kHz, Fmean: 35.9 ± 2.8 (31.1 - 39.3) kHz, Fknee: 37.6 ± 2.7 (32.1 - 40.8) kHZ, Fchar: 34.1 ± 2.6 (30.1 - 37.6) kHz, and duration: 0.74 ± 0.10 (0.65 - 0.95) msec. Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 20.3 kHz, Fstart: NA kHz, Fend: NA kHz, Band width: NA kHz, and duration: 8.1 msec. DIET: In fecal pellets from Maroua (Cameroon), Bol A Anong (2013: 32) found Coleoptera (64.75%), Hemiptera (25.00%), Diptera (9.53%) and Lepidoptera (0.23%). POPULATION: Structure and Density:- This is a locally common species that can be found in large colonies of up to hundreds of individuals within buildings, but is more often present in smaller numbers (Cotterill, 2008f; IUCN, 2009; Monadjem and Cotterill, 2017c). Trend:- 2016: Unknown (Monadjem and Cotterill, 2017c). 2008: Unknown (Cotterill, 2008f; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Burundi, Cameroon, Congo (Democratic Republic of the), Mozambique, Rwanda, Zambia, Zimbabwe. DENTAL FORMULA: Dorst (1957b: 134) indicated the milk dentition of Mops angolensis niveiventer shows some differences with the dentition of Mops angolensis he described previously (Dorst, 1949), a fact he attributed to it being a different geographic race. However, the latter form is currently considered to belong to a separate species: Mops (Mops) condylurus. The upper milk toothrow of niveiventer consists of three functional teeth wich are reduced to pegs, and two additional, vestigial teeth. In the lower toothrow there are three incisors of different shape and a remarkably developed canine. There is also one premolar, which is clearly developed, but not functional as it doesn't erupt through the gum. This tooth is resorbed in the young. Figure 183. Distribution of Mops (Mops) niveiventer African Chiroptera Report 2020 525 Mops (Mops) trevori J.A. Allen, 1917 *1917. Mops trevori J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 469, pl. 48, fig. 2. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Faradje [03 44 N 29 43 E] [Goto Description]. Holotype: AMNH 49250: ad ♀, skull and alcoholic. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 29 September 1912; original number: 1954. - Etymology: In honour of John B. Trevor, a Trustee of the American Museum of Natural History and Chairman of the Committee on African Exploration, whose enthousiasm and generosity in support of the Congo Expedition contributed to its success (see Allen, 1917: 469). 1996. Mops trevoli: Kityo and Kerbis, J. East Afr. Nat. Hist., 85: 63. (Lapsus) ? Mops (Mops) trevori: (Name Combination, Current Combination) ? Tadarida trevori: (Name Combination) TAXONOMY: Formerly included in niangarae; see Freeman (1981b: 111, 159). been recorded from some protected areas. Further studies are needed into the distribution, natural history and threats to this little-known bat. COMMON NAMES: Czech: morous Trevorův. English: Trevor's Mops Bat, Trevor's Free-tailed Bat, Trevor´s Bat. French: Tadaride d'Allen, Tadaride de Trevor. German: Trevors Bulldoggfledermaus. There appear to be no direct conservation measures in place, however, it has been recorded from some protected areas. Further studies are needed into the distribution, natural history and threats to this little-known bat. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since, although it has been recorded over a very wide area, there are only very few records, and very little is known about its status, threats and habitat requirements (Mickleburgh et al., 2008bv; IUCN, 2009; Mickleburgh et al., 2014a). GENERAL DISTRIBUTION: Mops trevori has been patchily recorded in parts of West and Central Africa. In West Africa It has been recorded from Guinea, Côte d'Ivoire, Comoe in Ghana, and Agege in Nigeria. In Central Africa it has been recorded from the Central African Republic, parts of eastern Democratic Republic of the Congo and western Uganda. Assessment History Global 2014: DD ver 3.1 (2001) [assessed as Mops trevori] (Mickleburgh et al., 2014a). 2008: DD ver 3.1 (2001) [assessed as Tadarida trevori] (Mickleburgh et al., 2008bv; IUCN, 2009). 2004: VU A4c ver 3.1 (2001) (Mickleburgh et al., 2004aw; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Native: Central African Republic; Congo (The Democratic Republic of the); Côte d'Ivoire; Ghana; Guinea; Nigeria; Sierra Leone (Weber et al., 2019: 28 - first record); Sudan; Uganda (Kityo and Kerbis, 1996: 63). Regional None known. MAJOR THREATS: This species is generally threatened by habitat loss, resulting from the conversion of land to agricultural use, and the extraction of firewood and timber (Mickleburgh et al., 2008bv; IUCN, 2009; Mickleburgh et al., 2014a). CONSERVATION ACTIONS: Mickleburgh et al. (2014a) repeats what stated by Mickleburgh et al. (2008bv) [in IUCN (2009)] who report that there appear to be no direct conservation measures in place, however, it has POPULATION: Structure and Density:- Little is known about the abundance of this apparently rare species, but it is presumed that colony size is likely to be small (Mickleburgh et al., 2008bv; IUCN, 2009; Mickleburgh et al., 2014a). Trend:- 2014: Decreasing (Mickleburgh et al., 2014a). 2008: Decreasing (Mickleburgh et al., 2008bv; IUCN, 2009). REPRODUCTION AND ONTOGENY: One of two females reported from Sierra Leone by Weber et al. (2019: 28) was pregnant on 25 March. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Nigeria, Sierra Leone, South Sudan, Uganda. 526 ISSN 1990-6471 Figure 184. Distribution of Mops (Mops) trevori Subgenus Mops (Xiphonycteris) Dollman, 1911 *1911. Xiphonycteris Dollman, Ann. Mag. nat. Hist., ser. 8, 7 (38): 210. Publication date: 1 February 1911 [Goto Description]. - Comments: Type species: Xiphonycteris spurrelli Dollman, 1911. ? Mops (Xiphonycteris): (Name Combination, Current Combination) ? Tadarida (Xiphonycteris): (Name Combination) Mops (Xiphonycteris) bakarii Stanley, 2009 *2008. Mops bakarii Stanley, Acta Chiropt., 10(2): 184, Figs 2 -6. Publication date: December 2008. Type locality: Tanzania: Pemba Island, Kipangani Village: in attic of hospital [4.96487 S 39.71456 E, 12m]. Holotype: FMNH 192895: ad ♂, skin and skull. Collected by: William ("Bill") T. Stanley; collection date: 5 August 2006; original number: WTS 7307. Paratype: FMNH 192824: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7312. Paratype: FMNH 192825: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7313. Paratype: FMNH 192826: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7327. Paratype: FMNH 192827: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7328. Paratype: FMNH 192828: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7329. Paratype: FMNH 192829: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7335. Paratype: FMNH 192830: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7336. Paratype: FMNH 192831: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7337. Paratype: FMNH 192832: ad ♀. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7338. Paratype: FMNH 192833: ad ♀. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7339. Paratype: FMNH 192834: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7345. Paratype: FMNH 192835: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7347. Paratype: FMNH 192836: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7348. Paratype: FMNH 192894: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7306. Paratype: FMNH 192896: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7308. Paratype: FMNH 192897: ad ♂. African Chiroptera Report 2020 ? 527 Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7322. Paratype: FMNH 192898: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7331. Paratype: FMNH 192899: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7332. Paratype: FMNH 192900: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7333. Paratype: FMNH 192901: ad ♂. Collected by: William ("Bill") T. Stanley; collection date: 5 - 6 August 2006; original number: WTS 7334. - Etymology: In honour of Dr. Bakari Asseid, director of the Department of Commercial Crops, Fruits and Forestry, Zanzibar, to recognize his significant contributions to the conservation of natural habitats and biota of Zanzibar (including both Pemba and Unguja Islands) (see Stanley, 2008: 185). Mops (Xiphonycteris) bakarii: (Name Combination, Current Combination) COMMON NAMES: Czech: morous Stanleyův. German: Bakaris Bulldoggfledermaus, Pemba-Bulldoggfledermaus. CONSERVATION STATUS: Global Justification Shapiro (2017) report this species is listed as Data Deficient due to only one known locality and lack of further data, while population size is not known. POPULATION: Structure and Density:- There is no information about the species population size and population trend (Shapiro, 2017). Trend:- 2016: Unknown (Shapiro, 2017). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Tanzania. Assessment History Global 2016: DD ver. 3.1 (2001) [assessed as Mops bakarii] (Shapiro, 2017). Regional None known. MAJOR THREATS: Specific threats to this species are known at present (Shapiro, 2017). CONSERVATION ACTIONS: Shapiro (2017) report there are no specific conservation measures are in place for this species. GENERAL DISTRIBUTION: O'Brien (2011: 288) reports it from Pemba. Figure 185. Distribution of Mops (Xiphonycteris) bakarii Mops (Xiphonycteris) brachypterus (Peters, 1852) *1852. Dysopes brachypterus Peters, Naturwissenschaftliche Reise nach Mossambique, Zoologie, Säugethiere, 59, pl. 15, fig. 1. Publication date: 1852. Type locality: Mozambique: Mozambique Island [15 03 S 40 46 E] [Goto Description]. Holotype: ZMB 536/85537: ad ♂, skin and skull. Collected by: Prof. Wilhelm Carl Hartwig Peters; collection date: between 1843 and 1847. See Turni and Kock (2008) [skin: ZMB 536 + skull: ZMB 85537 (=An 16356)]. Neotype: ROM 46721: ad ♂, skin and skull. Collected by: Terence Michael Shortt and A.E. Williams; collection date: 24 June 1968; original number: SG301. Presented/Donated by: ?: Collector Unknown. - Etymology: From the Greek adjective "βρᾰχύς" (brakhìs), meaning "short" and the Greek neuter substantive "πτερόν" (pteròn), meaning "wing"; but actually the species' wings are not particularly short (see Lanza et al., 2015: 295). 1877. Nyctinomus [(Nyctinomus)] brachypterus: Dobson, Proc. zool. Soc. Lond., 1876, IV: 722. Publication date: April 1877. (Name Combination) 528 ISSN 1990-6471 1908. 1917. 1980. ? ? ? ? ? ? ? ? ? ? ? Nyctinomus leonis Thomas, Ann. Mag. nat. Hist., ser. 8, 2 (10): 373. Publication date: 1 October 1908. Type locality: Sierra Leone: "Sierra Leone" [Goto Description]. Holotype: BMNH 1862.12.23.3: ad ♂, skin and skull. Presented/Donated by: J. Brown Esq. (Specimen "c" of Dobson's catalogue). Nyctinomus ochraceus J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 455. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Medje [02 25 N 27 18 E, 800 m] [Goto Description]. Holotype: AMNH 48821: ad ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 16 March 1910; original number: 745. Tadarida (Mops) brachyptera: El-Rayah, Systematics of African molossid bats of the subgenus Xiphonycteris of the genus Tadarida (Molossidae: Chiroptera) - PhD Thesis University of Toronto:, 231. Chaerephon leonis ochraeus: Mops (Xiphonycteris) brachypterus: (Name Combination, Current Combination) Mops brachypterus leonis: (Name Combination) Mops brachypterus ochraceus: (Name Combination) Mops brachypterus: (Name Combination) Tadarida brachyptera leonis: (Name Combination) Tadarida brachyptera: (Name Combination) Tadarida leonis leonis: Tadarida leonis ochraeus: Tadaride brachyptera: (Name Combination) Xiphonycteris leonis: TAXONOMY: Includes leonis; see El-Rayah (1981: 6). Stanley (2008: 190) points out that brachypterus is in need of revision, since the dental features (e.g. size of upper canines and first upper premolar) differ between for the holotype and continental specimens attributed to it. Monadjem et al. (2016y: 368) and Herkt et al. (2017: Appendix S9) found differences in the ecological niches and a clear geographic gap between the distribution areas of brachypterus and leonis sensu stricto, but as genetic tests evaluating the distinct species status are still lacking, they are still considered to represent the same species. COMMON NAMES: Castilian (Spain): Murciélago Guanero. Chinese: 短翼犬吻蝠. Czech: morous krátkokřídlý. English: Sierra Leone Mops Bat, Short-winged Mops Bat, Sierra Leone Free-tailed Bat, Whitefingered Free-tailed Bat. French: Tadaride de Peters, Tadaride à doigts blancs. German: Kurzflügel-Bulldoggfledermaus, Kurzflügeligen Doggengrämler. Italian: Carlìno brachìttero. CONSERVATION STATUS: Global Justification Listed as Least Concern in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bo; IUCN, 2009; Monadjem et al., 2017az). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops brachypterus] (Monadjem et al., 2017az). 2008: LC ver 3.1 (2001) [assessed as Tadarida brachyptera] (Mickleburgh et al., 2008bo; IUCN, 2009). 2004: LC ver 3.1 (2001) [assessed as Mops brachypterus] (Mickleburgh et al., 2004bo; IUCN, 2004). Regional None known. MAJOR THREATS: The species is threatened in parts of its range by deforestation, presumably mostly through logging and mining activities and the conversion of land to agricultural use (Mickleburgh et al., 2008bo; IUCN, 2009; Monadjem et al., 2017az). CONSERVATION ACTIONS: Mickleburgh et al. (2008bo) [in IUCN (2009)] and Monadjem et al. (2017az) report that it is not known if the species is present in any protected areas. Further studies are needed into the taxonomy, distribution, natural history, and threats to this species. GENERAL DISTRIBUTION: Mops brachypterus ranges widely from West Africa to the East African coast. In West Africa, it is distributed from the Gambia and Sierra Leone in the west, through to Nigeria and Cameroon in the east. From Cameroon, it occurs patchily eastwards through the Congo Basin to Uganda, African Chiroptera Report 2020 and to coastal parts of Kenya, Tanzania (including the islands of Unguja [=Zanzibar] [where its presence is questioned according to O'Brien (2011: 289)] and Mafia) and Mozambique in the south. Native: Cameroon; Central African Republic; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 543); Côte d'Ivoire; Equatorial Guinea; Gabon; Ghana; Guinea (Simons et al., 2014: 284); Kenya; Liberia; Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 543); Nigeria; Sierra Leone; Tanzania; Togo. Presence uncertain: Zanzibar (Stanley, 2008: 191). 529 POPULATION: Structure and Density:- This is a common species (Mickleburgh et al., 2008bo; IUCN, 2009; Monadjem et al., 2017az). Trend:- 2016: Unknown (Monadjem et al., 2017az). 2008: Unknown (Mickleburgh et al., 2008bo; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Burkina Faso, Cameroon, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Guinea, Liberia, Malawi, Mozambique, Nigeria, Rwanda, Senegal, Sierra Leone, Tanzania, Uganda. Bates et al. (2013: 339) reject the presence of this species in the Republic of Congo. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) described the karyotype as having a diploid number of 2n = 48, aFN = 54, BA = 8, and a presumably submetacentric X (one female studied). HABITAT: Monadjem et al. (2016y: 368) captured this bat flying over small forest streams in the Mount Nimba area, at altitudes between 450 and 550 m. PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. Figure 186. Distribution of Mops (Xiphonycteris) brachypterus Mops (Xiphonycteris) nanulus J.A. Allen, 1917 *1917. Mops (Allomops) nanulus J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 477. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Niangara [03 24 N 27 52 E] [Goto Description]. Holotype: AMNH 48864: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 12 December 1910; original number: 1318. Topotype: AMNH 48860: ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition. Presented/Donated by: ?: Collector Unknown. Topotype: AMNH 48861: ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition. Presented/Donated by: ?: Collector Unknown. Topotype: AMNH 48862: ♀, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition. Presented/Donated by: ?: Collector Unknown. Topotype: AMNH 48863: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition. Presented/Donated by: ?: Collector Unknown. Topotype: AMNH 48865: ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition. Presented/Donated by: ?: Collector Unknown. 1940. Mops calabarensis Hayman, in: Sanderson, Trans. Linn. Soc. Lond., 24 (7) 1: 677. Publication date: January 1940. Type locality: Nigeria: Eastern region, Calabar: Ikotmbo [05 03 N 18 19 E, 50 ft]. Holotype: BMNH 1939.318: ♂, skin and skull. Collected by: Ivan T. Sanderson; collection date: 18 September 1932; original number: 3. 530 ISSN 1990-6471 1940. 2015. ? ? ? ? Mops nannulus: Sanderson, Trans. Linn. Soc. Lond., 24 (7) 1: 677. Publication date: January 1940. - Comments: Lapsus (see Grubb et al. (1998: 96). (Lapsus) Mops annulus: Gunnel, Butler, Greenwood and Simmons, Am. Mus. Novit., 3846: 34. Publication date: 16 December 2015. (Lapsus) Mops (Xiphonycteris) nanulus: (Name Combination, Current Combination) Mops nanulus: (Name Combination) Tadarida nanula calabarensis: (Name Combination) Tadarida nanula: (Name Combination) TAXONOMY: Distinction from spurrelli not certain; see Koopman (1989a). COMMON NAMES: Chinese: 侏 犬 吻 蝠 . Czech: morous trpasličí. English: Dwarf Mops Bat, Dwarf Free-tailed Bat. French: Tadaride naine. German: ZwergBulldoggfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008w; IUCN, 2009; Monadjem et al., 2017l). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops nanulus] (Monadjem et al., 2017l). 2008: LC ver 3.1 (2001) [assessed as Tadarida nanula] (Mickleburgh et al., 2008w; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004o; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: In general, there are no threats to this widespread and somewhat adaptable species. In parts of its range it is locally threatened by habitat loss, primarily deforestation resulting from logging operations and the conversion of land to agricultural use (Mickleburgh et al., 2008w; IUCN, 2009; Monadjem et al., 2017l). CONSERVATION ACTIONS: Mickleburgh et al. (2008w) [in IUCN (2009)] and Monadjem et al. (2017l) report that it is not known if the species is present in any protected areas. Further studies are needed into the distribution, abundance, natural history, and threats to this species. GENERAL DISTRIBUTION: Mops nanulus is widely, but patchily, recorded in West, Central and East Africa. It ranges from Sierra Leone and Guinea in the west, through West Africa to Cameroon, and from here east to southern Sudan, Uganda, and western Kenya with an additional record from western Ethiopia and as far south as the southern portion of the Democratic Republic of the Congo. Start (1969: 222) suggests that the species' range might have been more continuous during the Pleistocene when there was a higher degree of rainfall, resulting in a continuous belt of lowland forest between Congo and Kenya. Native: Benin (Capo-Chichi et al., 2004: 163); Cameroon; Congo (The Democratic Republic of the) (Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 544); Côte d'Ivoire; Ethiopia; Gambia (Emms and Barnett, 2005: 50; Simmons, 2005); Ghana; Guinea (Fahr et al., 2006a: 73; Decher et al., 2016: 276); Kenya (Start, 1969: 220); Liberia (Fahr, 2007a: 104); Nigeria; Sierra Leone; Sudan; Uganda. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Start (1969: 221) indicates that there appears to be a general cline from pale-winged red forms in the west to dark-winged brown ones in the east, although this is not absolute. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) reported 2n = 48, FN = 54, BA = 8, a submetacentric X chromosome and an acrocentric Y chromosome. Protein / allozyme - Unknown. POPULATION: Structure and Density:- This is a common species (Mickleburgh et al., 2008w; IUCN, 2009; Monadjem et al., 2017l). Trend:- 2016: Unknown (Monadjem et al., 2017l). 2008: Unknown (Mickleburgh et al., 2008w; IUCN, 2009). REPRODUCTION AND ONTOGENY: Start (1969: 221) reports on six female specimens collected in West Pokot (Kenya) between 25 and African Chiroptera Report 2020 531 29 August 1967, of which five were carrying well developed embryos, whereas the sixth was lactating. Weber et al. (2019: 28) reported one pregnant female from Sierra Leone on 26 March. Krutzsch (2000: 127) indicates that "Tadarida (Mops) nanulus" is polyoestrous. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Benin, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ethiopia, Gabon, Ghana, Guinea, Kenya, Nigeria, Sierra Leone, South Sudan, Togo, Uganda. Figure 187. Distribution of Mops (Xiphonycteris) nanulus Mops (Xiphonycteris) petersoni (El Rayah, 1981) *1981. Tadarida [(Xiphonycteris)] petersoni El Rayah, Life Sci. Occ. Pap., R. Ont. Mus., 36: 3, figs 1 - 2. Publication date: 11 December 1981. Type locality: Cameroon: Kumba, 15 km S [04 39 N 09 26 E] [Goto Description]. Holotype: ROM 55813: ad ♂, skull and alcoholic. Collected by: Randolph Lee Peterson; collection date: 27 January 1970; original number: 0364. See: 3). Paratype: AMNH 236332: ♂, skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. 6 km SE Eseka(03°35'N 10°44'E) (See El-Rayah, 1981: 3). Paratype: AMNH 236333: ♂, skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. 6 km SE Eseka(03°35'N 10°44'E) (see: El-Rayah, 1981: 3). Paratype: AMNH 241075: ♂, skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Near Okola (04°00'N 11°33'E) (see El-Rayah, 1981: 3). Paratype: AMNH 241076: ♂, skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. 30 km E Nanga Emboko (04°38'N 12°21'E) (see El-Rayah, 1981: 3). Paratype: AMNH 241078: ♀, skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. 30 km E Nanga Emboko (04°38'N 12°21'E) (see El-Rayah, 1981: 3). - Etymology: In honour of Randolph L. Peterson, collector of the type specimen. ? Mops (Xiphonycteris) petersoni: (Name Combination, Current Combination) ? Mops petersoni: (Name Combination) ? Tadarida petersoni: (Name Combination) TAXONOMY: Described in Tadarida (Xiphonycteris), but see comments under Tadarida and Mops. primary forest habitat, which makes the species close to qualifying for Vulnerable (Mickleburgh et al., 2010). COMMON NAMES: Czech: morous kamerunský. English: Peterson's Mops Bat, Peterson's Free-tailed Bat. French: Tadaride de Peterson. German: Petersons Bulldoggfledermaus. Assessment History Global 2010: NT ver 3.1 (2001) [assessed as Mops pertersoni] (Mickleburgh et al., 2010). 2008: VU A4c ver 3.1 (2001) [assessed as Tadarida petersoni] (Mickleburgh et al., 2008bt; IUCN, 2009). 2004: VU A4c ver 3.1 (2001) (Mickleburgh et al., 2004av; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). CONSERVATION STATUS: Global Justification Listed as Near Threatened because this species is likely in significant decline (but probably at a rate of less than 30% over a 15 year period; i.e., three generations) because of ongoing loss of its Regional None known. 532 ISSN 1990-6471 MAJOR THREATS: The species is considered to be threatened by loss and degradation of forest habitats within its known range, largely through the conversion of land to agricultural use and the extraction of firewood and timber (Mickleburgh et al., 2008bt; IUCN, 2009; Mickleburgh et al., 2010). CONSERVATION ACTIONS: Mickleburgh et al. (2010) repeats what stated by Mickleburgh et al. (2008bt) [in IUCN (2009)] who report that there appear to be no direct conservation measures in place for this species. It is not known if the species is present within any protected areas. There is a need to maintain areas of suitable forest within the range of this species. Further studies are needed into the distribution, abundance, breeding biology and general ecology of this species. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) reported 2n = 48, FN = 54, BA = 8, a submetacentric X chromosome and an acrocentric Y chromosome. Protein / allozyme - Unknown. POPULATION: Structure and Density:- It is considered to be a very rare species (Mickleburgh et al., 2008bt; IUCN, 2009; Mickleburgh et al., 2010). Trend:- 2010: Unknown (Mickleburgh et al., 2010). 2008: Unknown (Mickleburgh et al., 2008bt; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Ghana. GENERAL DISTRIBUTION: Mops petersoni has only been recorded from Ghana and Cameroon (with type locality at '15 km S Kumba'), with no records from the intervening countries of Togo, Benin and Nigeria. Koopman (1993a) included "perhaps Sierra Leone" as part of the distribution, however, J. Fahr (pers. comm., 2004 in Mickleburgh et al., 2008bt; IUCN, 2009) indicates that there is no evidence of the species' presence. Native: Cameroon; Ghana. Presence uncertain: Sierra Leone (Koopman, 1993a). Figure 188. Distribution of Mops (Xiphonycteris) petersoni Mops (Xiphonycteris) spurrelli (Dollman, 1911) *1911. Xiphonycteris spurrelli Dollman, Ann. Mag. nat. Hist., ser. 8, 7 (38): 211. Publication date: 1 February 1911. Type locality: Ghana: Western province: 60 mi (90 km) W Kumasi: Bibianaha [=Bibiani] [06 28 N 02 20 W, 720 ft] [Goto Description]. Holotype: BMNH 1911.1.11.1: ad ♂. Collected by: Dr. H.G.F. Spurrell; collection date: 8 December 1910; original number: 34. - Etymology: In honour of dr. H.G.F. Spurrell, who collected the type specimen, and provided many rare and unique West-African mammals to National Collection (see Dollman, 1911: 211). 1973. Xiphonycteris spurelli: Eisentraut, Bonn. zool. Monogr., 3: 47. (Lapsus) ? Mops (Xiphonycteris) spurelli: (Lapsus) ? Mops (Xiphonycteris) spurrelli: (Name Combination, Current Combination) ? Mops spurrelli: (Name Combination) ? Tadarida spurelli: (Lapsus) ? Tadarida spurrelli: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: morous sahelský. English: Spurrell's Mops Bat, Spurrell's Free-tailed Bat. French: African Chiroptera Report 2020 Tadaride de Spurell. Bulldoggfledermaus. German: Spurrells CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008dm; IUCN, 2009; Monadjem et al., 2017ak). Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops spurrelli] (Monadjem et al., 2017ak). 2008: LC ver 3.1 (2001) [assessed as Tadarida spurrelli] (Mickleburgh et al., 2008dm; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cm; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The species is locally threatened by deforestation, often through conversion of land to agricultural use or by extraction of timber and firewood (Mickleburgh et al., 2008dm; IUCN, 2009; Monadjem et al., 2017ak). CONSERVATION ACTIONS: Mickleburgh et al. (2008dm) [in IUCN (2009)] and Monadjem et al. (2017ak) report that there appear to be no direct conservation measures in place for this species, and it is not known if the species is present within any protected areas. There is a need to maintain areas of suitable forest habitat for this species. Additional studies are needed into the distribution of this species. 533 Native: Cameroon; Central African Republic; Congo (The Democratic Republic of the); Côte d'Ivoire; Equatorial Guinea (Bioko); Ghana; Guinea (Fahr and Ebigbo, 2003: 128); Liberia; Sierra Leone; Togo. Presence uncertain: Benin; Congo; Nigeria. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Smith et al. (1986) reported 2n = 48, FN = 64, BA = 18, a submetacentric X chromosome for a female. Protein / allozyme - Unknown. HABITAT: At Mount Nimba, Monadjem et al. (2016y: 368) recorded this species over small streams and swamps at altitudes between 450 and 600 m. POPULATION: Structure and Density:- Within Central Africa this is a very common species and is believed to have quite large colonies (Mickleburgh et al., 2008dm; IUCN, 2009; Monadjem et al., 2017ak). Trend:- 2016: Unknown (Monadjem et al., 2017ak). 2008: Unknown (Mickleburgh et al., 2008dm; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Ghana, Liberia, Sierra Leone, Togo. GENERAL DISTRIBUTION: Mops spurrelli ranges through parts of West and Central Africa. It occurs from Sierra Leone in the west, through Liberia, Guinea, Côte d'Ivoire and Togo, to Cameroon, Equatorial Guinea (Bioko and Rio Muni), the Central African Republic with a record from the eastern part of The Democratic Republic of the Congo. It is possibly present in other countries, such as Benin, Nigeria and Congo, however, this needs to be confirmed. Figure 189. Distribution of Mops (Xiphonycteris) spurrelli Mops (Xiphonycteris) thersites (Thomas, 1903) *1903. Nyctinomus thersites Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (72): 634. Publication date: 1 December 1903. Type locality: Cameroon: Efulen [02 46 N 10 42 E] [Goto Description]. Holotype: BMNH 1904.2.8.4: ad ♂. Collected by: George Latimer Bates 534 ISSN 1990-6471 1917. 1973. 2018. ? ? ? ? ? ? Esq. 3 specimens: see Thomas (1903e: 635). - Etymology: Thersites was one of the members of the Greek army against Troy. He tried to stir up the army against its leaders, for which he got beaten by Odysseus. His name is a scorn: naughty boy. He was an excessive thwaddler, with deformed legs, feet, and bent over shoulders, a pointed head and thin flaky hair. Mops (Allomops) occipitalis J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 474, text-fig. 15. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Avakubi [01 18 N 27 32 E] [Goto Description]. Holotype: AMNH 48851: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 13 February 1914; original number: 2487. Tatarida theresites: Eisentraut, Bonn. zool. Monogr., 3: 113. (Lapsus) Mops therisites: Gunnell and Manthi, J. Hum. Evol., Suppl.. Publication date: 6 April 2018. (Lapsus) Mops (Xiphonycteris) thersites: (Name Combination, Current Combination) Mops (Xyphonycteris) thersites: (Lapsus) Mops thersites: (Name Combination) Tadarida (Mops) thersites: (Name Combination) Tadarida thersites thersites: (Name Combination) Tadarida thersites: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Chinese: 无 畏 犬 吻 蝠 . Czech: morous spílavý. English: Railer Mops Bat, Railer Bat. French: Tadaride de Railer. German: Thersites' Bulldoggfledermaus. Portuguese: Morcego de railer. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Gunnell et al. (2015a: 22) report on a number of humeri from a Pleistocene deposit at Olduvai Gorge (Tanzania), which they assigned to Mops cf. M. thersites. MAJOR THREATS: It is locally threatened by habitat destruction, often through conversion of land to agricultural use or timber extraction (Mickleburgh et al., 2008dn; IUCN, 2009; Monadjem et al., 2017al). CONSERVATION ACTIONS: Mickleburgh et al. (2008dn) [in IUCN (2009)] and Monadjem et al. (2017al) report that there appear to be no direct conservation measures in place for this species. It is not known if the species is present within any protected areas. Additional studies are needed into the distribution of this species. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, tolerance of a degree of habitat modification, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008dn; IUCN, 2009; Monadjem et al., 2017al). GENERAL DISTRIBUTION: Mops thersites has been recorded from much of Western and Central Africa. It ranges from Sierra Leone in the west, through West Africa (where it has been recorded from most countries) to Cameroon, and from here south into southern parts of the Democratic Republic of the Congo, and east into Uganda, Rwanda and western Kenya. It is present on Bioko Island, Equatorial Guinea. Assessment History Global 2016: LC ver 3.1 (2001) [assessed as Mops thersites] (Monadjem et al., 2017al). 2008: LC ver 3.1 (2001) [assessed as Tadarida thersites] (Mickleburgh et al., 2008dn; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cl; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Native: Cameroon; Central African Republic; Congo (The Democratic Republic of the) (Schouteden, 1944; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 544); Côte d'Ivoire; Equatorial Guinea [Bioko]; Gabon; Ghana; Guinea (Fahr et al., 2006b: 245; Denys et al., 2013: 284); Kenya; Liberia (Fahr, 2007a: 104); Nigeria; Rwanda; Sierra Leone; Uganda. Regional None known. Presence uncertain: Mozambique and Zanzibar. African Chiroptera Report 2020 Bates et al. (2013: 339) reject the presence of this species in the Republic of Congo as Malbrant and Maclatchy (1949) only suggested it might occur in that country and subsequent authors did not provide supporting records. MOLECULAR BIOLOGY: Denys et al. (2013: 284) report the karyotype for a single female specimen from the Guinean Mount Nimba area attributed to this species as: 2n = 48, NFa = 70, including four pairs of metacentric, eight pairs of subtelocentric, 11 pairs of acrocentric autosomes and a middle-sized submetacentric X chromosome. HABITAT: This species was captured by Monadjem et al. (2016y: 368) in the Mount Nimba area above water in forested and open habitats between 450 and 880 m. 535 SIPHONAPTERA Ischnopsyllidae: Lagaropsylla oblique Smit, 1957 from Sierra Leone, Cameroon and the Congo (Haeselbarth et al., 1966: 191). VIRUSES: Filoviridae - File viruses Marburgvirus Weiß (2013: 58, 66) found antibodies against MARV in one M. thersites from the Taï National Park (Côte d'Ivoire). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Gabon, Ghana, Guinea, Kenya, Liberia, Nigeria, Rwanda, Sierra Leone, Togo, Uganda. POPULATION: Structure and Density:- It appears to be quite abundant in parts of its range (Mickleburgh et al., 2008dn; IUCN, 2009; Monadjem et al., 2017al). Trend:- 2016: Stable (Monadjem et al., 2017al). 2008: Stable (Mickleburgh et al., 2008dn; IUCN, 2009). PARASITES: Polyctenidae: Hypoctenes clarus Jordan, 1922, from near Kribi, Cameroon (Haeselbarth et al., 1966: 18). Figure 190. Distribution of Mops (Xiphonycteris) thersites Genus Mormopterus Peters, 1865 *1865. Mormopterus Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 258. - Comments: Type species: Nyctinomus (Mormopterus) jugularis Peters, 1865. - Etymology: From the Greek "mormopterus" meaning phantom- or ghost-winged or winged goblin (Flannery, 1990: 367; 1995a: 408). (Current Combination) ? Tadarida (Mormopterus): (Name Combination) TAXONOMY: Meester et al. (1986) state that this treatment of Mormopterus is provisional; it has been regarded as a subgenus of Tadarida (Hayman and Hill, 1971) or as a vaild genus that includes Platymops and Sauromys as subgenera (Freeman, 1981b; Legendre, 1984), and the extralimital Micronomus as either a subgenus (Legendre loc. cit.) or a synonym (Freeman, 1981b; Peterson, 1985). Bronner et al. (2003) mention that the status of Sauromys, described as a monotypic subgenus of the extralimital (to southern Africa) Platymops for the South African flat-headed free-tailed bat, remains unclear. Peterson (1965) raised it to generic rank, a treatment endorsed by many subsequent authors, and corroborated by a limited multivariate analysis of wing bone and cranial characteristics (Peterson, 1985). However, morphometric studies by Freeman (1981b) and Legendre (1984) concluded that it is a subgenus of Mormopterus, a position followed by Koopman (1993a, 1994). Simmons (2005) retains Sauromys as a valid genus in the 3rd Edition of Mammal Species of the World, a treatment we favour because of the unique ecology and 536 ISSN 1990-6471 morphology of S. petrophilus. The separation between Mormopterus and Sauromys is also supported by the analysis of the Rag2 data by Ammerman et al. (2013: 309). Meester et al. (1986) furthermore state that it was originally described as a subgenus, but this genus was regarded as a synonym of Platymops by Shortridge (1942), Roberts (1951) and Ellerman et al. (1953). Harrison (1962) retains it as a subgenus, while Peterson (1965) raises it to generic rank, in which he is followed by Hayman and Hill (1971), Smithers (1971), Corbet and Hill (1980) and Rautenbach (1982). However, Freeman (1981b), followed by Koopman (1982) and Legendre (1984), conclude that both Sauromys and Platymops are subgenera of Mormopterus. We take the view that both Platymops and Sauromys are valid genera. From their molecular study, Lamb et al. (2010: 203; 2011: 10) found that M. jugularis and M. francoismoutoui form a monophyletic clade (dated about 31.2 MYR), whereas a third member of the genus (the New World M. kalinowskii) seems more related to the South American Nyctinomops laticaudatus and Nyctinomops aurispinosus. The analysis of the recombination activating gene 2 (Rag2) sequence data led Ammerman et al. (2013: 307) to conclude that the genus Cheiromeles is the most basal lineage within the Molossinae, and that Mormopterus is a sistergroup of the rest of the Molossinae. Key in Jacobs and Fenton (2002, Mammalian Species, 703). Currently (Simmons and Cirranello, 2020) recognized subgenera and species of the genus Mormopterus: Micronomus: norfolkensis (Gray, 1840) – Norfolk Isl, southeastern Queensland, eastern New South Wales (Australia) (Simmons, 2005: 445). Mormopterus: acetabulous (Hermann, 1804); doriae K. Andersen, 1907 – Sumatra (Indonesia) (Simmons, 2005: 445); francoismoutoui Goodman, Jansen Van Vuuren, Ratrimomanarivo, Probst and Bowie, 2008; jugularis (Peters, 1865); kalinowskii (Thomas, 1893) – Peru, northern Chile (Simmons, 2005: 445); minutus (Miller, 1899) – Cuba (Simmons, 2005: 445); phrudus Handley, 1956 – Peru (Simmons, 2005: 4435). Ozimops Reardon, McKenzie and Adams, 2014: beccarii Peters, 1881 – Molucca Isls, New Guinea, adjacent small islands, northern Australia (Simmons, 2005: 444); cobourgianus D.H. Johnson, 1959 – N Australia; halli Reardon, McKenzie and Adams, 2014 – N Australia; kitcheneri McKenzie, Reardon and Adams, 2014 – S West Australia; loriae Thomas, 1897 – northern Australia; New Guinea (Simmons, 2005: 445); lumsdenae Reardon, McKenzie and Adams, 2014 – N Australia; petersi (Leche, 1884) – Australia; planiceps (Peters, 1866) – southern and central Australia (Simmons, 2005: 446); ridei (Felten, 1964) – E Australia. Setirostris Reardon, McKenzie and Adams, 2014: eleryi Reardon and McKenzie, 2008 – C and E Australia. COMMON NAMES: Czech: Petersovi morousi. English: Phantomwinged Bats, Little Bulldog Bats, Little Mastiff Bats, Little Goblin Bats, Flat-headed Bats, Flat-headed Bulldog Bats, Goblin bats. French: Tadarides. German: Mastino-Fledermäuse. UTILISATION: See Goodman et al. (2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Mormopterus acetabulosus (Hermann, 1804) *1804. V[espertilio] acetabulosus Hermann, Obs. Zool. , 19. Publication date: 31 August 1804. Type locality: Mauritius: Port Louis [Goto Description]. Holotype: MNHN ZM-MO-1984368: ad? ♀. Collected by: M. Desjardins; collection date: 1829; original number: A488. Goodman et al. (2008c: 1324) incidcate that the specimen is labeled as type, but was collected 25 years after Hermann (1804) described it. Neotype: FMNH 187489: Collected by: Steven M. Goodman and V. Florens; collection date: 29 October 2006. Mauritius, Black River District, Palma, Palma Cave, 20 16'40.5''S 57 27'14.7''E. Goodman et al. (2008c: 1327) designated this as the neotype. Cheke (2009b) rejects the assignment of the neotype by Goodman et al. (2008c), based on article 73.1.4 and 75.8 (rules of Nomenclature - ICZN 1999). - Comments: Cheke (2009b: 1) rejects the assignment of the neotype, based on article 73.1.4 (rules of Nomenclature - ICZN 1999), which indicates that Paul Jossigny's drawings made of the specimen that Commerson once had before him in Mauritius should be considered the holotype of Hermann's Vespertilio acetabulosus, and that the designation of a neotype by Goodman et al. (2008c: 1324) is invalidated by article 75.8. The drawings on which Commerson based himself African Chiroptera Report 2020 1820. 1877. ? ? ? 537 (and which were referred to by Hermann, 1804: 19) are to be found amongst the Commerson papers, in the folder of other similar drawings at MS 282(II), labelled No. 51 (Moutou, 1982, pers. obs. 2006) in the archives of the Bibiothèque du Muséum; the folder is described in paragraph 73 of Laissus’s (1974) catalogue. Nyctinomus acetabulosus: Desmarest, Mammalogie, 117. (Name Combination) Nyctinomus [(Mormopterus)] acetabulosus: Dobson, Proc. zool. Soc. Lond., 1876, IV: 734. Publication date: April 1877. (Name Combination) Molossus acetabulosus: Commerson. - Comments: Dobson (1877: 734) refers for this combination to Peters (1869a: 402), and suggests it is only a manuscript name. (Name Combination) Mormopterus acetabulosus: (Name Combination, Current Combination) Tadarida acetabulosus natalensis: (Name Combination) TAXONOMY: See Simmons (2005: 444) and Goodman et al. (2008c). Cheke (2009a: 115) indicates that Jossigny’s drawing used by Hermann to describe the species in 1804 is to be considered as the lectotype of the species, which would overrule the need for a neotype as designated by Goodman et al. (2008c: 1324). COMMON NAMES: Afrikaans: Natalse losstertvlermuis. Czech: morous mauricijský. English: Natal Flat-headed Bat, Natal Wrinkle-lipped Bat, Natal Free-tailed Bat. French: Sauromys du Natal, Tadaride à lèvres ridées, Petit molosse de Port Louis. German: Mauritius Mastino-Fledermaus, Maurizische Doggengrämler. CONSERVATION STATUS: Global Justification Listed as Endangered (EN A2bc; B1ab(iii,v)+2ab(iii,v)) because this species has a restricted range and decline in population size. It occurs in less than five sites on Mauritius. It is restricted to lava tube caves. The population of the species has declined by over 80% during the last 18 years, however the rate of decline over the last three generations is unknown; it is likely that it approached 67% over three generations (11-12 years based on a generation length of 3.9 years; see Pacifici et al., 2013). The major roosting cave system for the species had been developing into a tourism attraction for the past three years, resulting in loss of over 10,000 bats from a single location. Assessment History Global 2016: EN A2bc; B1ab(iii,v)+2ab(iii,v) ver. 3.1 (2001) (Bergmans et al., 2017c). 2014: EN A2b; B1ab(iii)+2ab(iii) ver 3.1 (2001) (Bergmans et al., 2017c). 2008: VU D2 ver 3.1 (2001) (Hutson and Bergmans, 2008; IUCN, 2009). 2004: VU D2 ver 3.1 (2001) (Hutson and Bergmans, 2004; IUCN, 2004). 1996). 1996: VU (Baillie and Groombridge, Regional South Africa:- 2004: considered a vagrant not assessed, known from single locality (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: It is threatened by disturbance of roosting caves, and the removal of protective grills over the caves for landfill purposes Hutson and Bergmans, 2008; IUCN, 2009; Bergmans et al., 2017c). CONSERVATION ACTIONS: Hutson and Bergmans (2008) [in IUCN (2009)] and Bergmans et al. (2017c) report that it is not known if the species is present in any protected areas. There is a need to protect cave roosts of this species in Mauritius by re-grilling important sites, cleaning the caves from household waste, education of local community (Bergmans et al., 2017c). GENERAL DISTRIBUTION: Mormopterus acetabulosus is endemic to the Mascarene Islands (Mauritius) (see Goodman et al., 2008c: 1316; O'Brien, 2011: 288). Records from Réunion should be assigned to Mormopterus francoismoutoui). There is a single doubtful record for Madagascar (Peterson et al., 1995: 150). Taylor (1998: 60) mentioned a specimen from South Africa (Durban) collected by Andrew Smith in 1833, but also states that no further specimens have been collected since. An old record from between the district of Shoa and Lake Rudolph in southern Ethiopia (Hayman and Hill, 1971: 59), most probably should be attributed to M. francoismoutoui (Goodman et al., 2008c: 1325). Furthermore, it is now considered most likely that specimens taken outside of the Mascarene Islands were vagrants. 538 ISSN 1990-6471 Monadjem et al. (2010d) did not retain this species for south Central Africa, but did mention the specimen from Durban, South Africa in the appendix (p. 545). PARASITES: Dietrich et al. (2018a: 3) reported the presence of sequences closely related to the bacterium Leptospira borgpetersenii. Native: Mauritius (Smith, 1833: 54; Meester et al., 1986: 67; Simmons, 2005: 444; Goodman et al., 2008c: 1316). Laudisoit et al. (2012: 739) indicate that the type host for Araeopsylla martialis (Rothschild, 1903) (Siphonaptera; Ischnopsyllidae) was originally mentioned as "Nyctinomus acetabulosus", but this should actually be Mormopterus francoismoutoui. Presence uncertain: Madagascar (Dobson, 1878; Allen, 1939a; Hayman and Hill, 1971; Meester et al., 1986: 67; Peterson et al., 1995: 150; Simmons, 2005: 444); South Africa (Meester et al., 1986: 67; Taylor, 1998: 60; Taylor, 2000: 75; Simmons, 2005: 444; Monadjem et al., 2010d: 545); Sudan (Freeman, 1981b: 75). Presence incorrectly assigned: Ethiopia (Hayman and Hill, 1971: 59; Simmons, 2005: 444; Kaipf et al., 2015: 13); Réunion. ROOST: Middleton (1998a: 91) mention large colonies occurring in two caves in the Plaine des Roches area (Mauritius), and indicate that elsewhere the colonies seldom reach 2,000 animals. POPULATION: Structure and Density:- The largest known day roost colony on La Réunion (at Trois Bassins), with estimates from 25,000 to 10,000 individuals, is abandoned in austral winter, and it is not known where these bats spend the winter (Devaux, 2006; Issartel, 2004; Probst, 2003; Goodman et al., 2008c: 1325). Goodman (2007) found bats allocated to this species to be common at several sites on Mauritius. However, Taylor (2016) [in [Bergmans et al. (2017c)] found the species to be narrowed to only four caves in Mauritius. Based on the survey it has been estimated that the population decreased by over 80% during the last 18 years and it keeps declining. VIRUSES: Paramyxoviridae Wilkinson et al. (2012: 160) tested 55 individuals from the Mauritius using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 1 positive results for viral nucleic acids. Rhabdoviridae Of 31 specimens tested against Lagos bat lyssavirus and Duvenhage lyssavirus by Mélade et al. (2016a: 6), respectively five and six showed a seroreaction. This was also the case for one out of eight bats tested for European bat lyssavirus 1. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Ethiopia, Liberia, Madagascar, Mauritius, South Africa. Trend:- 2016: Decreasing (Bergmans et al., 2017c). 2008: Unknown (Hutson and Bergmans, 2008; IUCN, 2009). Figure 191. Distribution of Mormopterus acetabulosus Mormopterus acetabulosus acetabulosus (Hermann, 1804) *1804. V[espertilio] acetabulosus Hermann, Obs. Zool. , 19. Publication date: 31 August 1804. Type locality: Mauritius: Port Louis [Goto Description]. Holotype: MNHN ZM-MO-1984368: ad? ♀. Collected by: M. Desjardins; collection date: 1829; original number: A488. Goodman et al. (2008c: 1324) incidcate that the specimen is labeled as type, but was collected 25 years after Hermann (1804) described it. Neotype: FMNH 187489: Collected by: Steven M. Goodman and V. Florens; collection date: 29 October 2006. Mauritius, Black River District, Palma, Palma Cave, 20 16'40.5''S 57 27'14.7''E. Goodman et al. (2008c: 1327) designated this as the neotype. Cheke (2009b) rejects the African Chiroptera Report 2020 539 assignment of the neotype by Goodman et al. (2008c), based on article 73.1.4 and 75.8 (rules of Nomenclature - ICZN 1999). - Comments: Cheke (2009b: 1) rejects the assignment of the neotype, based on article 73.1.4 (rules of Nomenclature - ICZN 1999), which indicates that Paul Jossigny's drawings made of the specimen that Commerson once had before him in Mauritius should be considered the holotype of Hermann's Vespertilio acetabulosus, and that the designation of a neotype by Goodman et al. (2008c: 1324) is invalidated by article 75.8. The drawings on which Commerson based himself (and which were referred to by Hermann, 1804: 19) are to be found amongst the Commerson papers, in the folder of other similar drawings at MS 282(II), labelled No. 51 (Moutou, 1982, pers. obs. 2006) in the archives of the Bibiothèque du Muséum; the folder is described in paragraph 73 of Laissus’s (1974) catalogue. 1823-1824. Nyctinomus mauritianus Horsfield, Zoological Research in Java, Part 5, 9th page of text in account of N. tenuis. Publication date: 1823 - 1824. Type locality: Mauritius: Port Louis. ? Mormopterus acetabulosus acetabulosus: (Name Combination, Current Combination) ROOST: On Mauritius, these bats use caves as typical roosting sites (see Wilkinson et al., 2012: 160). VIRUSES: Wilkinson et al. (2012: 160) found one specimen out of 55 examined on Mauritius, that tested positive for Respirovirus / Morbillivirus / Henipahvirus (RMH) and Paramyxovirinae (PMV) DISTRIBUTION BASED ON VOUCHER SPECIMENS: Mauritius, Réunion. Mormopterus acetabulosus natalensis (A. Smith, 1847) 1833. Nyctinomus dubius A. Smith, S. Afr. Quart. J., ser. 2, 1 (2): 54. Publication date: November 1833. Type locality: South Africa: Cape Province and Natal, between [Goto Description]. *1847. Dysopes natalensis A. Smith, Illustrated Zoology of South African Mammals, pl. 49 and text. Publication date: December 1847. Type locality: South Africa: KwaZulu-Natal: Port Natal [=Durban], near [ca. 29 52 S 31 00 E]. - Comments: Dobson (1877: 734) mentions that the original name used by Smith (1847: pl. 49) was Nyctinomus natalensis, but this could not be checked yet. If this isn't the case, then this combination should probably be attributed to Dobson (1877). - Etymology: The name reflects the two specimens collected in KwaZulu-Natal. ? Mormopterus acetabulosus natalensis: (Name Combination, Current Combination) TAXONOMY: Allen (1939a) tentatively treats dubius as a species of Nyctinomus (= Tadarida), remarking that its status is uncertain; Roberts (1951), without giving reasons, regards it as a prior name for natalensis. Ellerman et al. (1953) are doubtful whether or not the two names are synonymous, and although they list dubius in the synonymy of natalensis they suggest that it may be necessary to discard the name as not being certainly identifiable. Hayman and Hill (1971: 65) list dubius as incertae sedis. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". Mormopterus francoismoutoui Goodman, Jansen Van Vuuren, Ratrimomanarivo, Probst, Bowie, 2008 *2008. Mormopterus francoismoutoui Goodman, Jansen Van Vuuren, Ratrimomanarivo, Probst and Bowie, J. Mamm., 89 (5): 1318, figs. 2 - 6. Type locality: Réunion: Commune de La Possession: Pont de Balthazar [2056.732S 5519.848E, 40m] [Goto Description]. Holotype: FMNH 193986: ad ♂, skull and alcoholic. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006; original number: SMG 15310. Paratype: FMNH 193982: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193983: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193984: ♂, skull only. Collected by: Steven M. 540 ISSN 1990-6471 Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193985: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193987: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193988: ♂, skull only. Collected by: Steven M. Goodman and JeanMichel Probst; collection date: 25 September 2006. Paratype: FMNH 193989: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193990: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193991: ♀, alcoholic (skull not removed). Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 25 September 2006. Paratype: FMNH 193992: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 193993: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 193994: ♂, skull only. Collected by: Steven M. Goodman and JeanMichel Probst; collection date: 26 September 2006. Paratype: FMNH 193995: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 193996: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 193997: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 193998: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 193999: ♀, skull only. Collected by: Steven M. Goodman and JeanMichel Probst; collection date: 26 September 2006. Paratype: FMNH 194000: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194001: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194002: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194003: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194004: ♀, skull only. Collected by: Steven M. Goodman and JeanMichel Probst; collection date: 26 September 2006. Paratype: FMNH 194005: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194006: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194007: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194008: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194009: ♂, skull only. Collected by: Steven M. Goodman and JeanMichel Probst; collection date: 26 September 2006. Paratype: FMNH 194010: ♂, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194011: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194012: ♀, skull only. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194013: complete skeleton. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194014: complete skeleton. Collected by: Steven M. Goodman and Jean-Michel Probst; collection date: 26 September 2006. Paratype: FMNH 194015: ♂, complete skeleton. Collected by: Steven M. Goodman. Paratype: FMNH 194016: ♀, complete skeleton. Collected by: Steven M. Goodman. Paratype: FMNH 74208: ♂, alcoholic (skull not removed). Collected by: G. Morel. Paratype: FMNH 74209: ♀, alcoholic (skull not removed). Collected by: G. Morel. Paratype: MNHN ZM-MO-1862362: Collected by: M. Millard. Paratype: MNHN ZM-MO-1865-24: Collected by: M. Lantz. Paratype: MNHN ZM-MO-1865-25: Collected by: M. Lantz. Paratype: MNHN ZM-MO-1865-26: Collected by: M. Lantz. Paratype: MNHN ZM-MO-1981-301: Collected by: Dr. Véto and F. Moutou; collection date: 23 March 1979. Paratype: MNHN ZM-MO-1981-302: Collected by: Dr. Véto and F. Moutou; collection date: 1 August 1980. Paratype: MNHN ZM-MO-1981-303: Collected by: Dr. Véto and F. Moutou; collection date: 1 August 1980. Paratype: MNHN ZM-MO-1981-304: Collected by: Dr. Véto and F. Moutou; collection date: 1 August 1980. Paratype: MNHN ZM-MO-1981-305: African Chiroptera Report 2020 541 Collected by: Dr. Véto and F. Moutou; collection date: 1 August 1980. Paratype: MNHN ZM-MO-1981-306: Collected by: Dr. Véto and F. Moutou; collection date: 1 August 1980. Paratype: MNHN ZM-MO-1981-307: Collected by: Dr. Véto and F. Moutou; collection date: 1 August 1980. Paratype: UMMZ 115813: Collected by: Frank B. Gill; collection date: 2 June 1967. Paratype: UMMZ 115814: Collected by: Frank B. Gill; collection date: 2 June 1967. Paratype: UMMZ 115815: Collected by: Frank B. Gill; collection date: 2 June 1967. Paratype: UMMZ 115816: Collected by: Frank B. Gill; collection date: 2 June 1967. Paratype: UMMZ 115817: ♀. Collected by: Frank B. Gill; collection date: 2 June 1967. Paratype: UMMZ 115818: Collected by: Frank B. Gill; collection date: 2 June 1967. Paratype: UMMZ 115819: Collected by: Frank B. Gill; collection date: 2 June 1967. - Etymology: The name francoismoutoui is a patronym in honour of Dr. François Moutou of the Unité d'Epidémiologie, Agence Française de Sécurité Sanitaire des Aliments, Maison Alfort, France. Dr. Moutou has made important contributions to the vertebrate fauna of the Mascarene Islands, particularly La Réunion (Goodman et al., 2008c: 1319). (Current Combination) TAXONOMY: Formerly identified as M. acetabulosus see Goodman et al. (2008c: 1316). COMMON NAMES: Czech: morous réunionský. English: Reunion Free-tailed bat. French: Petit molosse, Tadaride de La Réunion. German: Reunion MastinoFledermaus. CONSERVATION STATUS: Global Justification Goodman et al. (2008c: 1325) state it is under no serious threat of population decline in the near future. Augros et al. (2016: 372) refer to IUCN (2010) and mention "LC" as status. Goodman (2017f) report that this species is known from numerous sites on La Réunion, which include large maternity and day roosting colonies in caves and numerous synanthropique roost sites, such as under bridges and in houses, churches, and the principal airport (Dietrich et al., 2015b). No clear threats are known. Assessment History Global 2016: LC ver. 3.1 (2001) (Goodman, 2017f). Regional None known. MAJOR THREATS: No threats identified by Goodman (2017f). CONSERVATION ACTIONS: On the basis of territorial legislation, all bats on La Réunion are protected by law (Arrêté of 17 April 1981 and of 17 February 1989) (Goodman et al., 2008c: 1325). More details are needed on the ecology, bioacoustics, and dispersal patterns of La Réunion bats (Goodman, 2017f). GENERAL DISTRIBUTION: Mormopterus francoismoutoui has been recorded from a variety of localities on La Réunion from sea level to about 2,000 m (Moutou, 1982; Goodman et al., 2008c). Native: Réunion (Goodman et al., 2008c: 1316). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Mormopterus francoismoutoui is a relatively large member of the genus Mormopterus. Its dorsal body fur is short and dense with a rich dark brown color and its ventrum is slightly paler. Wing membrane and uropatagium are brownish black and show no noticeble change in pigmentation across their surface area. In adult males, there is a large glandular sac on the ventral portion of the lower neck; this gland is absent in females (Goodman et al. (2008c). An albino specimen was reported by Ramasindrazana et al. (2014: 103) and Lucati and López-Baucells (2016: Suppl.). DENTAL FORMULA: I 1/3, C 1/1, P 1/2, M 3/3 = 30 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Goodman et al. (2008c): The skull of M. francoismoutoui, as with most other members of this genus, is not noticebly flattened and has a long rostrum and well-developed lachrymal tubercles. The postorbital processes are well developed and distinctly shaped. The palatal emargination is well developed, similar to character state 4 for variable C65 of Freeman (1981b). Basisphenoid pits are present, but are not prominent, and there is a pronounced extension of the palatal spine descending onto the palatine bone. 542 ISSN 1990-6471 ECHOLOCATION: Barataud and Giosa (2013: 150) analyzed 63 sequences from M. francoismoutoui specimens on Réunion and found QFC = 77 kHz and FM = 176 kHz MOLECULAR BIOLOGY: Dietrich et al. (2019: 6) developed 12 polymorphic molecular markers to facilitate further investigations into population genetics of this species. HABITS: The holotype and associated specimens roosting in the expansion joint of a bridge were observed to roost in a vertical position (head down) in very narrow crevices (Goodman et al., 2008c: 1319). ROOST: Found in a variety of different roost sites, ranging from natural caves to synanthropic situations (Goodman et al., 2008c). The holotype and associated specimens were extracted by hand from the expansion fissures in the lower portion of a concrete bridge (Goodman et al., 2008c). Moutou (1982) found the species to roost in the crevices found in the rock faces around the island. It is also associated with a variety of human structures, including open roofs, roof slats, and window shutters (Goodman et al., 2008c). Augros et al. (2016: 374) provide an overview of 112 roosts, of which 74 (66 %) were artificial. MIGRATION: The bats abandon Trois Bassins (with estimates of 10,000 to 25,000 individuals) during the austral winter, and it is not known where these bats spend the winter (Probst, 2003; Issartel, 2004; Devaux, 2006; Goodman et al., 2008c). PARASITES: Laudisoit et al. (2012: 739) indicate that M. francoismoutoui is actually the type host for Araeopsylla martialis (Rothschild, 1903) (Syphonaptera; Ischnopsyllidae) and not "Nyctinomus acetabulosus". Dietrich et al. (2015b: 4280) found similar infection dynamics for Leptospira bacteria and paramyxoviruses, which showed peak infections during late pregnancy and two months after the initial birth period. Both pathogens, however, did not seem to interact with one another. Guernier et al. (2016: 1) found M. francoismoutoui on Réunion to be infected by Leptospira borgpetersenii bacteria, which also infect humans. However, these bacteria had different haplotypes, indicating that the bats do not have any role in human infections. Dietrich et al. (2018a: 2) mention the bacterium to be closely related to L. borgpetersenii. VIRUSES: Paramyxoviridae Nieto-Rabiela et al. (2019: Suppl.) reported Bat paramyxovirus to be present. Rhabdoviridae A large amount of bats from La Réunion were tested for lyssaviruses by Mélade et al. (2016a: 6), which resulted in the following numbers: Lagos bat lyssavirus: 3 seroreactions for 121 bats; Duvenhage lyssavirus: 14/117, and European bat lyssavirus 1: 9/82. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Réunion. POPULATION: Structure and Density:- The largest known day roost colony on La Réunion (at Trois Bassins), with estimates from 10,000 to 25,000 individuals, is abandoned in austral winter, and it is not known where these bats spend the winter (Devaux, 2006; Issartel, 2004; Probst, 2003; Goodman et al., 2008c: 1325) [as M. acetabulosus]. While Goodman (2017f) report that this species is not correctly estimated but presumably substantial, with one maternity cave-dwelling colony estimated at 66,500 animals (Ramasindrazana et al., 2014). In urban areas across the island, considerable numbers can be seen foraging at night around outside lighting (Goodman, 2017f). Trend:- 2016: Increasing (Goodman, 2017f). 2008: Unknown (Hutson and Bergmans, 2008; IUCN, 2009). Figure 192. Distribution of Mormopterus francoismoutoui African Chiroptera Report 2020 543 Mormopterus jugularis (Peters, 1865) *1865. Nyctinomus (Mormopterus) jugularis Peters, in: Sclater, Proc. zool. Soc. Lond., 1865, II: 468. Publication date: October 1865. Type locality: Madagascar: Antananarivo (formerly Tananarive) [18 55 S 47 31 E]. Holotype: ZMB 2977: ad ♂, skull and alcoholic. Collected by: H.R. Caldwell. Presented/Donated by: William Lutley Sclater. See Turni and Kock (2008: 70). 1877. Nyctinomus [(Mormopterus)] albiventer Dobson, Proc. zool. Soc. Lond., 1876, IV: 733. Publication date: April 1877. Type locality: Madagascar: "Madagascar" [Goto Description]. Holotype: BMNH 1863.12.26.1: ♀. Presented/Donated by: ?: Collector Unknown. - Comments: Type: BMNH ???, ad ♀, alcoholic, purchased. ? Mormopterus albiventer: ? Mormopterus jugularis: (Name Combination, Current Combination) TAXONOMY: See Simmons (2005: 445). COMMON NAMES: Czech: morous Petersův. English: Peters's Goblin bat, Peters's Wrinkle-lipped Bat, Peters' Wrinkle-lipped Bat. French: Sauromys à jugulaire, Sauromys de Peters, Tadaride de Madagascar. German: Madagaskar MastinoFledermaus. Japanese: Pita-ohiki-koumori. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Muldoon et al. (2009: 1114) report on subfossil remains from the Ankilitelo fauna (late Holocene, appr. 500 years ago). Gunnell et al. (2014: 3) refer to fossil material from Tsimanampetsotsa (one tooth and a few pieces of a forelimb according to Sabatier and Legendre (1985: 23)), Ankilitelo, and Andrahomana on Madagascar. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread distribution across Madagascar and because it is not thought to be facing any major threats across its range (Andriafidison et al., 2008g; IUCN, 2009; Monadjem et al., 2017cb). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cb). 2008: LC ver 3.1 (2001) (Andriafidison et al., 2008g; IUCN, 2009). 1996: VU A2c ver 2.3 (1994) (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: This species is not threatened by loss of native forests because it forages in open areas associated with agriculture. It is locally persecuted when roosting in buildings although this is not yet a major threat. Some colonies are subject to hunting and although this is though to have resulted in abandonment in at least two cases (Goodman et al., 2008d), harvest levels do not appear a major threat (Andriafidison et al., 2008g; IUCN, 2009; Monadjem et al., 2017cb). Fernández-Llamazares et al. (2018: 273) indicate that M. jugularis is often hunted in sacred caves in and around the Tsimanampetsotsa National Park, although in the park, it is generally restricted to periods of famine, e.g. in 2007, when the bats fed 200 people for two months. CONSERVATION ACTIONS: Andriafidison et al. (2008g) [in IUCN (2009)] and Monadjem et al. (2017cb) report that as this species is more commonly found roosting in buildings than natural settings it is not found in many of the protected areas in Madagascar (Goodman et al., 2005a). However, there is no need for any conservation measures at present as this species appears to be widely distributed and is locally abundant. GENERAL DISTRIBUTION: Mormopterus jugularis is endemic to Madagascar (Peterson et al., 1995) and very widespread across the island with a known distribution extending from the coast up to the high plateau at 1,400 m in Anjozorobe (J. Ranivo pers. comm., in Andriafidison et al., 2008g) in IUCN, 2009). Native: Madagascar (Peterson et al., 1995; Simmons, 2005: 445) DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 62) describe the baculum of this species as proportionately broad at the proximal, rounded base and tapering to a rounded distal tip. In lateral view, the structure has a slight curve to the 544 ISSN 1990-6471 main shaft; length: 0.69 ± 0.015 (0.68 - 0.71) mm, width: 0.24 ± 0.016 (0.22 - 0.26) mm. MOLECULAR BIOLOGY: DNA - Unknown Karyotype - Richards et al. (2010) reported one male with a 2n=48 and a Fna=54. They found one large metacentric, three medium-sized metacentric and 19 acrocentric autosomal pairs. The X chromosome is submetacentric and the Y chromosome is metacentric. FISH using Myotis probes was also applied. Trend:- 2016: Unknown (Monadjem et al., 2017cb). 2008: Unknown (Andriafidison et al., 2008g; IUCN, 2009). PARASITES: Lagadec et al. (2012: 1696) and Gomard et al. (2016: 5) report the presence of Leptospira sp bacteria in this species, which, according to Dietrich et al. (2018a: 3), were having sequences closely related to L. borgpetersernii HABITAT: Dammhahn and Goodman (2013: 108) indicate that this species' foraging habitat consists of open areas and the area above the forest canopy. Laudisoit et al. (2012: 739) refer to Lumaret (1962), who reported the present of Araeopsylla martialis (Rothschild, 1903) (Siphonaptera; Ischnopsyllidae) on "Tadarida albiventer" [= M. jugularis]. This flea was also reported by Hastriter (2016: 15). ROOST: Synantropic structures are the typical roosting sites for this species on Madagascar (see Wilkinson et al., 2012: 160). Durette-Desset and Chabaud (1975: 10) described the Nematode Molinostrongylus richardae from a "Mormopterus albiventer" from Anjiro, Madagascar. MIGRATION: Reher et al. (2019: 121) found that M. jugularis was only present in a southwestern Madagascan cave during the rainy season, whereas other species such as Triaenops menamena and Miniopterus mahafaliensis were present in both the dry and wet season. VIRUSES: Coronaviridae Joffrin et al. (2020: 5) reported an Alphacoronavirus from a bat vrom Madagascar. DIET: Kemp et al. (2018: Suppl.) used DNA metacarcoding to detect insect pest species in the diet of these bats and found the following prey orders (in descending order): Coleoptera (Xyleborus ferrugineus (Fabricius, 1801)), Ephemeroptera, Diptera (Simulium lineatum (Meigen, 1804), Culex annulioris Theobald, 1901, Culex MBI-27, Anopheles squamosus Theobald, 1901), Lepidoptera (Meyrickiella homosema (Meyrick, 1887), Herpetogramma licarsisalis (Walker, 1859)), Blattodea, Dermaptera, Hemiptera, Neuroptera, Orthoptera, Sarcoptiformes, Astigmata, Hymenoptera PREDATORS: Goodman et al. (2015c: 78) found the remains of three individuals in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. POPULATION: Structure and Density:- There are no quantitative data available on populations of this species; however, it is assumed to have a high population size because it is widespread and roosts in large colonies in buildings (Andriafidison et al., 2008g; IUCN, 2009; Monadjem et al., 2017cb). Flaviviridae Flavivirus Reynes et al. (2011: 3) reported the isolation of Dakar bat virus from M. jugularis. Paramyxoviridae Wilkinson et al. (2012: 160) tested 13 individuals from the Mauritius using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Wilkinson et al. (2014) tested 19 individuals in Madagascar with an RT-PCR specific for the Respiro-, Morbilli- and Henipavirus genera, two of the 19 individuals tested positive for paramyxovirus RNA. 12 out of 152 Madagascan specimens tested by Mélade et al. (2016b: 4) were positive for paramyxoviruses. Rhabdoviridae 58 specimens were tested by Mélade et al. (2016a: 6), which laid to three seroreactive results for Lagos bat lyssavirus and 17 for Duvenhage lyssavirus. UTILISATION: See Goodman et al. (2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar. African Chiroptera Report 2020 545 Figure 193. Distribution of Mormopterus jugularis Genus Myopterus E. Geoffroy St.-Hilaire, 1818 *1818. Myopterus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 113. Publication date: 1818 [Goto Description]. - Comments: Type species: Myopterus senegalensis Oken, 1816 (not available) (=Myopteris daubentonii Desmarest, 1820). - Etymology: From the Greek "μύξ" or éμυόξ", meaning mouse and "πτέρυξ", meaning wing (see Palmer, 1904: 440). (Current Combination) 1820. Myopteris Desmarest, Encyclopédie Méthodique, (Zoologie, Mammalogie), 1: 131. Publication date: 1820. - Comments: Type: Myopteris daubentonii Desmarest, 1820. 1837. Myoptera de Blainville, C. R. séances Acad. Sci., Paris, 5: 815. Publication date: 1837. 1905. Eomops Thomas, Ann. Mag. nat. Hist., ser. 7, 16 (95): 572. Publication date: 1 November 1905 [Goto Description]. - Comments: Type species: Mormopterus whitleyi Scharff, 1900. TAXONOMY: Includes Eomops; see Hayman and Hill (1971: 56). Grubb et al. (1998: 97) indicate that Myopterus Oken, 1816, Lehrbuch Naturgesch. Pt 3, Zool. Sect. 2: 952, type species Myopterus senegalensis Oken (=Myopteris daubentonii Desmarest, 1820), is not available according to Opinion 417 of the ICZN. Grubb et al. (1998: 97) indicate that Mahoney and Walton (1988a: 116) considered the date of Geoffroy's publication to be 1813. Gardner and Hayssen (2004: 12) mention that the correct date should be 1818, not 1813 as mentioned on the publication itself. Currently (Simmons and Cirranello, 2020) recognized species of the genus Myopterus: daubentonii Desmarest, 1820; whitleyi (Scharff, 1900). COMMON NAMES: Czech: krysoví morousi, spaky. English: Wingedmouse Bats, African Free-tailed Bats, Winged-rat Free-tailed Bats. French: Myoptères. German: Pergamentflügel-Fledermäuse. Myopterus daubentonii Desmarest, 1820 1816. Myopterus senegalensis Oken, Lehrbuch Naturgeschichte, Jena, 3 (2): 952. Type locality: Senegal. - Comments: Not available (Opinion 417 of the ICZN) see Grubb et al. (1998: 97). *1820. Myopterus daubentonii Desmarest, Encyclopédie Méthodique, (Zoologie, Mammalogie), 1: 132. Publication date: 1820. Type locality: Senegal: "Senegal". Holotype: [Unknown] lost:. See Koopman (1993a: 238). - Comments: Type specimen lost (see Koopman, 1993a: 238). - Etymology: In honour of the French naturalist Louis-Jean-Marie d'Aubenton [=Daubenton] (1716 - 1799/1800) (see Bogdanowicz, 1994: 5; Kozhurina, 2002: 15). (Current Combination) 546 ISSN 1990-6471 1915. ? Myopterus albatus Thomas, Ann. Mag. nat. Hist., ser. 8, 16 (96): 469. Publication date: 1 December 1915. Type locality: Congo (Democratic Republic of the): Uele River [04 09 N 22 26 E] [Goto Description]. Holotype: BMNH 1919.5.9.1: Collected by: M. Hutereau; original number: 17. Congo Museum o. 2911. Myopterus daubentonii albatus: (Name Combination) TAXONOMY: Koopman (1989a) and Simmons (2005) recognise albatus Thomas, 1915 as a subspecies. Allen (1939a: 110) used Myopterus senegalensis Oken, 1816 for this species. COMMON NAMES: Czech: morous krysový, spaka krysowá. English: Daubenton's Winged-mouse Bat, Daubenton's Free-tailed Bat, Senegal Mastiff Bat. French: Myoptère de Daubenton, Myoptère du Sénégal. German: Daubentons PergamentflügelFledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since, although it has been recorded over a very wide area, there are only very few records, and very little is known with certainty about its distribution, threats and habitat requirements (Mickleburgh et al., 2008bc; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008bc; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004ca; IUCN, 2004). 1996: DD (Baillie and Groombridge, 1996). Republic and northern Democratic Republic of the Congo. It is generally a lowland species, but has been found up to 1,250 m asl. Native: Central African Republic; Congo (The Democratic Republic of the); Côte d'Ivoire; Senegal. ROOST: Brosset and Vuattoux (1968: 85) reported five specimens from Côte d'Ivoire (as M. senegalensis), that were captured in a hollow trunk of a Borassus palm (Borassus aethiopum). The tree was standing in a savanna area, but only a few hundred meters from a galery forest. The bats were roosting at an altitude of about 12 meters. POPULATION: Structure and Density:- This appears to be a rare species, known from very few specimens (Mickleburgh et al., 2008bc; IUCN, 2009). Trend:- 2008: Unknown (Mickleburgh et al., 2008bc; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Senegal. Regional None known. MAJOR THREATS: The species is generally threatened by habitat loss, especially the removal of potential roost trees (Mickleburgh et al., 2008bc; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008bc) [in IUCN (2009)] report that there appear to be no direct conservation measures in place, and it is not known if the species is present within any protected areas. Further studies are needed into the distribution, natural history and threats to this bat. GENERAL DISTRIBUTION: Myopterus daubentonii is patchily recorded from eight localities in West and Central Africa. It has been reported from Côte d'Ivoire, Central African Figure 194. Distribution of Myopterus daubentonii African Chiroptera Report 2020 547 Myopterus whitleyi (Scharff, 1900) *1900. Mormopterus Whitleyi Scharff, Ann. Mag. nat. Hist., ser. 7, 6 (36): 569. Publication date: 1 December 1900. Type locality: Nigeria: Mid-Western Region, Nikrowa Forest Reserve, Sapoba: Benin City [06 19 N 05 41 E] [Goto Description]. Holotype: BMNH 1900.10.26.1: ad ♂, skull and alcoholic. Presented/Donated by: ?: Collector Unknown. ? Myopterus whitleyi: (Name Combination, Current Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: morous západoafrický. English: Bini Winged-mouse Bat, Whitley's Winged-mouse Bat, Bini Free-tailed Bat. French: Myoptère de Whitley, Myoptère de Bini. German: Whitleys Pergamentflügel-Fledermaus, Bini Pergamentflügel-Fledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008bd; IUCN, 2009; Monadjem et al., 2017ba). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ba). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008bd; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004cf; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). GENERAL DISTRIBUTION: Myopterus whitleyi is distributed in West and Central Africa. It has been recorded from Ghana in the west, through parts of Nigeria, Cameroon, Gabon, and the Democratic Republic of the Congo to western Uganda in the east. Native: Cameroon; Central African Republic (Barrière et al., 2002: 234); Congo (The Democratic Republic of the) (Hayman et al., 1966; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 545); Gabon; Ghana; Nigeria; Uganda. POPULATION: Structure and Density:- It is generally considered to be a rare species. It is found roosting singly or in small numbers (Mickleburgh et al., 2008bd; IUCN, 2009; Monadjem et al., 2017ba). Trend:- 2016: Decreasing (Monadjem et al., 2017ba). 2008: Decreasing (Mickleburgh et al., 2008bd; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Central African Republic, Congo (Democratic Republic of the), Gabon, Ghana, Nigeria, Uganda. Regional None known. MAJOR THREATS: This species is presumably threatened by general conversion of forest to agricultural land and logging activities (Mickleburgh et al., 2008bd; IUCN, 2009; Monadjem et al., 2017ba). CONSERVATION ACTIONS: Mickleburgh et al. (2008bd) [in IUCN (2009)] and Monadjem et al. (2017ba) report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further research is needed into the species distribution, natural history, and adaptability to habitat degradation. Figure 195. Distribution of Myopterus whitleyi 548 ISSN 1990-6471 Genus Otomops Thomas, 1913 *1913. Otomops Thomas, J. Bomb. nat. Hist. Soc., 22: 91. Publication date: April 1913. Comments: Type species: Nyctinomus wroughtoni Thomas, 1913. - Etymology: From the Greek substantive "οὖς" (genitive "ώτός", "ûs", "ōtós"), meaning "ear" and the masculine German substantive Mops, as the genus has outstandingly large ears (see Lanza et al., 2015: 309). (Current Combination) TAXONOMY: Richards et al. (2012: 910) suggest that three distinct taxa might be recognized in Africa: Otomops madagascariensis from Madagascar; O. martiensseni s.s. from southern, eastern, central, and western Africa; and an "undescribed taxon" (now identified as O. harrisoni) from north-east Africa and the Arabian Peninsula. Currently (Simmons and Cirranello, 2020) recognized species of the genus Otomops: formosus Chasen, 1939 – Java (Simmons, 2005: 447); harrisoni Ralph, Richards, Taylor, Napier, and Lamb, 2015; johnstonei Kitchener, How and Maryanto, 1992 – Alor Isl (Indonesia) (Simmons, 2005: 447); madagascariensis Dorst, 1953; martiensseni (Matschie, 1897); papuensis Lawrence, 1948 – southeastern New Guinea (Simmons, 2005: 448); secundus Hayman, 1952 – northeastern New Guinea (Simmons, 2005: 448); wroughtoni (Thomas, 1913) – southern and northeastern India, Cambodia (Simmons, 2005: 448). Bats. French: Molosses à grandes oreilles. German: Riesen-Bulldoggfledermäuse. Italian: Otocarlìno. ETYMOLOGY OF COMMON NAME: So named because they have outstanding large round-pointed ears that are attached along the nose and lie in the plane of the face (see Taylor, 2005). BIOGEOGRAPHY: Lamb et al. (2008) suggests that African and Oriental Otomops lineages diverged around 4.2 Myr, while Malagasy and African lineages split around 1.5 Mya. MOLECULAR BIOLOGY: Patterson et al. (2018: 7) found the average interspecific genetic distance (uncorrected cyt-b pdistance) between O. harrisoni and O. martiensseni to be 2.5 %. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burundi, Kenya, Rwanda. COMMON NAMES: Czech: velkouší morousi. English: Big-eared Bulldog Bats, Big-eared Free-tailed Bats, Mastiff Otomops harrisoni Ralph, Richards, Taylor, Napier and Lamb, 2015 *2015. Otomops harrisoni Ralph, Richards, Taylor, Napier and Lamb, Zootaxa, 4057 (1): 1, 18, figs 8 - 10. Publication date: 9 December 2015. Type locality: Ethiopia: Bale district: Sof Omar Cave [06 54 N 40 48E, 1340 m] [Goto Description]. Holotype: HZM 60.36217: ad ♂, skull and alcoholic. Collected by: Paul J.J. Bates, Malcolm J. Pearch and O. Nurhussein; collection date: 19 July 1998; original number: A51. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 40.31315: ad ♂. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 41.31316: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 42.31317: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 43.31318: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 44.31328: ad ♂. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 56.36213: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 57.36214: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 july 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 58.36215: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 59.36216: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. African Chiroptera Report 2020 ? 549 Paratype: HZM 61.36218: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 62.36219: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Ralph et al. (2015: 19) mention "(also DM 14750)". Paratype: HZM 63.36220: ad ♀. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. Paratype: HZM 64.36221: ad ♂. Collected by: ?: Collector Unknown; collection date: 19 July 1998. Presented/Donated by: ?: Collector Unknown. - Etymology: In honour of Dr. David L. Harrison (1926 - 2015), who's numerous publications on Afro-Arabian Chiroptera, in particular the Molossidae, have significantly improved our knowledge of this poorly known family (Ralph et al. (2015: 20). Otomops martiensseni: COMMON NAMES: Chinese: 哈 氏 巨 犬 吻 蝠 . English: Harrison’s large-eared giant mastiff bat. German: HarrisonRiesen-Bulldoggfledermaus SIMILAR SPECIES: Ralph et al. (2015: 25) report that O. harrisoni is in general larger than other Otomops species. CONSERVATION STATUS: Patterson et al. (2018: 11) report the status of O. harrisoni as Vulnerable. GENERAL DISTRIBUTION: Native: Djibouti; Eritrea (Kock and Zinner, 2004: 3 as O. martiensseni); Ethiopia (in a disused railway tunnel 8 km east of Asmara) for the first time (Kock and Zinner, 2004: 3 as O. martiensseni); Kenya (Patterson et al., 2018: 1), Rwanda (Patterson et al., 2018: 1) and Yemen (Hud Sawa cave, ArRayadi Al-Gharbi Mountains) on the Arabian Peninsula (Al-Jumaily, 1999; Hutson et al., 2001 as O. martiensseni). Ralph et al. (2015: 27) suggest that the range might include additional localities in Uganda, northern areas of Tanzania and southern areas of Somalia. Patterson et al. (2018: 1) found both O. martiensseni and O. harrisoni in a cave in the Musanze district, Rwanda. As they found both species in Kenya, they also caution the identifications of Otomops specimens from that country, especially those from Kwale, Lake Baringo, Nairobi, Naivasha, and Wei-Wei River (p. 11). DENTAL FORMULA: The dental formula of O. harrisoni is I 1/2, C 1/1, P 2/2, M 3/3, characteristic of molossid bats (see Ralph et al., 2015: 26). ECHOLOCATION: Ralph et al. (2015: 26) indicate that Taylor et al. (2005) reported the following values for O. martiensseni specimens from Bungule, Kenya: Fpeak: 12.00 kHz, Fmin: 10.50 kHz, Fmax: 50 kHz, and duration of 9.00 msec. However, since this locality is equidistant from localities were either O. martiensseni or O. harrisoni was found, the taxonomic assignment of the specimens involved remains uncertain. Patterson et al. (2018: Suppl.) recorded the following parameters for 10 Kenyan specimens identified as "O. cf. harrisoni: Fmax: 13.99 ± 2.93 (9.52 - 17.95) kHz, Fstart: 19.35 ± 3.50 (10.72 21.90) kHz, Fend: 11.58 ± 1.17 (9.63 - 13.13) kHz, duration: 10.13 ± 2.46 (7.04 - 15.16) msec. However, they also found (p. 7) differences between O. harrisoni populations from two localities in Kenya: Chyulu Hills: Fpeak: 17.18 ± 0.42 (16.40 - 17.95) kHz, Fmax: 21.67 ± 0.10 (21.48 21.90) kHz, Fmin: 12.53 ± 0.29 (11.80 - 13.13) kHz and call duration: 10.72 ± 1.10 (7.57 - 12.67) msec; and Mount Suswa: Fpeak: 11.86 ± 0.72 (9.52 13.80) kHz, Fmax: 17.81 ± 1.70 (10.72 - 21.62) kHz, Fmin: 110.94 ± 0.47 (9.63 - 12.80) kHz and call duration: 9.74 ± 1.21 (7.04 - 15.16) msec. MOLECULAR BIOLOGY: Karyotype: Warner et al. (1974) reported 2n = 48, FN = 58, BA = 12, a submetacentric X chromosome and a submetacentric Y chromosome for one male O. martiensseni specimen from Kenya, which differed from previously reported karyotypes. It is therefore possible that this male might have been a representative of O. harrisoni. HABITAT: Ralph et al. (2015: 27) mention that these bats are found at high altitudes (above 1,000 m) characterised by relatively drier climates (< 500 mm annual precipitation), including warm semiarid, tropical savanna, warm desert and temperate oceanic climates (e.g. the Ethiopian highlands). 550 ISSN 1990-6471 ROOST: Ralph et al. (2015: 27) indicate that individuals roost predominantly in mountain-associated cave systems and lava caves. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Djibouti, Ethiopia, Kenya. DIET: Feacal data from Ethiopia indicate that medium (size range: 1 - 5 cm) to large (wing span size range: 2.5 - 30 cm) Lepidoptera (Noctuidae, Geometridae and Saturniidae) form 97 % by volume of this bat's diet (see Ralph et al., 2015: 26). PARASITES: At Mount Suswa (Kenya), Patterson et al. (2018: 10, Suppl.) found the following ectoparasites: Brachytarsina alluaudi (Falcoz, 1923) [Diptera, Hippoboscidae], Hypoctenes clarus (Jordan, 1922) [Hemiptera, Polyctenidae], Lagaropsylla sp. [Siphonaptera, Ischnopsyllidae], and some unidentified Argasidae and Macronyssidae. Figure 196. Distribution of Otomops harrisoni Otomops madagascariensis Dorst, 1953 *1953. Otomops madagascariensis Dorst, Mém. Inst. Scient. Madagascar, (A) 8: 236. Type locality: Madagascar: S of Soalala: Namoroka, Réserve Naturelle: Cave P. Saboureau, No. 8 [16 23 S 45 28 E]. Holotype: MNHN ZM-MO-1953-1590: ad ♀, skull and alcoholic. Collected by: Renaud Paulian; collection date: September 1952. See Peterson et al. (1995: 181), Richards et al. (2012: 912). - Etymology: Referring to the island where this taxon occurs. (Current Combination) 2000. Otomops martiensseni madagascarensis: Horácek, Hanák and Gaisler, Bats of the Palearctic region. (Lapsus) ? Otomops madagascariensis madagascariensis: (Name Combination) ? Otomops martiensseni madagascariensis: (Name Combination) TAXONOMY: Previously recognised as a subspecies or synonym of Otomops martiensseni (Koopman, 1993a: 239). Valdivieso et al. (1979) listed it as a distinct species, as did Peterson et al. (1995: 183) and Russ et al. (2001). We are (yet) unable to confirm whether the original spelling is madagascariensis (see Koopman, 1993a: 239, Peterson et al., 1995, Russ et al., 2001, Simmons, 2005: 447) or madagascarensis (see Horácek et al., 2000: 137; Fenton et al., 2002: 1584). COMMON NAMES: Czech: morous madagaskarský. English: Madagascar Free-tailed Bat, Madagascar Mastiff Bat. French: Molosse à grandes oreilles de Madagascar. German: Madagassische RiesenBulldoggfledermaus. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Muldoon et al. (2009: 1114) report on subfossil remains from the Ankilitelo fauna (late Holocene, appr. 500 years ago). CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread but patchy distribution in Madagascar. It is not well known in Madagascar, but it is likely distributed more widely than current records suggest and it is unlikely that it is declining at a rate that would warrant listing in a higher category of threat (Andriafidison et al., 2008I; IUCN, 2009; Monadjem et al., 2017cd). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cd). 2008: LC ver 3.1 (2001) (Andriafidison et al., 2008I; IUCN, 2009). African Chiroptera Report 2020 Regional None known. MAJOR THREATS: The threats to this species are unclear, but more research is needed into possible disturbance at its roost sites (Andriafidison et al., 2008I; IUCN, 2009; Monadjem et al., 2017cd). CONSERVATION ACTIONS: Andriafidison et al. (2008I) [in IUCN (2009)] and Monadjem et al. (2017cd) report that this species roosts in one cave that is locally protected by 'fady' or taboo (Andriafidison et al., 2007). It has been recorded from five protected areas: Parc National du Tsingy de Bemaraha, Parc National de Namoroka, Parc National d’Isalo, Réserve Spéciale d’Ankarana and Réserve Spéciale d’Analamerana (Goodman et al., 2005a). A total of nine roosting colonies are currently known: six in Réserve Spéciale d’Ankarana (Cardiff, 2006), two in Parc National du Tsingy de Bemaraha (Andriafidison et al., 2007) and one in Sarodrano (Goodman et al., 2005a; Andriafidison et al., 2007), and all receive some form of protection. More research is needed on its roosting ecology and surveys are required in areas where this species is expected to occur. GENERAL DISTRIBUTION: Otomops madagascariensis is endemic to the island of Madagascar where it has a disjunct distribution from the north to the south-west of the island. It has an elevation range of 5 m to 800 m above sea level (Goodman et al., 2005a). O'Brien (2011: 289) reports it from the Tanzanian island of Mafia. Native: Madagascar (Valdivieso et al., 1979; Peterson et al., 1995: 183; Russ et al., 2001; Simmons, 2005: 447). DETAILED MORPHOLOGY: Baculum: The baculum of O. madagascariensis is an elongated rod with a notable twist to the mid-shaft. In lateral view, its proximal portion is distinctly wider than the proximal end (Rakotondramanana and Goodman, 2017: 62); length: 0.74 ± 0.057 (0.65 - 0.80) mm, width: 0.13 ± 0.028 (0.09 - 0.18) mm. MOLECULAR BIOLOGY: DNA - See Lamb et al. (2008). Karyotype - Unknown. Protein / allozyme - Unknown. 551 HABITAT: Dammhahn and Goodman (2013: 108) report that this species' foraging habitat includes open areas and the area above the forest canopy. ROOST: Cave roosts in limestone or sandstone (Goodman et al., 2005a; Andriafidison et al., 2007). POPULATION: Structure and Density:- The colony of O. madagascariensis in a cave in the south of Parc National du Tsingy de Bemaraha consisted of between 90 and 100 individuals in 2003 (Andriafidison et al., 2007). The Sarodrano roost in the south of Madagascar contained at least 67 animals in November 2003 (Andriafidison et al., 2007). The maximum colony size at Réserve Spéciale d’Ankarana was 97 individuals (S. G. Cardiff pers. comm. [in Andriafidison et al. (2008I) in IUCN (2009); Monadjem et al., 2017cd]). Trend:- 2016: Unknown (Monadjem et al., 2017cd). 2008: Unknown (Andriafidison et al., 2008I; IUCN, 2009). PARASITES: Lagadec et al. (2012: 1696), Lei and Olival (2014: Suppl.) and Gomard et al. (2016: 5) report the presence of Leptospira sp. bacteria in this species, which were identified as having sequences closely related to L. borgpetersenii by Dietrich et al. (2018a: 3). Laudisoit et al. (2012) describe a new species Lagaropsylla makay (Siphonaptera; Ischnopsyllidae), and they also mention a new parasite-host association between Araeopsylla martialis (Rothschild, 1903) (Siphonaptera; Ischnopsyllidae) and O. madagascariensis. Ramasindrazana et al. (2016: 6) report the presence of an unnamed filariod in this bat species. VIRUSES: Paramyxoviridae Wilkinson et al. (2012: 160) tested 8 individuals from the Madagascar using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Wilkinson et al. (2014) tested 18 individuals from Anjohikinakina in Madagascar with an RT-PCR specific for the Respiro-, Morbilli- and Henipavirus genera, five of the 18 individuals tested positive for paramyxovirus RNA. Seven out of 39 Madagascan specimens tested by Mélade et al. (2016b: 4) were positive for paramyxoviruses. 552 ISSN 1990-6471 Rhabdoviridae Mélade et al. (2016a: 6) examined 20 specimens for Lagos bat lyssavirus and got three positive results. Four out of 18 animals had a positive outcome for Duvenhage lyssavirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Figure 197. Distribution of Otomops madagascariensis Otomops martiensseni (Matschie, 1897) *1897. Nyctinomus martiensseni Matschie, Arch. Naturgesch. Berlin, 63 (1): 84. Publication date: October 1897. Type locality: Tanzania: 25 mi W of Tanga, SE portion of W Usambara Mts: Magrotto Plantation [05 07 S 38 45 E] [Goto Description]. Holotype: ZMB 97523: ♂, skin and skeleton and skull. Collected by: Martienssen; collection date: August 1897. See Turni and Kock (2008: 71). - Comments: Horácek et al. (2000: 137) mention the type locality as "Usanbara Mts., W of Tanga, NE Tanzania". Grubb et al. (1998: 98) indicate that the locality has been defined by Swynnerton, 1945, Proc. Zool. Soc. Lond., 115: 63). - Etymology: In honour of "Herr Martienssen of the Königlich Museum für Naturkunde" who found the type specimen (see Long, 1995: 4). . 1966. Otomops maertiensseni: Hayman, Misonne and Verheyen, Ann. Kon. Mus. Mid. Afr., Zool. Wetensch., (8) 154: pl. XX. Publication date: December 1966. (Lapsus) 2017. O[tomops] martiensenni: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 10. Publication date: 6 April 2017. (Lapsus) 2017. Otomops martiensenii: Waruhiu, Ommeh, Obanda, Agwanda, Gakuya, Ge, Yang, Wu, Zohaib, Hu and Shi, Virol. Sin., 32 (2): 7. Publication date: 6 April 2017. (Lapsus) 2017. Otomops martienssi: Tao, Shi, Chommanard, Queen, Zhang, Markotter, Kuzmin, Holmes and Tong, J. Virol., 91 (5) e01953-16: Suppl. Publication date: 11 January 2017. (Lapsus) ? Otomops martiensseni: (Name Combination, Current Combination) TAXONOMY: Koopman (1993a: 239) included madagascariensis, but see Peterson et al. (1995: 183), Long (1995) and Simmons (2005: 447). Lamb et al. (2008: 30) indicate that there are two clades in martiensseni, of which "clade 1" (northeast Africa + Yemen) would constitute an undescribed taxon. Additionally, Lamb et al. (2008: 31) indicate that the population from Côte d'Ivoire may represent a distinct lineage. Ralph et al. (2015) proved that the specimens from Ethiopia, Eritrea, Somalia, Djibouti and Yemen represent a different species: O. harrisoni, which is larger and more robust than any other Otomops. Ammerman et al. (2012: 23) indicate that the separation between martiensseni and madagascariensis took place less than 2 million years ago. Simmons (2005: 447) recognises one subspecies icarus Chubb, 1917. COMMON NAMES: Afrikaans: Bakoor-losstertvlermuis. Chinese: 大 耳犬吻蝠. Czech: morous Martienssenův. English: Martienssen's Big-eared Bulldog Bat, Large-eared Free-tailed Bat, Giant Mastiff Bat, Martiensen's Free-tailed Bat, Martienssen bat, Large-eared Giant Mastiff Bat. French: Grand molosse à grandes oreilles, Chauve-souris bouledogue géante. German: Großohrige African Chiroptera Report 2020 Riesen-Bulldoggfledermaus. Italian: Otocarlìno di Martienssèn. Kinande (DRC): Mulima. CONSERVATION STATUS: Global Justification Listed as Near Threatened (NT ver 3.1 (2001)) because this species is probably still in significant decline globally (but probably at a rate of less than 30 % over ten years), owing to the probable continued loss of the known major East African populations. Smaller populations from southern Africa may be increasing. The status of populations in Central and West Africa, where the species is still known from very few specimens, remains unclear. Almost qualifies as threatened under criterion A2c (Mickleburgh et al., 2008dd; IUCN, 2009; Richards et al., 2017). Assessment History Global 2016: NT ver 3.1 (2001) (Richards et al., 2008: NT ver 3.1 (2001) (Mickleburgh 2008dd; IUCN, 2009). 2004: NT ver 3.1 (Mickleburgh et al., 2004cq; IUCN, 2004). VU (Baillie and Groombridge, 1996). 2017). et al., (2001) 1996: Regional South Africa: 2016: LC ver 3.1 (2001) (Richards et al., 2016e). 2004: VU D2 ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). Legal Status Richards (2017) report that it is a listed Threatened or Protected Species (ToPS) under the South African National Environmental Management (NEMBA): Biodiversity Act (No. 10 of 2004) and is protected by provincial ordinance in the KwaZuluNatal province. MAJOR THREATS: The leading threat to this species appears to be roost disturbance (Ralph et al., 2015). Populations in Durban (South Africa) have been found to use old buildings with attics as roosts, and it appears that the main threat to these populations is indirect poisoning through the application of toxic timber treatments (Fenton et al., 2002). CONSERVATION ACTIONS: Mickleburgh et al. (2008dd) [in IUCN (2009)] and report that there is a need to reassess the status of all known roosts (and to locate additional localities) to ascertain numbers and status of colonies, so that key sites can be identified. While Richards et al. (2017) report that species has been recorded from Comoe National Park, Côte d'Ivoire (Fahr and Kalko 2011 in Richards et al. (2017), Garamba National Park, Democratic Republic of Congo (Verschuren, 1957), and a 553 national park in Rwanda (Hutson et al., 2001). The species occurs in several protected areas in South Africa (Adams et al., 2015). An IUCN Species Action Plan has been developed for this species (Hutson et al., 2001); this action plan requires revision given the reviewed species limits of O. martiensseni (Lamb et al., 2008; Richards et al., 2012) and the description of new northeast African species, O. harrisoni (Ralph et al., 2015). GENERAL DISTRIBUTION: Otomops martiensseni has been widely, but patchily, recorded from much of sub-Saharan Africa. In Africa, it has been reported from Côte d'Ivoire and Ghana in West Africa, through Central Africa (recorded from Central African Republic, the Democratic Republic of the Congo, Rwanda and western Uganda) to East Africa (Kenya and Tanzania) to southern Africa (Malawi, central Angola [Chitau], Zimbabwe, Zambia and eastern South Africa). It is found at elevations from sea level to 2,900 m asl, although the highest elevations might need to be referred to O. harrisoni. Records from Madagascar are now refered to O. madagascariensis (see Taxonomy section above). Lamb et al. (2008: 30, 39) mention it from Côte d'Ivoire, Ghana, and Burundi, but also see Taxonomy paragraph. Records from Djobuti, Eritrea, Ethiopia, Somalia, Yemen are now referred to O. harrisoni, as might be the case for some of the records from Kenya, Tanzania and Uganda (see Taxonomy section above). Patterson et al. (2018: 1) found both O. martiensseni and O. harrisoni in a cave in the Musanze district of Rwanda. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 545; Taylor et al., 2018b: 63); Botswana (Adams et al., 2015: 711); Central African Republic; Congo (The Democratic Republic of the) (Ansell, 1974; Hayman et al., 1966; Monadjem et al., 2010d: 545); Côte d'Ivoire; Ghana; Kenya (Patterson et al., 2018: 5); Malawi (Monadjem et al., 2010d:454); Rwanda (Patterson et al., 2018: 1); South Africa (Monadjem et al., 2010d:545;Adams et al., 2015: 710-711); Tanzania; Uganda; Zambia Monadjem et al., 2010d: 545); Zimbabwe (Lamb et al., 2008; Monadjem et al., 2010d: 545). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Large species, with narrowest wings of any bat for fast flying in open areas (Anonymus, 2004f: 57). FUNCTIONAL MORPHOLOGY: Based on the analysis of wing dimensions, Thollesson and Norberg (1991: 34) suggest that 554 ISSN 1990-6471 the longer hand wings and more pointed wing tips of O. martiensseni indicate that this is a species primarily flying in open areas at a high altitude. ECHOLOCATION: From Bungule in Kenya Taylor et al. (2005) reported a maximum frequency of 16.5 kHz. However, due to the split off of O. harrisoni, the taxonomic status of the specimens involved is uncertain (see Ralph et al., 2015: 26). At Sengwa in Zimbabwe, Jacobs (1996) and Fenton and Bell (1981) recorded a highest frequency of 17 kHz. Taylor (1999b) recorded maximum frequencies of 29.5 (2.4) kHz (open flight) and 14.7 (1.5) (emergence) from two localities in the Durban area in South Africa. Whereas, Fenton et al. (2004) reported a highest frequency of 14.7 (1.68) kHz from the Durban area in South Africa. From the same area, Schoeman and Waddington (2011: 291) mention a peak frequency of 10.7 ± 0.4 kHz and a duration of 27 ± 1.5 msec. Fenton (2004b: 4) states that there is evidence that, in this species, calls originally associated with echolocation may actually serve more for communication. Adams et al. (2015: 709) report the following values from South Africa: Fchar: 11.70 ± 3.61 (10.51 - 13.54) kHz; Fhigh: 13.19 ± 1.08 (11.01 - 15.67) kHz; Flow: 10.82 ± 0.65 (9.11 - 12.23) kHz; Band width: 2.40 ± 0.90 (0.65 - 5.64) kHz; Fmax: 12.41 ± 0.90 (10.76 - 14.55) kHz; Duration: 21.18 ± 3.61 (11.79 - 27.81) msec. Also from the RSA ( Mapungubwe National Park), Parker and Bernard (2018: 57) reported on 20 calls: Fchar: 10.74 ± 0.63 kHz, Fmax: 11.52 ± 0.60 kHz, Fmin: 10.28 ± 0.50 kHz, Fknee: 10.81 ± 0.64 kHz, duration: 22.68 ± 13.89 msec, with 3.98 ± 2.40 calls/sec. Nine calls from the Okavango River Basin reported by Weier et al. (2020: Suppl.) had the following characteristics: Fmax: 13.16 ± 1.93 kHz, Fmin: 9.75 ± 2.64 kHz, Fknee: 12.61 ± 1.63 kHz, Fchar: 11.77 ± 1.02 kHz, slope: 16.52 ± 6.04 Sc, duration: 12.99 ± 5.28 msec. For seven calls from the Aberdares Range in Kenya, Eisenring et al. (2016: SI 2) reported: PF: 14.8 ± 2.8 (9.9 - 17.4), HF: 22.0 ± 4.2 (14.5 - 26.4), LF: 9.9 ± 2.5 (5.2 - 13.2), DT: 0.1 ± 0.0 (0.1 - 0.2), DF: 12.1 ± 2.4 (9.3 - 16.0), IPI: 3.3 ± 2.0 (1.5 - 5.5). Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 11.8 kHz and duration: 27 msec. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003), Lamb et al. (2006) and Lamb et al. (2008). Karyotype - Ðulic and Mutere (1973a) reported 2n = 48, FN = 56, BA = 10, a submetacentric X chromosome, and an acrocentric Y chromosome. However, Warner et al. (1974) reported 2n = 48, FN = 58, BA = 12, a submetacentric X chromosome and a submetacentric Y chromosome for one male specimen from Kenya. In view of the split off of O. harrisoni the Kenyan karyotype might represent the latter species rather than O. martiensseni. Protein / allozyme - Unknown. HABITS: O. martiensseni is probably a long-range forager, which hunts over a wide diversity of habitats, e.g. semi-arid areas and montane forests up to 2,000 m (Anonymus, 2004f: 57). Adams et al. (2015), referring to Fenton and Griffin (1997), indicate that foraging occurs at altitudes exceeding 600 m above ground. Actually, Fenton and Griffin (1997: 248) reported one call in the 10 - 13 kHz range (used by O. martiensseni) at an altitude between 500 and 550 m. Foord et al. (2018: 5) indicate that open-air feeders (e.g. O. martiensseni) may commute over 10 km between roosts and feeding areas, but do prefer smaller feeding areas. ROOST: In the Durban area, Taylor et al. (1999: 67) found roosts on the brick surface of the inner gable apex wall within the roof space, with entrances height between 3 and 12 m. The colony size varied between one and 30. Richardson and Taylor (1995: 73) already remarked that the exit holes did not have obstructing trees in the way. Anonymus (2004f: 57) indicates that O. martiensseni is generally colonial with larger colonies in underground sites (e.g. caves and lava tubes), whereas in South Africa small colonies occur in houses. POPULATION: Structure and Density:- Although O. martiensseni was once considered to be rare, with gaps in distribution, additional collecting has demonstrated local abundance in several areas (Long, 1995; Taylor, 1998; Fenton et al., 2002; Skinner and Chimimba, 2005). For example, it is common around Durban in KwaZulu Natal Province of South Africa (Fenton et al., 2002), and is fairly common in Comoe National Park, Côte d'Ivoire (Richards et al., 2017). African Chiroptera Report 2020 Trend:- 2016: Decreasing (Richards et al., 2017). 2008: Decreasing (Mickleburgh et al., 2008dd; IUCN, 2009). ACTIVITY AND BEHAVIOUR: Richardson and Taylor (1995: 73) found that males might be chased from their birth colony once they become mature. REPRODUCTION AND ONTOGENY: Martin and Bernard (2000: 32) indicate that the corpus luteum undergoes luteolysis in late pregnancy. Krutzsch (2000: 126) further mentions that the species exhibits classical mammalian male-female synchrony of reproductive cycles. Richardson and Taylor (1995: 72) reported births in Durban (RSA) in early to mid-December, and that juveniles were of almost adult size by midJanuary. The young could then still be distinguished by their greyer colour. Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] indicated that juveniles have been recorded from October through May. MATING: Ober et al. (2016: 375) mention that O. martiensseni forms single male/mulitple female groups. PARASITES: SIPHONAPTERA Ischnopsyllidae: Beaucournu and Kock (1996) reported Araeopsylla scitula Rothschild, 1909 from a bat from Kenya. However, in view of the split off of O. harrisoni, this latter species might have been the host for this ectoparasite. ACARI Trombiculidae: Stekolnikov (2018a: 189) mentioned Willmannium suswaensis (Vercammen-Grandjean & Langston, 1976) from a bat collected at Mount Suswa (Kenya). 555 SARS-CoV - Tong et al. (2009) tested two bats collected in Kenya during 2006 positive for the presence of coronavirus RNA in fecal samples. 28.6 % (10 out of 35) of the Kenyan O. martiensseni specimens were positive for CoV (Tao et al., 2017: 3). Flaviviridae Hepacivirus (BHV) - Quan et al. (2013: Table S5) report that 3 of 15 Kenyan specimens were infected with clade C Hepacivirus (20.0 %). Pegivirus (BPgV) - 2 of the 15 specimens examined by Quan et al. (2013: Table S5) were also infected with clade K Pegivirus (13.3 %). Paramyxoviridae: Paramyxovirus - Five of the 19 specimens from the Suswa Cave by Conrardy et al. (2014: 259) tested positive for this virus, and this was also the case for nine out of 40 specimens from Kenya, which were tested by Mortlock et al. (2015: 1481). Polyomaviridae: Orthopolyomavirus - Initially, three novel orthopolyomaviruses were detected in 2006 from Kenya by Tao et al. (2012). Additional surveillance detected 6 of the 19 specimens from the Suswa Cave to be positive tested by Conrardy et al. (2014: 259). Rhabdoviridae Lyssavirus - Rabies related viruses Horton et al. (2014: Table S1) tested 16 Kenyan O. martiensseni specimens but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). Remark: as most of these viruses were found on "O. martiensseni" specimens from Kenya, the actual host species might be O. harrisoni. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa". VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in Otomops martiensseni: Adenoviruses and Polyomaviruses. Adenoviridae: Adenovirus - Two of the 19 specimens from the Suswa Cave (Kenya) by Conrardy et al. (2014: 259) tested positive for this virus. Coronaviridae - Coronaviruses Figure 198. Distribution of Otomops martiensseni 556 ISSN 1990-6471 Otomops martiensseni icarus Chubb, 1917 *1917. Otomops icarus Chubb, Ann. Durban Mus., 1: 433, pl. 21. Publication date: 21 May 1917. Type locality: South Africa: KwaZulu-Natal: Durban [29 52 S 31 00 E] [Goto Description]. Holotype: BMNH 1916.10.9.1: ♂. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. - Etymology: Named after Icarus, son of Daidalos, who fled from their prison with wings made of wax and feathers. ? Otomops martiensseni icarus: (Name Combination, Current Combination) GENERAL DISTRIBUTION: From Angola to KwaZulu-Natal, Malawi (Horácek et al., 2000: 137). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Côte d'Ivoire, Ghana, South Africa, Uganda. Otomops martiensseni martiensseni (Matschie, 1897) *1897. Nyctinomus martiensseni Matschie, Arch. Naturgesch. Berlin, 63 (1): 84. Publication date: October 1897. Type locality: Tanzania: 25 mi W of Tanga, SE portion of W Usambara Mts: Magrotto Plantation [05 07 S 38 45 E] [Goto Description]. Holotype: ZMB 97523: ♂, skin and skeleton and skull. Collected by: Martienssen; collection date: August 1897. See Turni and Kock (2008: 71). - Comments: Horácek et al. (2000: 137) mention the type locality as "Usanbara Mts., W of Tanga, NE Tanzania". Grubb et al. (1998: 98) indicate that the locality has been defined by Swynnerton, 1945, Proc. Zool. Soc. Lond., 115: 63). - Etymology: In honour of "Herr Martienssen of the Königlich Museum für Naturkunde" who found the type specimen (see Long, 1995: 4). . ? Otomops martiensseni martiensseni: (Name Combination, Current Combination) GENERAL DISTRIBUTION: Central African Republic to Yemen and south to Zimbabwe (Horácek et al., 2000: 137). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Burundi, Central African Republic, Congo (Democratic Republic of the), Ethiopia, Kenya, Malawi, Rwanda, South Africa, Tanzania, Zimbabwe. Genus Platymops Thomas, 1906 *1906. Platymops Thomas, Ann. Mag. nat. Hist., ser. 7, 17 (101): 499. Publication date: 1 May 1906 [Goto Description]. - Comments: Type species: Platymops Macmillani Thomas, 1906 (=Mormopterus setiger Peters, 1878). - Etymology: From the Greek adjective "πλατύς" (platýs), meaning "flat" and the masculine German substantive Mops, referring to the extremely flattened skull of the genus (see Lanza et al., 2015: 315). (Current Combination) ? Mormopterus (Platymops): (Name Combination) TAXONOMY: Considered a subgenus of Mormopterus by Koopman (1993a: 238), Czaplewski et al. (2003: 68), but a valid genus by Kock (1969a: 155), Simmons (2005). Included in Mormopterus by Jacobs and Fenton (2002: 1). Currently (Simmons and Cirranello, 2020) recognized species of the genus Platymops: setiger (Peters, 1878). COMMON NAMES: Czech: ploší morousi. English: East African Flatheaded Bats, Peters's flat-headed Bats. French: Tadarides à tête plate. German: FlachkopfBulldoggfledermäuse. Italian: Platicarlìni. African Chiroptera Report 2020 557 Platymops setiger (Peters, 1878) *1878. Mormopterus setiger Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 196, pl. 1, fig. 2. Type locality: Kenya: Taita district: Ndi [03 14 S 38 30 E, 2 000 m] [Goto Description]. Syntype: ZMB 5265a: ♀, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008). Syntype: ZMB 5265b: ♀, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008). - Comments: Allen (1939a: 109) mentioned 1881; Monatsb. K. Preuss. Akad. Wiss., Berlin, p. 483, pl., fig. 3. - Etymology: From the masculine Latin adjective sètiger, meaning "covered with coarse hair, bristly", referring to the coarse fur of the species Lanza et al., 2015: 316). 1960. Platymops setiger: Harrison and Fleetwood, Durban Mus. Novit., 5 (20): 269. (Name Combination, Current Combination) TAXONOMY: See Freeman (1981b: 133, 161). Included in Platymops by Harrison and Fleetwood (1960: 277 - 278). COMMON NAMES: Chinese: 彼 德 犬 吻 蝠 . Czech: morous plochý. English: East African Flat-headed Bat, Peters's Flat-headed Bat, Flat-headed free-tailed bat. French: Sauromys à tête plate, Tadaride à tête plate de Peters. German: Peters' FlachkopfBulldoggfledermaus. Italian: Carlìno setolóso. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008al; IUCN, 2009; Monadjem et al., 2017aj). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017aj). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008al; IUCN, 2009). 2004: DD ver 3.1 (2001) (Mickleburgh et al., 2004ay; IUCN, 2004). 1996: LR/lc [assessed as Mormopterus setiger] (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: There are presumably no major threats to this species (Mickleburgh et al., 2008al; IUCN, 2009; Monadjem et al., 2017aj). CONSERVATION ACTIONS: Mickleburgh et al. (2008al) [in IUCN (2009)] and Monadjem et al. (2017aj) report that there appear to be no direct conservation measures in place. In view of the species' East African range, it seems likely that it is present within some protected areas, although this needs to be confirmed. Further studies are needed into the distribution, natural history and potential threats to this species. GENERAL DISTRIBUTION: Platymops setiger is an East African species present in Kenya, southwestern Ethiopia (Glass, 1965: 179), southeastern Sudan and is possibly present in adjacent Uganda, although this needs to be confirmed. It has been recorded at elevations of up to 2,000 m asl. Native: Ethiopia; Kenya; Sudan. Presence uncertain: Uganda. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Warner et al., 1974) reported 2n = 48, FN = 54, BA = 8 and submetacentric X and acrocentric Y chromosomes. Protein / allozyme - Unknown. POPULATION: Structure and Density:- Little information is available on the population abundance or size of this species (Mickleburgh et al., 2008al; IUCN, 2009; Monadjem et al., 2017aj). Trend:- 2016: Unknown (Monadjem et al., 2017aj). 2008: Unknown (Mickleburgh et al., 2008al; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Kenya. 558 ISSN 1990-6471 Figure 199. Distribution of Platymops setiger Platymops setiger macmillani Thomas, 1906 *1906. Platymops Macmillani Thomas, Ann. Mag. nat. Hist., ser. 7, 17 (101): 500. Publication date: 1 May 1906. Type locality: Ethiopia: Between Addis Ababa and Lake Rudolf [Goto Description]. Holotype: BMNH 1906.11.1.10: ad ♂, skull and alcoholic. Collected by: Ph.C. Zaphiro. Presented/Donated by: W.N. McMillan Esq. 7 specimens. - Etymology: In honour of Mr. W.N. McMillan, to whose liberality science is indebted for the exploration of which it is part of the outcome (see Thomas, 1906b: 501). 1956. Platymops barbatogularis Harrison, Ann. Mag. nat. Hist., ser. 12, 9 (104): 549. Publication date: 18 December 1956. Type locality: Sudan: SE Sudan, Ilemi triangle: Lokomarinyang Marsh [05 02 N 35 33 E] [Goto Description]. Holotype: BMNH 1967.1232: ad ♂, skin and skull. Collected by: John George Williams; collection date: 10 June 1953. Formerly HZM 2.1897 (see Harrison (1956e: 549). Paratype: BMNH 1957.258:. - Comments: Evenhuis (2003: 54) indicates that ser. 12, 9 (104) is the "August 1956" issue which was published on "18 December 1956". 1960. Platymops barbatogularis parkeri Harrison and Fleetwood, Durban Mus. Novit., 5 (20): 270, figs 1 - 6. Publication date: 30 April 1960. Type locality: Kenya: Lualeni borehole: Maktau [03 24 S 38 08 E] [Goto Description]. Holotype: BMNH 1959.524: ad ♀, skin and skull. Collected by: Ian Parker; collection date: 12 February 1958; original number: 2. ? Mormopterus setiger macmillani: (Name Combination) ? Platymops setiger macmillani: (Name Combination, Current Combination) TAXONOMY: Kock (1969a: 155) considers barbatogularis to be a synonym of macmillani. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia, Kenya, South Sudan. Platymops setiger setiger (Peters, 1878) *1878. Mormopterus setiger Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 196, pl. 1, fig. 2. Type locality: Kenya: Taita district: Ndi [03 14 S 38 30 E, 2 000 m] [Goto Description]. Syntype: ZMB 5265a: ♀, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008). Syntype: ZMB 5265b: ♀, skull and alcoholic. Collected by: Dr. Johann Maria Hildebrandt; collection date: July 1877. See Turni and Kock (2008). - Comments: Allen (1939a: 109) mentioned 1881; Monatsb. K. Preuss. Akad. Wiss., Berlin, p. 483, pl., fig. 3. - Etymology: From the masculine Latin adjective sètiger, meaning "covered with coarse hair, bristly", referring to the coarse fur of the species Lanza et al., 2015: 316). ? Mormopterus setiger setiger: (Name Combination) ? Platymops setiger setiger: (Name Combination, Current Combination) African Chiroptera Report 2020 559 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, South Sudan. Genus Sauromys Roberts, 1917 1917. Platymops Roberts, Ann. Transv. Mus., 6(1): 5. Publication date: 28 June 1917. Comments: Type species: Platymops (Sauromys) petrophylus Roberts, 1917. Not of Thomas, 1906. *1917. Platymops (Sauromys) Roberts, Ann. Transv. Mus., 6(1): 5. Publication date: 28 June 1917. - Comments: Type species: Platymops (Sauromys) haagneri Roberts, 1917 (=Platymops petrophilus Roberts, 1917). As a subgenus of Platymops Thomas, 1906. Etymology: From Greek "sauros" meaning lizard and "mys" meaning mouse (see Jacobs and Fenton, 2002: 1). ? Mormopterus (Sauromys): (Name Combination) ? Sauromys: (Current Combination) ? Tadarida (Sauromys): (Name Combination) TAXONOMY: Bronner et al. (2003) state that the status of Sauromys, described as a monotypic subgenus of the extralimital Platymops for the South African flat-headed free-tailed bat, remains unclear. Peterson (1965) raised it to generic rank, a treatment endorsed by many subsequent authors, and corroborated by a limited multivariate analysis of wing bone and cranial characteristics (Peterson, 1985). However, morphometric studies by Freeman (1981b) and Legendre (1984) concluded that it is a subgenus of Mormopterus, a position followed by Koopman (1993a, 1994), and Jacobs and Fenton (2002: 1). Simmons (2005) retains Sauromys as a valid genus in the 3rd Edition of Mammal Species of the World, a treatment we favour because of the unique ecology and morphology of S. petrophilus. Analysis of the Rag2 data confirmed the split between Mormopterus and Sauromys (see Ammerman et al., 2013: 309). Meester et al. (1986) also state that originally described as a subgenus, this genus was regarded as a synonym of Platymops by Shortridge (1942), Roberts (1951) and Ellerman et al. (1953). Harrison (1962) retains it as a subgenus, while Peterson (1965) raises it to generic rank, in which he is followed by Hayman and Hill (1971), Smithers (1971), Corbet and Hill (1980) and Rautenbach (1982). However, Freeman (1981b), followed by Koopman (1982) and Legendre (1984), conclude that both Sauromys and Platymops are subgenera of Mormopterus. We take the view that both Platymops and Sauromys are valid genera. Generic and species classification of African flatheaded molossids is unclear. Roberts (1917c) originally described Sauromys as a subgenus of Platymops. Later Roberts (1951, 1954) dropped Sauromys as a subgenus, as Shortridge (1942) reported that a minute upper premolar was found on the type Platymops macmillani Thomas, 1906. Peterson (1965) indicates that the minute premolar on the type P. macmillani is probably a milk premolar and that normal adult dental formula should be considered as having only one upper premolar. For this reason and others (e.g. glands not on wings) Peterson (1965) raises Sauromys to generic status. Koopman (1975) indicates that Sauromys is a subgenus within Tadarida, however Freeman (1981b) found that all the so-called flatheaded bats; Sauromys, Platymops, and Neoplatymops, are phenotypically related to Mormopterus. While the phenogram in Freeman (1981b) also indicates that Tadarida aegyptiaca (E. Geoffroy 1818) and T. brasiliensis (I. Geoffroy St.-Hilaire 1824) are phenetically similar to Mormopterus, this grouping is explained due to convergence of shape due to similar life styles (Freeman, 1981b). Meester et al. (1986) did not follow Freeman (1981b) and rather followed Peterson (1965) recognizing Sauromys, Mormopterus and Platymops as valid genera. While Koopman (1993a) followed Freeman (1981b) in classifying Sauromys and Platymops as subgenera in the genus Mormopterus, Simmons (2005) recognizes Sauromys as a distinct genus. Peterson (1965) suggests that a detailed comparison between Sauromys and Tadarida needs to be performed, including an investigation into the specific status and geographic variation. Ammerman et al. (2012: 21) suggest that Sauromys is more appropriately recognized as a member of the genus Tadarida, which split off from T. aegyptiaca some 6.5 million years ago. However, based on their morphological data, Gregorin and Cirranello (2015: 10) still find it premature to transfer Sauromys into Tadarida. 560 ISSN 1990-6471 Currently (Simmons and Cirranello, 2020) recognized species of the genus Sauromys: petrophilus (Roberts, 1917). COMMON NAMES: Czech: ještěří morousi. French: Sauromys. German: Flachkopf-Bulldoggfledermäuse. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Avery (2007: 619) reported on the presence of Sauromys sp. in Pleistocene deposits at Wonderwerk Cave, South Africa. Sauromys petrophilus (Roberts, 1917) *1917. Platymops (Sauromys) petrophilus: Roberts, Ann. Transv. Mus., 6(1): 5. Publication date: 28 June 1917. 1917. Platymops petrophilus Roberts, Ann. Transv. Mus., 6(1): 5. Publication date: 28 June 1917. Type locality: South Africa: Western Transvaal: near Rustenburg: Bleskop [Goto Description]. Holotype: TM X835: ♂. Collected by: G.P. Van Dam; collection date: 2 February 1917; original number: 16. Presented/Donated by: ?: Collector Unknown. Etymology: From the Greek "petros" meaning rock and "philos" meaning loving. (Name Combination) ? Mormopterus petrophilus petrophilus: (Name Combination) ? Mormopterus petrophilus: (Name Combination) ? Platymops haagneri haagneri: ? Sauromys petrophilus: (Name Combination, Current Combination) TAXONOMY: (Shortridge and Carter, 1938). Included in Mormopterus by Jacobs and Fenton (2002: 1). Jacobs and Fenton (2002: 1) considered M. petrophilus to be monotypic. COMMON NAMES: Afrikaans: Platkop-losstertvlermuis. Czech: morous ještěří. English: Rock-loving Flat-headed Bat, Rock-dwelling Flat-headed Bat, Roberts' Flatheaded Bat, Roberts's Flat-headed Bat, Flatheaded Free-tailed Bat. French: Sauromys de Roberts. German: Roberts' FlachkopfBulldoggfledermaus. Portuguese: Morcego de cabeca curta. Figure 200. Sauromys petrophilus umbratus from Kliphuis, Cederberg, Western Cape South Africa. Roberts (1951, 1954) recognised two species and five subspecies of flat-headed bats in southern Africa: Platymops haagneri Roberts, 1917 (haagneri, umbratus Shortridge and Carter, 1938) and Platymops petrophilus Roberts, 1917 (petrophilus, erongensis Roberts, 1946, fitsimonsi Roberts, 1946). Ellerman et al. (1953) suggested that the two species are conspecific, and rather recognized five subspecies of Platymops petrophilus (petrophilus, haagneri, umbratus, erongensis and fitzsimonsi). Meester et al. (1986) also recognized the five subspecies suggested by Ellerman et al. (1953) for petrophilus but within the genus Sauromys, following Peterson (1965: 12). Freeman (1981b) indicates that erongensis (Roberts, 1946) may be a synonmy of haagneri (Roberts, 1917) and fitzsimonsi (Roberts, 1946) is possibly a synonym of umbratus PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: No fossils are known (Jacobs and Fenton, 2002: 1). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Mickleburgh et al., 2008an; IUCN, 2009; Monadjem et al., 2017bh). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bh). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008an; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bc; IUCN, 2004). 1996: LR/lc [assessed as Mormopterus petrophilus] (Baillie and Groombridge, 1996). African Chiroptera Report 2020 Regional South Africa:- 2016: LC ver 3.1 (2001) (Jacobs et al., 2016f). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). 561 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: As indicated by Rautenbach and Nel (1980: 112), the extremely flattened skull is an adaptation to roosting in the narrowest of rock crevices. For further details, see Jacobs and Fenton (2002). MAJOR THREATS: There appear to be no major threats to this species. In parts of its range it is believed to be threatened by considerable habitat change resulting from deforestation (Mickleburgh et al., 2008an; IUCN, 2009; Monadjem et al., 2017bh). DETAILED MORPHOLOGY: Baculum - Unknown For a description of the crania, ears and tragus, and wingshape and aspect ratio, see Jacobs and Fenton (2002). CONSERVATION ACTIONS: Mickleburgh et al. (2008an) [in IUCN (2009)] and Monadjem et al. (2017bh) report that this species is present in several protected areas in Namibia and Angola. In South Africa it has been reported from the protected Algeria forestry station. No direct conservation measures are currently needed for this species as a whole. Curtis and Simmons (2018: 1390) found a relatively large, anteroposteriorly elongated, single, mineralized midline element positioned anterior to the dorsal margin of the nasal aperture in the cupular cartilages. Their CT-microscans showed a variation in the grayscale, suggesting porosity or trabeculae, which in turn might suggest a bony structure. GENERAL DISTRIBUTION: Sauromys petrophilus is found in southern Africa, ranging along the west coast from the Angola and Namibia border area, south into southern South Africa, and from here into northeastern South Africa, Zimbabwe, and western Mozambique. It is found between 100 m and 2,000 m asl. In the RSA, its distribution is best predicted by geology (Babiker Salata, 2012: 50). FUNCTIONAL MORPHOLOGY: Toussaint and McKechnie (2012: 1138) found that the rapid increases in Evaporate Water Loss at high ambient temperatures in S. petrophilus may imply that, notwithstanding having a comparatively heat tolerance, this species may also be most vulnerable to lethal dehydration during future heat waves. Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 915,652 km2. The Ghanaian specimen which should be in the USNM (see Meester et al., 1986), could not be located in the collection (see Jacobs and Fenton, 2002: 2). Native: Angola; Botswana (Monadjem et al., 2010d: 545); Mozambique (Monadjem et al., 2010d: 545; Monadjem et al., 2010c: 382); Namibia (Monadjem et al., 2010d: 545); South Africa (Seamark and Kearney, 2007; Monadjem et al., 2010d: 545)); Zimbabwe (Monadjem et al., 2010d: 545). Presence uncertain: Ghana. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Jacobs and Fenton (2002). Schoeman and Jacobs (2002: 157) mention the following wing parameters for 51 specimens from the Algeria Forestry Station (RSA): Fa: 39.0 ± 1.0 mm, Wing area: 93.1 ± 11 cm 2, Wing span: 26.3 ± 1.9 cm, Wing loading: 11.2 ± 1.1 N/m2, and aspect ratio: 7.4 ± 0.6. ECHOLOCATION: See Jacobs and Fenton (2002). From Algeria Forest Station (RSA), Schoeman and Jacobs (2002: 157) reported the type of calls to be low duty echolocation dominated by frequency modulated calls, with a Fpeak: 29.2 ± 1.8 kHz. Two calls by bats from the Kalahari Desert had the following characteristics: Fchar: 24.0 ± 0.2 kHz, Fmax: 30.3 ± 3.2 kHz, Fmin: 23.7 ± 0.1 kHz, duration: 4.3 ± 0.7 msec (Adams and Kwiecinski, 2018: 4). MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Rautenbach et al. (1993) reported 2n = 48, FN = 62, BA = 16, a submetacentric X chromosome, and an acrocentric Y chromosome for specimens from Namibia and South Africa. However, this did not include specimens from the Western Cape that Rautenbach et al. (1993) included with Chaerephon pumilus, but have subsequently been reidentified as S. petrophilus (Jacobs and Fenton, 2001). Protein / allozyme - Unknown. HABITAT: Sirami et al. (2013: 34) recorded in the Western Cape Province, South Africa that S. petrophilus 562 ISSN 1990-6471 activity was significantly and positively influenced by size of wetland, while habitats 100 m surrounding wetlands were also significantly and positively influenced by the water body. S. petrophilus favoured open habitats surrounding wetlands (Sirami et al., 2013: 35) . HABITS: See Jacobs and Fenton, 2002). ROOST: See Jacobs and Fenton (2002). DIET: See Jacobs and Fenton (2002). From the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) reported the following prey groups based on 464 faecal pellets from 51 bats (in volume percent): Diptera (37.8 ± 31.2), Hemiptera (32.9 ± 23.7), Coleoptera (19.5 ± 23.1), Hymenoptera (7.9 ± 16.2), Neuroptera (0.6 ± 3.0), Trichoptera (0.4 ± 2.9), Lepidoptera (0.3 ± 2.2), and unknown (1.0 ± 3.3). ACTIVITY AND BEHAVIOUR: See Jacobs and Fenton, 2002). REPRODUCTION AND ONTOGENY: See Jacobs and Fenton (2002). VIRUSES: Rhabdoviridae Lyssavirus - Rabies related viruses Rabies - Oelofsen and Smith (1993) tested 12 individuals from Riemvasmaak, Northern Cape, South Africa using "Trousse Platelia Rage" ELISA kit (Diagnostic Pasteur), none tested positive for antibodies to rabies virus glycoprotein G. Oelofsen and Smith (1993) tested 12 individuals from Riemvasmaak, Northern Cape, South Africa by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Namibia, South Africa. PREDATORS: Mikula et al. (2016: Supplemental data) mention S. petrophilus as possible prey of Lanner falcon (Falco biarmicus Temminck, 1825) and Peregrine falcon (Falco peregrinus Tunstall, 1771), POPULATION: Structure and Density:- This is a locally common species. It is generally found in small numbers, with colonies thought to be tens of animals rather than hundreds (Jacobs and Fenton, 2002). Trend:- 2016: Stable (Monadjem et al., 2017bh). 2008: Stable (Mickleburgh et al., 2008an; IUCN, 2009). Figure 201. Distribution of Sauromys petrophilus Sauromys petrophilus erongensis (Roberts, 1946) *1946. Platymops petrophilus erongensis Roberts, Ann. Transv. Mus., 20 (4): 308. Type locality: Namibia: Damaraland: Omaruru district: Eronga Mountains: Ombu Farm [21 35 S 15 43 E] [Goto Description]. Holotype: TM 9494: ♂. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 19 September 1941; original number: 112 (14). Presented/Donated by: ?: Collector Unknown. Holotype: TM 9570: ♂. Collected by: Austin Roberts; collection date: 22 September 1941; original number: 128. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9495: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 14 September 1941; original number: 77. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9496: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 17 September 1941; original number: 93. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9497: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 17 September 1941; original number: 95. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9497a: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 14 September 1941. Presented/Donated by: ?: African Chiroptera Report 2020 ? ? 563 Collector Unknown. Paratype: TM 9498: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 18 September 1941; original number: 105. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9499: ♂. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 18 September 1941; original number: 106. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9500: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 18 September 1941; original number: 108. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9501: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 18 September 1941; original number: 107. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9502: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 19 September 1941; original number: 113. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9503: ♀. Collected by: The Barlow TM-Expedition and Austin Roberts; collection date: 19 September 1941; original number: 115. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9504: ♂. Collected by: Austin Roberts; collection date: 19 September 1941; original number: 116. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9505: ♂. Collected by: Austin Roberts; collection date: 19 September 1941; original number: 117. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9506: ♂. Collected by: Austin Roberts; collection date: 19 September 1941; original number: 118. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9507: ♀. Collected by: Austin Roberts; collection date: 20 September 1941; original number: 124. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9508: ♂. Collected by: Austin Roberts; collection date: 20 September 1941; original number: 125. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9511: ♂. Collected by: Austin Roberts; collection date: 22 September 1941; original number: 129. Presented/Donated by: ?: Collector Unknown. Paratype: TM 9512: ♀. Collected by: Austin Roberts; collection date: 22 September 1941; original number: 129. Presented/Donated by: ?: Collector Unknown. - Etymology: Referring to the Eronga mountains, where the type specimen was collected. Mormopterus petrophilus erongensis: (Name Combination) Sauromys petrophilus erongensis: (Name Combination, Current Combination) DISTRIBUTION BASED ON VOUCHER SPECIMENS: Namibia. Sauromys petrophilus haagneri (Roberts, 1917) 1917. Platymops (Sauromys) haagneri: Roberts, Ann. Transv. Mus., 6 (1): 5. Publication date: 28 June 1917. (Name Combination) *1917. Platymops haagneri Roberts, Ann. Transv. Mus., 6 (1): 5. Publication date: 28 June 1917. Type locality: Namibia: Great Namaqualand: Keetmanshoop [26 35 00 S 18 08 00 E, 984 m] [Goto Description]. Holotype: TM X472: ad ♂. Collected by: Mr. Sigmund Haagner. Presented/Donated by: Mr. Sigmund Haagner. ? Sauromys haagneri: (Name Combination) ? Sauromys petrophilus haagneri: (Name Combination, Current Combination) COMMON NAMES: Afrikaans: Haagnerse Platkopvlermuis. English: Haagner's Flat-headed Bat. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Namibia, South Africa. Sauromys petrophilus petrophilus (Roberts, 1917) *1917. Platymops (Sauromys) petrophilus: Roberts, Ann. Transv. Mus., 6(1): 5. Publication date: 28 June 1917. ? Sauromys petrophilus petrophilus: (Name Combination, Current Combination) 564 ISSN 1990-6471 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Botswana, Mozambique, South Africa, Zimbabwe. Sauromys petrophilus umbratus (Shortridge & Carter, 1938) *1938. Platymops haagneri umbratus Shortridge and Carter, Ann. S. Afr. Mus., 32 (4): 282. Publication date: July 1938. Type locality: South Africa: Western Cape province: 11 mi [=18 km] NE Clanwilliam, Pakhuis Pass: Kliphuis [32 08 S 19 00 E] [Goto Description]. Holotype: KM 2827: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. 1946. Platymops petrophilus fitzsimonsi Roberts, Ann. Transv. Mus., 20 (4): 308. Type locality: South Africa: SW Cape province: near Ceres: Mitchell's Pass [Goto Description]. Holotype: TM 8968: ♀. Collected by: Austin Roberts; collection date: 7 November 1940; original number: 63. Presented/Donated by: ?: Collector Unknown. Holotype: TM 8969: ♀. Collected by: Austin Roberts; collection date: 7 November 1940. Presented/Donated by: ?: Collector Unknown. ? Sauromys petrophilus umbratus: (Name Combination, Current Combination) DISTRIBUTION BASED ON VOUCHER SPECIMENS: South Africa. Genus Tadarida Rafinesque, 1814 *1814. Tadarida Rafinesque, Précis des découvertes et travaux somiologiques, 55. Publication date: 3 June 1814. - Comments: Type species: Cephalotes teniotis Rafinesque, 1814. Etymology: Meaning unknown (Flannery, 1990: 366). Possibly from the Greek "data" or Latin "taeda" and "nukterida" (see Kozhurina, 2002: 18). Braun and Mares (1995: 182) refer to Gotsch (1979), who states this is a word created by C.S. Rafinesque. Lanza et al. (2015: 317 - 318) gives the following explanation: "From the feminine Greek substantive "νυκτερίς" (nykterís) [genitive "νυκτερίδος" (nikterídos)], meaning "bat"; by aphaeresis ] and dialectal deformation, the above genitive turned into "tadarida" in southern Italy. Some examples of similar dialect deformations are according to Forsyth Major (1893): tagddariti (Naples); taddarita, taddarida and taddarito (Calabria), tardaritòla and tardarira (Sicily)". (Current Combination) 1818. Nyctinomus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 114, pl. 2. Publication date: 1818 [Goto Description]. - Comments: Type species: Nyctinomus aegyptiacus E. Geoffroy St. Hillaire, 1818. In the original description, Geoffroy Saint-Hilaire (1818) mentions three species: Nyctinomus Ægyptiacus [=Tadarida aegyptiaca, Vesp. acetabulosus Hermann [=Mormopterus acetabulosus] and Vesp. plicatus Buchanan [=Chaerephon plicatus], which would make Nyctinomus a partial synonym of these three genera. - Etymology: From the Greek "νύξ" or "νυκτός", meaning night and "νομός", meaning habitation (see Palmer, 1904: 466). 1821. Nyctimones Gray, London Med. Repos., 15: 299. Publication date: 1 April 1821. Comments: Generally, the name is reported as Nyctinomes, but Gray (1821: 299) actually mentioned Nyctimones, with one species Vesp. acetabulosus. 1821. Nyctinoma Bowdich, Analysis of the Natural Classification of Mammalia, p. 28. 1822. Nyctinomia Fleming, The Philosophy of Zoology, 2: 178 [Goto Description]. 1825. Dinops Savi, Nuov. Giorn. Letter. Pisa, Sci., 10: 229. - Comments: Type species: Dinops cestoni Savi, 1824 (=Cephalotes teniotis Rafinesque, 1814). - Etymology: From the Greek "δεινός", meaning terrible and "ώψ", meaning face or aspect, probably referring to the deeply grooved or wrinkled face (see Palmer, 1904: 234). 1830-1831. Dysopes Cretzschmar, in: Rüppell, Atlas Reise Nördlichen Afrika, Zoologie Säugethiere, 1: 69. Publication date: 1830-1831. - Comments: Not of Illiger, 1811. The publication date of Cretzschmar's work is sometimes indicated as "1826" (as mentioned on the title page), but also as "1827", "1828", "1830-1831". 1934. Austronomus Iredale and Troughton, Mem. Aust. Mus., p. 100. - Comments: Type species: Nyctinomus australis atratus Thomas, 1924 (=Molossus australis Gray, 1839). (Nomen Nudum). African Chiroptera Report 2020 1941. 1973. 1984. 1995. ? 565 Austronomus Troughton, Animals of Australia, 1st ed, 360. - Comments: Type species: Nyctinomus australis atratus Thomas, 1924 (=Molossus australis Gray, 1839). Validation of Austronomus Iredale and Troughton, 1934 (see Meester et al., 1986: 68; Simmons (2005). Tatarida: Eisentraut, Bonn. zool. Monogr., 3: 113. (Lapsus) Rhizomops Legendre, Rev. suisse Zool., 91 (2): 427. Publication date: June 1984. Comments: Type species: Rhizomops brasiliensis (I. Geoffroy Saint-Hilaire, 1824). Etymology: From the Greek "riza" meaning root (see Legendre, 1984: 427). Dynops: Pavlinov, Borissenko, Kruskop and Jahonton, Arch. Zool. Mus., Moscow State Univ., 133: ???. (Lapsus) Tadarida sp.: TAXONOMY: Formerly included Chaerephon, Mops, Mormopterus, and Nyctinomops; see Freeman (1981b: 133). Includes Rhizomops, but see Legendre (1984); see also Owen et al. (1990). Mahoney and Walton (1988c) regarded Nyctinomus as the prior name for this genus. Based on their analysis of molecular data, Lamb et al. (2011: 9) suggest that the genus Tadarida - as currently recognized - is not a natural group. Tadarida teniotis - the type species of the genus forms a separate clade from T. aegyptiaca - the type species of Nyctinomus - and "C. jobimena". They suggest that the two latter species - together with other species - might be placed in a separate genus: Nyctinomus, but they await further taxonomic sampling and resolution of the phylogeny of other African taxa before making this formal. Currently (Simmons and Cirranello, 2020) recognized species within the genus Tadarida: aegyptiaca (E. Geoffroy Saint-Hilaire, 1818); brasiliensis (I. Geoffroy Saint-Hilaire, 1824) – southern Brazil, Bolivia, Argentina and Chile to Oregon, south Nebraska and Ohio (USA), Greater and Lesser Antilles (Simmons, 2005: 450); fulminans (Thomas, 1903); insignis (Blyth, 1862) – Japan, Taiwan, Korea, southern China (Simmons, 2005: 450); latouchei Thomas, 1920 – northern China, Thailand, Laos, Japan (Simmons, 2005: 450); lobata (Thomas, 1891); teniotis (Rafinesque, 1814); ventralis (Heuglin, 1861). COMMON NAMES: Czech: širokouší morousi, hroznohledové, netopýři morousovití. English: Guano Bats, Freetailed Bats, Tadarine Bats. French: Tadarides. German: Gewöhnliche Bulldoggfledermäuse. Italian: Tadarìde. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Avery (2007: 619) reported on the presence of Tadarida sp. in Pleistocene deposits at Wonderwerk Cave, South Africa. PARASITES: Barbosa et al. (2016: 215) reported Trypanosoma erneyi from a Tadarida sp. from Mozambique. ACARI Myobiidae: Fain (1994: 1280) reports one species of Schizomyobia, living on Old World Tadarida. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Congo (Democratic Republic of the), Egypt, Kenya, Niger, Somalia, South Africa, South Sudan, Tanzania, The Gambia, Uganda, Zambia. †Tadarida engesseri Rachl, 1983 *1983. Tadarida engesseri Rachl, Die Chiroptera (Mammalia) aus den mittelmiozänen Kalken des Nordlinger Rieses (Suddeutschland). Thesis Doct, Ludwig-Maximilians. Universität München.. Type locality: Germany: Steinberg (Ries). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Middle Miocene (Astaracanian). GENERAL DISTRIBUTION: Europe; Morocco Hugueney et al. (2015: 472). Tadarida aegyptiaca (E. Geoffroy St.-Hilaire, 1818) *1818. Nyctinomus ægyptiacus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 128, pl. 2, No 2. Publication date: 1818. Type locality: Egypt: Giza province: Giza [30 01 N 31 13 E]. Lectotype: MNHN ZM-MO-1986-1084 (A.467): ad ♂, 566 ISSN 1990-6471 1855. 2012. 2018. ? ? skin and skull. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Number 228 in Rode (1941: 251). - Comments: Type locality restricted from "Egypt" to Giza by Koopman (1975: 422): see Qumsiyeh (1985: 73), Meester et al. (1986: 70). - Etymology: From the scientific Latin feminine adjective aegyptìacus, meaning "Egyptian", as the species was decribed from an Egyptian specimen (see Lanza et al., 2015: 319). D[ysopes] aegyptiacus: Giebel, Die Säugethiere., 957. (Name Combination) Nyctinomus aegyptiacus: Benda, Faizolâhi, Andreas, Obuch, Reiter, Ševcík, Uhrin, Vallo and Ashraf, Acta Soc. Zool. Bohem., 76: 163, 516. Publication date: 27 December 2012. (Alternate Spelling) Tadarida aegyptica: Lazzeroni, Burbrink and Simmons, Ecol. Evol., 8 (24): 12584. Publication date: 28 November 2018. (Lapsus) Tadarida (Tadarida) aegyptiaca: (Name Combination) Tadarida aegyptiaca: (Name Combination, Current Combination) TAXONOMY: Benda et al. (2012a: 516) place this taxon in the genus Nyctinomus, a view followed by a.o. Bendjeddou et al. (2016b: 23). Regional South Africa:- 2016: LC ver 3.1 (2001) (MacEwan et al., 2016a). 2004: LC ver 3.1 (2001) (Friedmann and Daly, 2004). COMMON NAMES: Afrikaans: Egiptiese losstertvlermuis. Bengali: Lomba-leji Chamchika. Chinese: 北 非 犬 吻 . Czech: morous egyptský, příšerec egipecký, tadarida jižní, morous jižní. Dutch: Egyptischte bulvleermuis. English: Egyptian Free-tailed Bat, Egyptian Guano Bat, Egyptian Nyctinome. French: Tadaride d'Egypte, Molosse d'Egypte. German: Ägyptische Bulldoggfledermaus. Italian: Tadarìda egiziàna. Portuguese: Morcego de cauda livre do egipto. Tamil: எகிப் தி ய ொலற் ற வெளொல் , Ekiptiya Vāaṟṟa Veḷavā. MAJOR THREATS: No serious threats other than roost disturbance from human interference is noted for this species (Molur et al., 2002). Pesticides used against locusts are a threat as for all bat species found in the Saharan belt (Mickleburgh et al., 2008ar; IUCN, 2009; Monadjem et al., 2017bi). Furthermore, Doty and Martin (2012: 75) report on fatalities due to wind turbines in the Eastern Cape, South Africa. ETYMOLOGY OF COMMON NAME: The original specimens were collected in Egypt during the Napoleonic wars by E. Geoffroy SaintHilaire, who at that time served under Napoleon (see Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Avery and Avery (2011: 18) report on Holocene and Pleistocene remains from Wonderwerk (Northern Cape province, South Africa). CONSERVATION STATUS: Global Justification Broadly distributed and locally common, hence is listed as Least Concern (LC ver 3.1 (2001)) (Mickleburgh et al., 2008ar; IUCN, 2009; Monadjem et al., 2017bi). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bi). 2008: LC ver 3.1 (2001) (Mickleburgh et al., 2008ar; IUCN, 2009). 2004: LC ver 3.1 (2001) (Mickleburgh et al., 2004bb; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). CONSERVATION ACTIONS: Mickleburgh et al. (2008ar) [in IUCN (2009)] and Monadjem et al. (2017bi) report that the species is protected by national legislation in some of the Mediterranean range states as well as in South Africa. It is likely to be found in protected areas. A survey using bat detectors should be undertaken to help clarify the distribution limits and population size of this species as it is probably more widespread than current records indicate. A study on the impacts of pesticides is required, especially ways in which the impact might be minimised. GENERAL DISTRIBUTION: Tadarida aegyptiaca is found throughout Africa, and in the Arabian Peninsula through to India, Sri Lanka and Bangladesh. In the Mediterranean region there are isolated records from southern Morocco and western Algeria, but it is more widely distributed in Egypt along the Nile River valley eastwards to the Red Sea coast and south to the Sudanese border. The species is thought to be more widely distributed than is currently known and may well occur elsewhere in the region. In South Asia, this widely distributed species is presently known from Afghanistan (Kabul Province) (Habibi, 2003), Bangladesh (no exact African Chiroptera Report 2020 location) (Khan, 2001; Srinivasulu and Srinivasulu, 2005), India (Andhra Pradesh, Gujarat, Karnataka, Kerala, Madhya Pradesh, Maharashtra, Rajasthan, Tamil Nadu and West Bengal), Pakistan (Punjab and Sind) and Sri Lanka (Central and Uva Provinces) (Molur et al., 2002). Elevation: sea level up to 2,100 m asl. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,214,127 km 2. In the RSA, the species' distribution is linked with land use/land cover (Babiker Salata, 2012: 50). Native: Afghanistan; Algeria; Angola (CrawfordCabral, 1989; Hayman, 1963; Monadjem et al., 2010d: 545; Taylor et al., 2018b: 63); Bangladesh; Botswana (Monadjem et al., 2010d: 545); Congo (The Democratic Republic of the); Egypt; Ethiopia; Eritrea (Lavrenchenko et al., 2004b: 132); India; Iran, Islamic Republic of; Kenya; Lesotho (Lynch, 1994: 197; Monadjem et al., 2010d: 545); Mauritania; Morocco (Benda et al., 2004d; El Ibrahimi and Rguibi Idrissi, 2015: 365); Mozambique (Monadjem et al., 2010d: 545; Monadjem et al., 2010c: 382); Namibia (Monadjem et al., 2010d: 545); Nigeria; Oman; Pakistan; Saudi Arabia; South Africa (Seamark and Kearney, 2007; Monadjem et al., 2010d: 546); Swaziland (Monadjem et al., 2010d: 546); Sri Lanka; Sudan; Tanzania; Tunisia (Bendjeddou et al., 2016b: 23); Uganda; Yemen (Benda et al., 2011b: 45); Zambia (Cotterill, 2004a: 260; Monadjem et al., 2010d: 546); Zimbabwe (Monadjem et al., 2010d: 546). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Schoeman and Jacobs (2002: 157) reported the following parameters for 43 specimens from the Algeria Forestry Station (RSA): Fa: 45.1 ± 1.2 mm, Wing area: 115.2 ± 8.4 cm 2, Wing span: 30.8 ± 1.0 cm, Wing loading: 13.1 ± 1.6 N/m 2, and aspect ratio: 8.3 ± 0.6. ECHOLOCATION: From Algeria Forest Station (RSA), Schoeman and Jacobs (2002: 157) reported the type of calls to be low duty echolocation dominated by frequency modulated calls, with a Fpeak: 22.8 ± 1.9 kHz. Schoeman and Waddington (2011: 291) mention a peak frequency of 21.8 ± 1.7 kHz and a duration of 12.8 ± 0.7 msec for specimens from Durban, South Africa. From Waterberg, South Africa, Taylor et al. (2013b: 18) recorded the following parameters for 8 calls: Fmax: 28.3 ± 3.36 (24.8 - 34.1) kHz, Fmin: 21.6 ± 2.73 (17.2 - 24.0) kHz, Fknee: 24.2 ± 1.47 (21.9 - 26.1) kHz, Fchar: 22.8 ± 1.39 (20.5 - 24.5) kHz, duration: 7.2 ± 3.10 (3.3 - 10.9) msec. 567 At Farm Welgevonden, South Africa, Taylor et al. (2013a: 556) report a Fknee value of 24 (22 - 26) kHz. Linden et al. (2014: 40) reported the following parameters from the Soutpansberg range (RSA): Fmin:17 - 24 kHz, Fchar: 20 - 24 kHz, Fknee: 22 - 26 kHz, Slope: 65 - 221 OPS, duration: 3 - 11 msec. In the Mapungubwe National Park (RSA), Parker and Bernard (2018: 57) recorded 20 calls with the following characteristics: Fchar: 23.68 ± 1.51 kHz, Fmax: 29.23 ± 5.77 kHz, Fmin: 22.47 ± 1.74 kHz, Fknee: 25.72 ± 2.92 kHz, duration: 9.87 ± 3.93 msec, with 4.10 ± 2.72 calls/sec. Moores and Brown (2017: 612) recorded the following parameters for 25 calls from southern Morocco: Fstart: 26.0 ± 2.6 (22.2 - 30.4) kHz, Fmax: 19.6 ± 1.6 (17.4 - 22.3) kHz, Fend: 18.4 ± 0.8 (17.2 - 19.5) kHz, duration: 9.4 ± 1.4 (8.1 - 12.2) msec, interpulse interval: 373 ± 153 (220 - 780) msec. Adams and Kwiecinski (2018: 4) reported the following values for 5 calls from the Kalahari Desert: Fchar: 21.6 ± 1.2 kHz, Fmax: 25.2 ± 1.1 kHz, Fmin: 21.2 ± 1.1 kHz, duration: 4.9 ± 0.6 msec. Weier et al. (2020: Suppl.) reported on 16 calls from the Okavango River Basin with the following characteristics: Fmax: 21.69 ± 1.48 kHz, Fmin: 19.83 ± 3.23 kHz, Fknee: 21.25 ± 0.89 kHz, Fchar: 20.65 ± 0.71 kHz, slope: 8.58 ± 9.28 Sc, duration: 6.07 ± 2.83 msec. Luo et al. (2019a: Supp.) reported the following data (bats leaving their roost): Fpeak: 19.44 kHz, Fstart: 24.44 kHz, Fend: 16.67 kHz, Band width: 7.77 kHz, and duration: 15.5 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Both Nagorsen et al. (1976) and Rautenbach et al. (1993) reported 2n = 48 and an actrocentric Y chromosome. However, while Nagorsen et al. (1976) reported FN = 54, BA = 8, and a submetacentric X chromosome, Rautenbach et al. (1993) reported FN = 68, BA = 22, and a metacentric X chromosome, for specimens from Namibia and South Africa. Protein / allozyme - Unknown. HABITAT: Sirami et al. (2013: 34) recorded in the Western Cape Province, South Africa that T. aegyptiaca activity was significantly and positively influenced by wetland size, while habitats 100 m surrounding wetlands were also significantly and positively influenced by the water body. T. aegyptiaca favoured open habitats surrounding wetlands (Sirami et al., 2013: 35). 568 ISSN 1990-6471 HABITS: Rautenbach and Nel (1980: 112) indicate that this is fast flying bat which follows a relatively straight flight path. Thomas and Jacobs (2013: 129) report that, at De Hoop Nature Reserve, South Africa, bats feeding on Diptera (such as T. aegyptiaca) emerged later than bats feeding on Lepidoptera (e.g. Rhinolophus capensis and Nycteris thebaica). ROOST: In the Durban area (RSA), Taylor et al. (1999: 68) found these bats roosting in high-rise buildings of the Central Business District, and the colony sizes varied between 30 and an estimated 300. Toussaint et al. (2010) examined temperature in Pretoria, South Africa. roost DIET: In Zimbabwe, Fenton and Thomas (1980: 86) report that the diet of T. aegyptiaca consists of Lepidoptera and Coleoptera. From the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) reported the following prey groups based on 314 faecal pellets from 43 bats (in volume percent): Diptera (34.6 ± 33.4), Coleoptera (25.5 ± 30.7), Hemiptera (19.2 ± 24.6), Hymenoptera (8.4 ± 22.2), Lepidoptera (5.4 ± 13.4), Ephemeroptera (2.7 ± 7.7), Neuroptera (1.5 ± 6.5), Orthoptera (0.6 ± 3.4), and unknown (1.4 ± 7.1). For Saudi-Arabia, Benda et al. (2012a: 524) indicate that some diet samples were dominated by Lepidoptera, whereas others were dominated by Coleoptera and Hymenoptera or by Heteroptera and Brachycera. In Iran, three digestive tracts were analysed: One tract was empty, one contained Lepidoptera (30 % of volume), Coleoptera (Scarabaeidae) (30 %), Neuroptera (Myrmeleontidae) (10 %), and Orthoptera (Ensifera) (30 %), and the third tract contained Coleoptera (Elateridae, Carabidae) (60 %) and Orthoptera (Ensifera) (40 %). PREDATORS: Avery et al. (2005: 1054) found this species to be present in pellets from Tyto alba in South Africa. Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- This species is widespread and common in its South Asian range, where its population seems to be stable (Molur et al., 2002). Trend:- 2016: Unknown (Monadjem et al., 2017bi). 2008: Unknown (Mickleburgh et al., 2008ar; IUCN, 2009). ACTIVITY AND BEHAVIOUR: Toussaint et al. (2010) examined heterothermy in T. aegyptiaca in Pretoria, South Africa. REPRODUCTION AND ONTOGENY: In Pretoria (South Africa), Tsita (1993), Bernard and Tsita (1995: 18) and le Grange et al. (2011: 170 - 171) found spermatogenesis occurring in January, and spermatozoa were first found in the epididymis in May, where they were stored until the end of September. In females, follicular development was already taking place in January. Copulation, ovulation and fertilization took place in August to early September; and births took place in December. Their conclusion was that T. aegyptiaca is a monoestrous, monotocous seasonal breeder. le Grange et al. (2011: 173) also indicate that, at 33° S, spermatogenesis only started in February, and that follicular development only started in April, but copulation, ovulation, and fertilization also occurred at the end of August. Monadjem et al. (2010d) [in Weier et al. (2018: Suppl.)] mentioned births in November or December. PARASITES: Copeland et al. (2011: 363) reported the rediscovery of the "terrible hairy fly" (Mormotomyia hirsuta (after 62 years) in guano produced by C. cf. bivittatus and T. aegyptiaca at Ukasi Hill, Kenya. From Algeria, the following ectoparasites were reported by Bendjeddou et al. (2017: 15): Nycteribiidae: Nycteribia (Nycteribia) pedicularia Latreille, 1805 and Phthiridium biarticulatum Hermann, 1804; Streblidae: Brachytarsina (Brachytarsina) flavipennis Macquart, 1851: and Arachnida: Spinturnix myoti (Kolenati, 1856) and Eyndhovenia euryalis (G. Canestrini, 1885). Orlova et al. (2020b: 1) reported the first case of an infection by Parasteatonyssus nyctinomi in Namibia (Macronyssidae, Arachnida). VIRUSES: Bunyaviridae Phlebovirus Rift Valley Fever Virus (RVFV) - Oelofsen and Van der Ryst (1999) tested 24 individuals from two localities in the Free State Province, South Africa for RVFV antigen using an enzyme linked immunosorbet assay (ELISA), none tested positive. African Chiroptera Report 2020 569 Rhabdoviridae Lyssavirus - Rabies related viruses Rabies - Oelofsen and Smith (1993) tested 55 individuals from four localities using "Trousse Platelia Rage" ELISA kit (Diagnostic Pasteur), none tested positive for antibodies to rabies virus glycoprotein G. Oelofsen and Smith (1993) tested 89 individual brains, from 9 localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Mauritius, Niger. Figure 202. Distribution of Tadarida aegyptiaca Tadarida aegyptiaca aegyptiaca (E. Geoffroy St.-Hilaire, 1818) *1818. Nyctinomus ægyptiacus E. Geoffroy Saint-Hilaire, Description des Mammifères qui se trouve en Egypte, 2: 128, pl. 2, No 2. Publication date: 1818. Type locality: Egypt: Giza province: Giza [30 01 N 31 13 E]. - Comments: Type locality restricted from "Egypt" to Giza by Koopman (1975: 422): see Qumsiyeh (1985: 73), Meester et al. (1986: 70). - Etymology: From the scientific Latin feminine adjective aegyptìacus, meaning "Egyptian", as the species was decribed from an Egyptian specimen (see Lanza et al., 2015: 319). 1827. Dysopes geoffroyi Temminck, Monogr. Mamm, 1: 226, pls 19, 23. Publication date: 1827. - Comments: Allen (1939a: 110), Qumsiyeh (1985: 73) mentioned 1827 as year of publication. Kock (1969a: 137), Meester et al. (1986: 70) mention 1826. Substitute for aegyptiacus E. Geoffroy St. Hilaire (see Ellerman and Morrison-Scott, 1951: 135; Harrison, 1964a: 109; Kock, 1969a: 137; Qumsiyeh, 1985: 73; Meester et al., 1986: 70). 1877. Dysopes talpinus Heuglin, Reise in Nordost Afrika, 2: 28. Publication date: 1877. Type locality: Sudan: Bahr-el-Ghazal province: Kidj district: Bahr el Jebel, West bank: "Kidj district" [ca. 07 06 N 30 46 E]. 1916. Nyctinomus (Nyctinomus) tongaënsis Wettstein, Anz. kais. Akad. Wiss., Wien, 53: 192. Publication date: 1916. Type locality: Sudan: Upper White Nile: Tonga [09 30 N 31 03 E] [Goto Description]. Syntype: NMW 8406: ♂, alcoholic (skull not removed). Collected by: Prof. Dr. Franz Werner; collection date: 16 April 1916. Presented/Donated by: Otto Ritter von Westersheim Wettstein. See Benda et al. (2012a: 582). Syntype: NMW 8407: ♂, skull and alcoholic. Collected by: Prof. Dr. Franz Werner; collection date: 16 April 1916. Presented/Donated by: Otto Ritter von Westersheim Wettstein. See Benda et al. (2012a: 582). - Comments: Wettstein (1916: 189, 192) mentions two male type specimens, collected by Prof. Dr. Franz Werner on 16 April 1914. 1938. [Nyctinomus] tongaënsis: Frechkop, Exploration du Parc National Albert, 54. (Name Combination) ? Tadarida aegyptiaca aegyptiaca: (Name Combination, Current Combination) ? tongaensis: (Current Spelling) GENERAL DISTRIBUTION: Arabia, N Africa (Horácek et al., 2000: 136). Transvaal Provinces of South Africa, Lesotho and Botswana (Haeselbarth et al., 1966: 189). PARASITES: SIPHONAPTERA Ischnopsyllidae: Araeopsylla scitula (Rothschild 1909) found in southern Africa from Cape, Natal, DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Côte d'Ivoire, Egypt, Ethiopia, Guinea, Kenya, Mali, Morocco, Nigeria, South Africa, South Sudan, Sudan, Tanzania, Uganda. 570 ISSN 1990-6471 Tadarida aegyptiaca bocagei (Seabra, 1900) 1900. Nyctinomus Anchietæ Seabra, J. Sci. mat. phys. nat., ser. 2, 6: 82. Publication date: August 1900. Type locality: Angola: W Angola: Quibula [Goto Description]. *1900. Nyctinomus Bocagei Seabra, J. Sci. mat. phys. nat., ser. 2, 6: 84. Publication date: August 1900. Type locality: Angola: Benguela district: East of Hanha: Galanga [12 04 S 15 09 E] [Goto Description]. 1900. Nyctinomus brunneus Seabra, J. Sci. mat. phys. nat., ser. 2, 6: 83. Publication date: August 1900. Type locality: Angola: NE of Benguela: Quissange [12 30 S 14 05 E] [Goto Description]. 1941. Tadarida bocagei: Hill and Carter, Bull. Am. Mus. Nat. Hist., 78 (1): 55. Publication date: 25 July 1941. (Name Combination) ? Tadarida (Tadarida) aegyptiaca bocagei: (Name Combination) ? Tadarida aegyptiaca bocagei: (Name Combination, Current Combination) TAXONOMY: If page precedence is to be taken into account, anchietae might be the senior synonym for this form. COMMON NAMES: Afrikaans: Bocagese Losstertvlermuis. Bocage's Free-tailed Bat. English: GENERAL DISTRIBUTION: Angola and Zambia to South Africa (Horácek et al., 2000: 136). PREDATORS: Dean (1975: 218) reports on remains being found in barn owl (Tyto alba) pellets in South Africa. PARASITES: HEMIPTERA Cimicidae: Stricticimex transverses Ferris and Usinger, 1957 from Kanye, Botswana (Haeselbarth et al., 1966: 15). SIPHONAPTERA Ischnopsyllidae: Araeopsylla scitula (Rothschild, 1909) found in southern Africa from Cape, Natal, Transvaal Provinces of South Africa, Lesotho and Botswana (Haeselbarth et al., 1966: 189). ACARI Trombiculidae: Microtrombicula kanyei Vercammen-Grandjean, 1965 was reported by Stekolnikov (2018a: 155) from bats from Kanye, Botswana. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Botswana, Eswatini, Lesotho, Malawi, Mozambique, Namibia, South Africa, Zambia, Zimbabwe. Tadarida fulminans (Thomas, 1903) *1903. Nyctinomus fulminans Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (71): 501. Publication date: 1 November 1903. Type locality: Madagascar: Eastern Betsileo: Fianarantsoa [21 36 S 47 05 E, 1 130 m] [Goto Description]. Holotype: BMNH 1882.3.1.34: ♂, skull and alcoholic. Collected by: Rev. W. Deans Cowan; original number: 4. - Etymology: From the Latin "fulminate" meaning strike with lightning. 1946. Nyctinomus mastersoni Roberts, Ann. Transv. Mus., 20 (4): 306. Type locality: Zimbabwe: Bindura district: Masembura Nature Reserve: Chikupu caves [17 24 S 31 20 E] [Goto Description]. Holotype: TM 9976: ad ♂, skin and skull. Collected by: H.B. Masterson; collection date: 10 July 1945. Topotype: ROM 64580: ad ♂, skin and skull. Collected by: Randolph Lee Peterson and G.E. Turner; collection date: 11 May 1972. Presented/Donated by: ?: Collector Unknown. 2004. Tadarida fulmilans: Goodman and Cardiff, Acta Chiropt., 6 (2): 227. (Lapsus) ? Tadarida fulminans mastersoni: (Name Combination) ? Tadarida fulminans: (Name Combination, Current Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Afrikaans: Madagaskarse grootlosstertvlermuis. Chinese: 岛 犬 吻 蝠 . Czech: morous východoafrický. English: Large Free-tailed Bat, Lightning Guano Bat, Large Guano Bat, African Chiroptera Report 2020 Madagascan Large Guano Bat, Madagascan Large Free-tailed Bat, Madagascan Free-tailed Bat, Madagascar Large Free-tailed Bat, Malagasy Free-tailed Bat. French: Tadaride de Thomas, Grande Tadaride de Madagascar, Molosse de Madagascar. German: Madagassische Bulldoggfledermaus. ETYMOLOGY OF COMMON NAME: The colloquial name refers to the island where the species occurs. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its wide range across east and southern Africa. Although it is not a common species and can be patchily distributed, it is not thought to be facing any major threats (Cotterill et al., 2008; IUCN, 2009; Monadjem et al., 2017bu). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bu). 2008: LC ver 3.1 (2001) (Cotterill et al., 2008; IUCN, 2009). 2004: LC ver 3.1 (2001) (Cotterill, 2004b; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016z). 2004: Considered a vagrant. Not assessed, due to known form a single locality (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: There appear to be no major threats to this species (Cotterill et al., 2008; IUCN, 2009; Monadjem et al., 2017bu). CONSERVATION ACTIONS: Cotterill et al. (2008) [in IUCN (2009)] and Monadjem et al. (2017bu) report that it has been recorded from one protected area in Madagascar and is presumed to exist in many protected areas within its range in eastern and southern Africa, however, there appear to be no active conservation measures in place. Further studies are needed into the distribution and taxonomic status of bats allocated to Tadarida fulminans. GENERAL DISTRIBUTION: Tadarida fulminans ranges through East Africa, southern Africa and a few localities on the island of Madagascar. Populations have been recorded through eastern and southeastern Africa, from around the border of Kenya and Uganda as far south as the border between Zimbabwe and South Africa. The most westerly record is from eastern 571 Democratic Republic of the Congo. In Madagascar, it is only known from a few records, mostly from the central-south region near to Fianarantsoa and Isalo National Park, and from records at Tolagnaro near the southeast coast (Jenkins et al., 2007a). Further surveys are needed within Madagascar (Goodman and Cardiff, 2004). It ranges from about sea level (at Fort Dauphin in Madagascar) to close to 2,000 m asl (Albertine Rift). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 541,523 km2. Native: Congo (The Democratic Republic of the); Kenya; Madagascar; Malawi (Happold et al., 1988; Monadjem et al., 2010d: 546); Mozambique (Monadjem et al., 2010d: 546; Monadjem et al., 2010c: 382); Rwanda; South Africa (Monadjem et al., 2010d: 546); Tanzania; Zambia (Ansell, 1957; Ansell, 1967; Monadjem et al., 2010d: 547); Zimbabwe (Monadjem et al., 2010d: 547). ECHOLOCATION: See Taylor (1999b). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Peterson and Nagorsen (1975) reported 2n = 48, FN = 54, BA = 8, a submetacentric X chromosome, and an acrocentric Y chromosome, for specimens from Zimbabwe. However, Rautenbach et al. (1993) reported 2n =48, FN =66, BA =20, a metacentric X chromosome and an acrocentric Y chromosome for specimens from South Africa. Protein / allozyme - Unknown. POPULATION: Structure and Density:- Locally common, but patchily distributed in Africa. On Madagascar this bat is an uncommon, patchily distributed species. Colonies of this species may consist of dozens of individuals (although always fewer than 100 animals) (Cotterill et al., 2008; IUCN, 2009; Monadjem et al., 2017bu). Trend:- 2016: Stable (Monadjem et al., 2017bu). 2008: Stable (Cotterill et al., 2008; IUCN, 2009). PARASITES: Polyctenidae: Hypoctenes fani Benoit 1958, a single female collected from Rwanda (Haeselbarth et al., 1966: 18). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Kenya, Madagascar, Malawi, Mozambique, 572 ISSN 1990-6471 Namibia, Rwanda, Zambia, Zimbabwe. South Africa, Tanzania, Figure 203. Distribution of Tadarida fulminans Tadarida lobata (Thomas, 1891) *1891. Nyctinomus lobatus Thomas, Ann. Mag. nat. Hist., ser. 6, 7 (39): 303. Publication date: 1 March 1891. Type locality: Kenya: West Pokot: upper reaches of Turkwell: Turkwell Gorge, at or near [ca. 01 52 N 35 22 E] [Goto Description]. Holotype: BMNH 1893.2.3.7: ♂, skull and alcoholic. Collected by: Sir Frederic John Jackson. Peterson and Harrison (1970: 1) indicate that this specimen is probably a male, although they could not find any trace of scrotal development. - Etymology: From "lobata" meaning lobed. ? Tadarida (Chaerephon) pumila limbata: ? Tadarida lobata: (Name Combination, Current Combination, Current Spelling) ? Tadarida pumila limbata: TAXONOMY: See Simmons (2005). (Cotterill, 2008e; IUCN, 2009; Monadjem and Cotterill, 2017). COMMON NAMES: Afrikaans: Grootoor-losstervlermuis. Chinese: 大 耳 犬 吻 蝠 . Czech: morous širokouchý. English: Big-eared Guano Bat, Big-eared Kenyan Guano Bat, Kenyan Big-eared Free-tailed Bat, Bigeared Free-tailed Bat, Big-eared Kenya Free-tailed Bat, Eastern Africa Free-tailed Bat. French: Tadaride du Kenya, Tadaride de l'Afrique orientale, Molosse à grandes oreilles. German: Großohr-Bulldoggfledermaus. Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Cotterill, 2017). 2008: LC ver 3.1 (2001) (Cotterill, 2008e; IUCN, 2009). 2004: DD ver 3.1 (2001) (Cotterill, 2004e; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). ETYMOLOGY OF COMMON NAME: The colloquial name perhaps refers to the country from which the species was originally described (see Taylor, 2005). MAJOR THREATS: There appear to be no identifiable threats to this species (Cotterill, 2008e; IUCN, 2009; Monadjem and Cotterill, 2017). CONSERVATION STATUS: Global Justification Although this species is known mainly from isolated records from a large area, it is listed as Least Concern (LC ver 3.1 (2001)) because it is associated with rocky areas in dry savanna habitat, and is unlikely to be declining fast enough to qualify for listing in a more threatened category CONSERVATION ACTIONS: Cotterill (2008e) [in IUCN (2009)] and Monadjem and Cotterill (2017) report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further surveys are needed to better determine the range of this species, and to Regional None known. African Chiroptera Report 2020 573 understand whether it is restricted to rocky areas or can persist in modified habitats. Pellérdy (1974) mentioned the bat as "Tadarida limbata" this might not be correct. GENERAL DISTRIBUTION: Tadarida lobata has only been recorded from four localities in Kenya and four localities in Zimbabwe (Skinner and Chimimba, 2005). It is found between 600 and 2,000 m asl. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, Tanzania, Zimbabwe. Native: Kenya; Zimbabwe (Monadjem et al., 2010d: 547). POPULATION: Structure and Density:- It appears to be a rare species found in small colonies (Cotterill, 2008e; IUCN, 2009; Monadjem and Cotterill, 2017). The apparent rarity might be attributed to their highflying habits and inaccessible daylight roosts (Cotterill, 1996b). Trend:- 2016: Unknown (Monadjem and Cotterill, 2017). 2008: Unknown (Cotterill, 2008e; IUCN, 2009). Figure 204. Distribution of Tadarida lobata PARASITES: Duszynski (2002: 20) indicates that T. lobata might be a host for Eimeria dukei Lavier, 1927, but as Tadarida teniotis (Rafinesque, 1814) *1814. Cephalotes teniotis Rafinesque, Précis des découvertes et travaux somiologiques, 12, 55. Publication date: 3 June 1814. Type locality: Italy: Sicily [Goto Description]. Etymology: From the Greek "teniotis" meaning striped, banded. 1825. Dinops Cestonii Savi, Nuov. Giorn. Letter. Pisa, Sci., 10: 230. Type locality: Italy: Tuscany: Pisa [43 43 N 10 24 E]. Syntype: ZFMK MAM-1983.0173: ♂, alcoholic (skull not removed). Collected by: Prof. Paulo Savi; collection date: 1846. On label: "Nyctinomus cestonii (Savi) Dobson (In der Sammlg. Von Berthold als Dysops Savii angegeben) Toscana"; ex Göttingen Museun 1978: see Hutterer (1984: 32). 1827. Dysopes rüpelii Temminck, Monogr. Mamm, 1: 224, pl. 18. Publication date: 1827. Type locality: Egypt: Cairo [30 03 N 31 15 E]. Lectotype: SMF 12380: skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell. Presented/Donated by: ?: Collector Unknown. Locality: Egypt original number: II.K.1.b. Paralectotype: SMF 12379: skin and skull. Collected by: Wilhem Peter Edward Simon Rüppell. Presented/Donated by: ?: Collector Unknown. Locality: Egypt old catalog number: II.K.1.a. - Comments: Type locality restricted from "Egypt" to Cairo by Qumsiyeh (1985: 71). Incorrect original spelling (rüpelii) corrected by Qumsiyeh (1985: 71) as rueppellii. 1840. Dysopes Savii Schinz, Europäische Fauna oder Verzeichniss der Wirbelthiere Europa's, 1: 138. - Comments: Substitute for cestoni (see Ellerman and Morrison-Scott, 1951: 134; Harrison, 1964a: 106). - Etymology: In honour of Gaetano Savi (1769 - 1844) (see Kozhurina, 2002: 17). 1855. D[ysopes] Rüppelli: Giebel, Die Säugethiere., 957. (Name Combination) 1855. Dysopes cestonii: Wagner, in: Schreber, Die Säugethiere in Abbildungen nach der Natur, mit Beschreibungen, Suppl. 5: 702. 1876. Nyctinomus cestonii: Dobson, Monogr. Asiatic Chiropt, 180. 1877. Dysopes rüppellii: Dobson, Proc. zool. Soc. Lond., 1876, IV: 719. Publication date: April 1877. - Comments: Correction of Temminck's Dysopes rüpelii. 1891. Nyctinomus taeniotis: Thomas, Proc. zool. Soc. Lond., 1891, II: 182. (Lapsus) 574 ISSN 1990-6471 1897. 1914. 1985. 1988. 2018. ? ? ? ? ? midas Schulze, Helios, Berlin, 14: 95. - Comments: Preoccupied by midas Sundevall, 1843 (See Pavlinov et al., 1995: 91). Cephalotes tæniotis: Cabrera, Fauna Ibérica, 145. (Lapsus) rueppellii Qumsiyeh, Spec. Publ. Mus. Texas Tech. Univ., 23: 71. - Comments: Emendation of rüpelii Temminck, 1827. (Emendation) Tadarida taeniotis: Estrada-Peña and Sánchez, Rev. Ibér. Parasitol., 48 (3): 310. (Lapsus) Tadarida tenioti: Lazzeroni, Burbrink and Simmons, Ecol. Evol., 8 (24): 12584. Publication date: 28 November 2018. (Lapsus) Tadarida (Tadarida) teniotis rueppelli: (Name Combination) Tadarida (Tadarida) teniotis: (Name Combination) Tadarida teniotis rueppelli: (Name Combination, Alternate Spelling) Tadarida teniotis rueppellii: (Name Combination) Tadarida teniotis: (Name Combination, Current Combination) TAXONOMY: Revised by Aellen (1966a) and Kock and Nader (1984). Populations in Japan, Taiwan and Korea are now treated as a separate species, T. insignis (Simmons, 2005). COMMON NAMES: Albanian: Lakuriq nate bishtlirë, Lakuriq nate bishtIire. Arabian: Khaffash. Armenian: Լայնականջ ծալքաշուրթ չղջիկ. Azerbaijani: Bükükdodaq enliqulaq. Basque: Saguzar buztanluze europar, Saguzar buztanluzea. Belarusian: Тадарыда еўрапейская. Bosnian: Golorepi šišmiš. Breton: Molos Cestoni. Bulgarian: Булдогов прилеп. Castilian (Spain): Murciélago rabudo. Catalan (Spain): Ratpenat cuallarg, Rat penat cuallarg. Croatian: Sredozemni slobodnorepac. Czech: Tadarida evropská, Morous evropský, Hroznohled pyskatý, Psohubec myšowý, Netopýr vrásopyskatý. Danish: Europæisk bulldogflagermus. Dutch: Zuideuropese bulvleermuis, Bulveermuis, Europese bulvleermuis. English: European Free-tailed Bat, European Guano Bat, Striped or Banded Guano Bat, Wrinkle-lipped Bat, Eurasian Free-tailed Bat. Estonian: Euroopa kurdmokk. Finnish: Doggilepakko. French: Molosse de Cestoni, Tadaride d'Europe, Molosse d'Europe, Tadaride de Cestoni. Frisian: Muskusbulflearmûs. Galician (Spain): Morcego rabudo. Georgian: ევროპული ტადარიდა. German: Bulldoggfledermaus, Europäische Bulldoggfledermaus. Greek: Νυχτονόμος. Hebrew: Pashaf Matsui. Hungarian: Európai szabadfarkú-denevér, Nycteris i urophoros. Irish Gaelic: Ialtóg earrshaor Eorpach. Italian: Molosso del Cestoni. Latvian: Brivastes sikspārnis. Lithuanian: Europinis raukšlėtalŪpis. Luxembourgish: Europäesch Bulldogfliedermaus. Macedonian: Опашест лилјак [= Opashest Liljak]. Maltese: Tadarida. Montenegrin: Dugorepi slijepi miš. Norwegian: Middelhavsfrihale. Polish: Molos europejski. Portuguese: Morcego-rabudo. Rhaeto-Romance: Moloss buldoc. Romanian: Lilacul cu coadă liberă europeană. Russian: Складчатогуб широкоухий. Serbian: Средоземни репаш [= Sredozemni repaš], Широкоухий складчатогуб. Scottish Gaelic: Ialtag eàrr-shaor Eòrpach. Slovak: Tadarida buldogovitá. Slovenian: Dolgorepi netopir. Swedish: Veckläppad fladdermus. Turkish: Avrupa Serbest Kuyruklu Yarasası. Ukrainian: Тадарида європейська. Welsh: Ystlum cynffonog. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Amorim et al. (2013: 7) indicate that palaeodistribution models predict that during the Last Glacial Maximum (LGM: 23,000 - 18,000 YBP) the climatic conditions for T. teniotis were favorable in southern Europe and northern Africa. CONSERVATION STATUS: Global Justification The species is widely distributed over a large extent of occurrence. It occurs in urban areas and forages in other modified habitats. Population trends are not known but are not believed to approach the threshold for the population decline criterion of the IUCN Red List. Consequently, it is assessed as Least Concern (LC ver 3.1 (2001)) (Aulagnier et al., 2008e; IUCN, 2009; Benda and Piraccini, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Benda and Piraccini, 2016). 2008: LC ver 3.1 (2001) (Aulagnier et al., 2008e; IUCN, 2009). 1996: LR: lc (Baillie and Groombridge, 1996). Regional None known. African Chiroptera Report 2020 MAJOR THREATS: It is negatively affected by disturbance and loss of roosts in buildings, and by use of pesticides. It is also potentially threatened by wind farms (GMA Europe Workshop, 2006), and deforestation affects the species in some parts of its range (Z. Amr pers. comm., 2005 [in Aulagnier et al. (2008e in IUCN, 2009]). However, none of these are considered to be major threats at present (Aulagnier et al., 2008e; IUCN, 2009; Benda and Piraccini, 2016). Sherwin et al. (2012: 174) report aerial hawking and long-distance dispersal to be risk factors related with climatic change. CONSERVATION ACTIONS: Aulagnier et al. (2008e) [in IUCN (2009)] and Benda and Piraccini (2016) report that it is protected by national legislation in a number of range states and receives international legal protection through the Bonn Convention (Appendix II and Eurobats Agreement) and Bern Convention in parts of its range where these apply. It occurs in several protected areas across its range. GENERAL DISTRIBUTION: Tadarida teniotis is mainly a Palaearctic species, although the south-eastern edge of its range extends into the Indomalayan region. It is well known in the Mediterranean basin, occuring from Portugal, Spain eastwards through southern Europe to the Balkans, Turkey, Israel, Palestine and Jordan. In North Africa it has been recorded from Morocco, Algeria, Tunisa, Libya and Egypt. It is possibly present on Madeira (to Portugal) as there was a supposed old record, but it has not been recorded from there again. It occurs on all the Canary Islands (to Spain) except for Fuerteventura and Lanzarote. It is also recorded from a number of Mediterranean islands (Hutson, 1999; Simmons, 2005). It occurs from sea level to 3,100 m. Populations in Japan, Taiwan and Korea are now considered to be a separate species, T. insignis (Simmons, 2005). Native: Afghanistan; Albania; Algeria (Horácek et al., 2000: 136); Andorra; Armenia; Azerbaijan; Bangladesh; Bhutan; Bosnia and Herzegovina; Bulgaria; Croatia; Cyprus; Egypt; France [Corse]; Georgia; Gibraltar; Greece [Kriti]; Holy See (Vatican City State); India; Iran, Islamic Republic of; Iraq; Israel; Italy [Sardegna, Sicilia]; Jordan; Kazakhstan; Kyrgyzstan; Lebanon; Libyan Arab Jamahiriya (Horácek et al., 2000: 136); Macedonia, the former Yugoslav Republic of; Malta; Monaco; Montenegro; Morocco (Benda et 575 al., 2010a: 161; El Ibrahimi and Rguibi Idrissi, 2015: 364); Myanmar; Nepal; Portugal; Russian Federation; San Marino; Saudi Arabia; Serbia; Spain [Baleares, Canary Is.]; Switzerland; Syrian Arab Republic; Tajikistan; Tunisia (Horácek et al., 2000: 136; Dalhoumi et al., 2014: 53; 2016b: 867; 2019b: 26); Turkey; Turkmenistan; Uzbekistan. Presence uncertain: China. FUNCTIONAL MORPHOLOGY: Maniakas and Youlatos (2012: 574) studied the external and muscular forelimb features that could be associated with high-speed flight in open air. They found (p. 576) that the mass of the muscles responible for the downstroke represented 59.3 % of the total forelimb musculature, and that more than half of these muscles are formed by the M. pectoralis. The muscles responsible for the upstroke are comparatively small, which might indicate that the upstroke in this bat is less important than for other bat groups (p.580). All these features result in a fast, steady enduring flight with relatively limited maneuverability. ECHOLOCATION: For 10 Greek specimens, Papadatou et al. (2008b: 133) report a start frequency of 15.4 ± 3.28 kHz, a terminating frequency of 11.1 ± 1.34 kHz, a peak frequency of 13.2 ± 1.18 kHz, and a bandwidth of 5.3 ± 1.49 kHz. The duration of the call is 18.4 ± 4.74 msec, and the interpulse interval is 764.6 ± 259.37 msec. Walters et al. (2012: suppl.) report the following figures for 35 calls from bats from France, Greece, Italy and Switzerland: duration: 15.03 ± 3.47 msec, Fmax: 14.61 ± 4.95 kHz, Fmin: 10.64 ± 1.51 kHz, bandwidth: 3.97 ± 3.76 kHz, Fpeak: 11.76 ± 2.28 kHz. In Jordan, Benda et al. (2010b: 204) recorded the following parameters for seven calls: Fstart: 15.7 ± 3.8 (12.4 - 22.6) kHz, Fend: 9.8 ± 0.4 (9.4 - 10.5) kHz, Fpeak: 11.7 ± 0.4 (11.1 - 12.6) kHz, duration: 16.3 ± 2.2 (13.1 - 19.3) msec, and interpulse duration: 571.8 ± 254.6 (163.0 - 800.0) msec. 14 calls from Iranian specimens were reported by Benda et al. (2012a: 183): Fstart:19.0 ± 2.6 (14.1 22.2) kHz, Fend: 11.8 ± 2.2 (8.9 - 14.2) kHz, Fpeak: 14.5 ± 1.3 (10.6 - 15.9) kHz, duration: 10.6 ± 0.8 (9.5 - 11.8) msec, and interpulse interval: 368.7 ± 89.8 (266.0 - 484.0) msec. Hackett et al. (2016: 223) recorded 44 calls in Israel with the following values: Pulse duration: 12.17 ± 3.34 msec, Fstart: 22.73 ± 5.02 (17.5 - 29.2) kHz, Fend: 15.90 ± 1.44 (13.1 - 17.5) kHz, Fpeak: 18.49 ± 1.57 (15.1 - 21.4) kHz. Disca et al. (2014: 226) indicate that in Morocco the type of call is FM-QFC, with the following 576 ISSN 1990-6471 parameters: Fstart: 22.3 ± 4.2 kHz, Fend: 13.3 ± 0.7 kHz, Fpeak: 14.4 ± 0.8 kHz, bandwidth: 9.0 ± 3.9 kHz, duration: 18.7 ± 2.7 msec. Luo et al. (2019a: Supp.) reported the following data (Free flying bats; 2 calls): Fpeak: 13.2, 13 kHz, Fstart: 15.4, 17 kHz, Fend: 11.1, 12.1 kHz, Band width: 4.3, 4.9 kHz, and duration: 18.4, 16.6 msec. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Ðulic and Mrakovcic (1980) reported 2n = 48, FN = 76, BA = 30, a submetacentric X chromosome and a subtelocentric Y chromosome for one male from Kroatia. Arroyo Nombela et al. (1986), however, reported 2n = 48, FN = 78, BA = 32 , a submetacentric X and an acrocentric Y for specimens from Spain. Volleth and Eick (2012: 167) mention 2n = 48 and a segment number of 25. The autosomal complement consists of five meta- and submetacentrics, and 19 subtelocentric and acrocentric pairs, of which the smallest is dotlike (Arslan and Zima, 2014: 14). These authors also indicate that the X chromosome was found to be subtelocentric in some studies. Combining mtDNA fragments, 14 microsatellites and ecological modelling tools, Amorim et al. (2017: 38) found three main haplogroups in Mediterranean T. teniotis: one restricted to the eastern Mediterranean and two that are widespread across the Iberian Peninsula and Morocco. The nuclear data indicated that the bats from the Canary Islands showed the lowest signs of gene flow with the continental populations. Furthermore, they found that gene flow between Morocco and the central Mediterranean area is higher than that between the Iberian Peninsula and the central Mediterranean. Amorim et al. (2019: 1) confirmed that the nuclear DNA of bats from the Canary Islands is highly differentiated and isolated from that the two other genetic populations they distinguished: one from Morocco, Iberia and France, and the other from Italy eastwards to Anatolia and the Middle East. Mata et al. (2017: 152) reported the mitogenome to be 16,869 bp long, containing 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNA genes, and 1 control region (d-loop). All proteincoding genes start with the ATG start codon, except for ND2, ND3, and ND5 which begin with ATA or ATT. Eleven protein-coding genes terminated in a canonical stop codon, TAA or TAG, two contain incomplete stop codons, T or TA. Cytochrome b terminates in the mitochondriaspecific stop codon AGA. Protein / allozyme - Unknown. HABITAT: In her study on the summer foraging habitats of bats in the Mediterranean region of the Iberian Peninsula, Rainho (2007: 176) found that T. teniotis showed low activity over fresh-water habitats, and primarily seemed to use pine-tree woodlands and Mediterranean shrublands. For southern Portugal, Marques et al. (2004: 99) reported that T. teniotis foraged preferentially over forested areas, particularly pine and cork oak woodlands. They used both alluvial plains and the valleys of a mountainous area, but not its ridges. HABITS: In southern Portugal, Marques et al. (2004: 99) found that T. teniotis is a late emerger, leaving its roost about one hour after sunset. They also found that these bats have only one foraging bout, which lasted on average 6 hours and 39 minutes. Its flight speed can reach over 50 km/h, and flying can be maintained for over 10 hours. Ancillotto and Russo (2014: 221) found that that T. teniotis is able to discriminate between familiar and stranger individuals. They also found that the animals show higher rates of aggressive behaviours towards the strangers. Females show less aggressiveness than males, especially towards other females, and both sexes behave less aggressive towards juveniles. ROOST: In Libya, Benda et al. (2014c: 119) found a roosting colony in a fissure in the ceiling in the portal of a large, but shallow cave DIET: Rydell and Arlettaz (1994: 175) suggest that the intense narrowband echolocation calls with low frequency (11 - 12 kHz) without strong harmonics emitted by T. teniotis seems to be a specialization for long-range detection of large, tympanate insects such as Lepidoptera and Neuroptera, which are less well represented in the diet of most other aerial-hawking bats. Based on faecal analyses, they found the following volume percents: southeastern France: Lepidoptera (68.3 %), Neuroptera (24.3 %), others (4.3 %), Acari (0.5 %), and unidentified (2.6 %); Kirghizstan: Lepidoptera (86.8 %), others (12.8 %), unidentified (0.4 %). Benda et al. (2014c: 124) examined four sets of faecal pellets from Libya. Ten pellets from Wadi an Nazrat contained only large moths (Lepidoptera) as did one pellet from the quarry at Wadi al Kuf. Twenty pellets from another location at Wadi al Kuf contained 97 % by volume medium- African Chiroptera Report 2020 577 sized moths and 3 % beetles (Coleoptera). Eight pellets from one animal from Wadi Darnah contained 56 % by volume medium-sized moths and 44 % lacewings (Neuroptera: Chrysopidae). 1.4); Diptera (10.1, 12.2), Hemiptera (2.9, 8.1), Neuroptera (7.2, 12.2), unidentified Insecta (1.4, 1.4), and also some unidentified Arthropoda (7.2, 5.4). Benda et al. (2012a: 515) indicate that Orthoptera formed the largest contributing group in the diet of T. teniotis in Turkey and Jordan. On Rhodes, Coleoptera (especially Curculionidae) formed the most important group. In Iran, Lepidoptera were the most important diet item and Coleoptera, Orthoptera (Ensifera) and Heteroptera also contributed substantially to the diet. Neuroptera (Chrysopidae) and Auchenorrhyncha were almost negligible. PREDATORS: In Isreal, Reuven (1991: 79) observed T. teniotis being preyed upon by a Lanner Falcon (Falco biarmicus Temminck, 1825). Balmori (2012: 9) furthermore reports bats being taken by Peregrine Falcon (Falco peregrinus Tunstall, 1771) and Common Kestrel (Falco tinnunculus Linnaeus, 1758) as well as by Barn Owl (Tyto alba (Scopoli, 1769)) and Tawny Owl (Strix aluco Linnaeus, 1758), but also by a Garden Dormouse (Eliomys quercinus (Linnaeus, 1766)). Mikula et al. (2016: Supplemental data) mention the following diurnal avian predators: Lanner falcon (Falco biarmicus Temminck, 1825), Peregrine falcon (Falco peregrinus Tunstall, 1771) and Eurasian hobby (Falco subbuteo Linnaeus, 1758), In northeastern Portugal, Mata et al. (2016: 1, 2) found significant dietary differences between males and females, where females ate more larger moths and more moths with migratory behaviour (* in below listing). In guano pellets of 69 females and 74 males, Lepidoptera were present in 98.6 % of the females pellets and in 100 % of the males pellets. Split over the various familes and species (which all occur in northern African too), they found (wingspan; % in females, % in males): Crambidae: Nomophila noctuella* (Denis and Schiffermüller, 1775) (rush veneer) (26 - 32 mm; 7.2, 14.9), other Crambidae 6 5.8 9.5; Gelechiidae: Mirificarma mulinella (Zeller, 1839) (5.5 - 7.5 mm; 7.2, 14.9), other Gelechiidae (0.0, 2.7); Geometridae: Aspitates ochrearia (Rossi, 1794) (yellow belle) (25 - 34 mm; 4.3, 10.8), Rhodometra sacraria* (Linnaeus, 1767) (vestal) (22 - 28 mm; 14.5, 36.5), other Geometridae (20.3, 13.5); Noctuidae: Agrotis ipsilon* (Hufnagel, 1766) (dark sword-grass) (40 45 mm; 21.7, 4.1), Agrotis puta (Hübner, 1803) (shuttle-shaped dart) (30 - 32 mm; 13.0, 18.9), Agrotis segetum (Denis & Schiffermüller, 1775) / Agrotis trux (Hübner, [1824]) (turnip/crescent dart) (32 - 42 mm; 37.7, 36.5), Autographa gamma* (Linnaeus, 1758) (silver y) (30 - 45 mm; 43.5, 23.0), Hoplodrina ambigua (Denis & Schiffermüller, 1775) (vine's rustic) (32 - 34 mm; 18.8, 35.1), Mythimna albipuncta (Denis & Schiffermüller, 1775) (white-point) (30 - 35 mm; 5.8, 9.5), Mythimna vitellina (Hübner, 1808) (delicate) (36 - 43 mm; 24.6, 28.4), Noctua pronuba (Linnaeus, 1758) / Noctua janthe Borkhausen, 1792 (large/lesser broad-bordered yellow underwing) (50 - 60 mm / 30 - 40 mm; 14.5, 13.5), Peridroma saucia* (Hübner, 1808) (pearly underwing) (43 - 50 mm; 24.6, 12.2), Phlogophora meticulosa* (Linnaeus, 1758) (angle shades) (45 52 mm; 18.8, 8.1), other Noctuidae (31.9, 36.5); Tortricidae: Tortrix viridana Linnaeus, 1758 (European oak leafroller) (18 - 23 mm; 5.8, 13.5), other Tortricidae (2.9 5.4); other Lepidoptera (17.4, 21.6); unidentifiedLepidoptera (13.0, 23.0). Other orders present in the pellets were: Coleoptera (1.4, POPULATION: Structure and Density:- It is a common species in suitable habitats. Summer and winter colonies typically number 5 - 100 individuals, although colonies of up to 300 - 400 animals have been recorded. It is probably sedentary, although seasonal in some areas (e.g., Malta). It is not abundant in the Caucasus, nor is it highly gregarious - large colonies are not known in this region (K. Tsytsulina pers. comm., 2005 in Aulagnier et al., 2008e [in IUCN, 2009) and Benda and Piraccini (2016)]). There are only six records for Iran, however, there have not been extensive survey efforts there (M. Sharifi pers. comm., in Aulagnier et al., 2008e [in IUCN, 2009] and Benda and Piraccini (2016)). Trend:- 2016: Unknown (Aulagnier et al., 2008e). 2008: Unknown (Aulagnier et al., 2008e; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Suppl.) and Lagunas-Rangel (2019: 2) report a maximum longevity of 13 years. REPRODUCTION AND ONTOGENY: In Spain, Balmori (2003: 37) reported births taking place from the end of June to the beginning of July. In Portugal, Amorim et al. (2015: 228) found that drought had a severe negative influence on the reproduction of T. teniotis. This drought probably resulted in reduced food resources early in the prebreeding season, which limited the ability to restore the bat's body condition after the winter and before the breeding season. 578 ISSN 1990-6471 MATING: In a colony in Valladolid, Spain, Balmori (2003: 37; 2017: 460) found that T. teniotis bats were organized in 'harems' made up by a dominant male together with a variable number of females, in which the male defends the refuge ("resource defence polygyny"). The males were emitting mating calls to attract passing-by females. These harems were formed between August and October. When competitors were present, the males reacted by noisy and fast chasing-flights and the releasing of a strong scent, probably used to mark their territory. Balmori (2017: 465) found two peaks of sexual activity: one in spring (Apri - –June) and another in summer/autumn (August - October), but also indicated that each female might only enter heat once a year. PARASITES: In Italy, Voyron et al. (2011: 193) reported the following fungal entities from live T. teniois specimens: Aspergillus fumigatus var. fumigatus and Trichosporon chiropterorum. Benda et al. (2010b: 315) only found the mite Parasteatonyssus hoogstraali Keegan, 1956 parasiting on Tadarida teniotis in Jordan. They do report on other ectoparasites found previously: the bat fly Nycteribia pedicularia Latreille, 1805 found in Palestine, bat fleas Araeopsylla wassifi Traub, 1954 in Turkey and Egypt, A. gestroi (Rotschild, 1906) in Lebanon, and a tick Argas vespertilionis (Latreille, 1802) in Egypt. Balmori (2012: 9) also mentioned Steatonyssus periblepharus Kolenati, 1858, Ewingana baekelandae Estrada-Peña, 1992, and unidentified Trombiculidae, as well als the bat bug Cimex pipistrelli Jenyns, 1839 and the flea Araeopsylla gestroi (Rothschild, 1906). 1886, Litomosa aelleni Tibayrenc, Bain & Ramachandran 1979, and further unidentified Litomosa sp. Benda et al. (2014c: 124) reported the following ectoparasites from Libya: Ischnopsyllidae: Araeopsylla gestroi (Rothschild, 1906), Macronyssidae: Parasteatonyssus hoogstraali (Keegan, 1956) [1st record from Libya], Spinturnicidae: Spinturnix myoti (Kolenati, 1856). From Algeria, Bendjeddou et al. (2017: 15) reported one ectoparasite: Arachnida: Ixodes ricinus (Linnaeus, 1758). VIRUSES: Herpesviridae Cytomegalovirus Nieto-Rabiela et al. (2019: Suppl.) mentioned this virus. Reoviridae Orthoreovirus This virus was reported by Kohl and Kurth (2014: 3113) from Italian bats. Rhabdoviridae Lyssavirus - Rabies related viruses European bat lyssavirus 1 was reported by Luis et al. (2013: Suppl.), Willoughby et al. (2017: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Egypt, Libya, Morocco, Sudan. Parasteatonyssus hoogstraali Keegan, 1956 was already reported by Estrada-Peña and Sanchez (1988: 310) from T. teniotis on Frontera, Canary Islands. The following enteroparasites were reported by Balmori (2012: 9): Trematodes Plagiorchis vespertilionis (Müller, 1784), Prosthodendrium ascidia (Van Beneden, 1873) and Lecithodendrium linstowi Dollfus, 1931; Nematodes: Molinostrongylus alatus (Ortlepp, 1932), Pseudophysaloptera formosana (Yokogawa, 1922), Rictularia bovieri Blanchard, Figure 205. Distribution of Tadarida teniotis Tadarida ventralis (Heuglin, 1861) *1861. Nyctinomus (Dysopes) ventralis Heuglin, Nov. Act. Acad. Cæs. Leop.-Carol., 29 (8): 4, 11. Publication date: 1861. Type locality: Eritrea: Kérén [ca. 15 45 N 38 20 E] [Goto African Chiroptera Report 2020 1862. 1876. 1877. 1877. 1891. 1901. 1928. 1939. 1951. 1953. 1974. 2015. ? ? 579 Description]. Lectotype: SMNS 982: ad ♀, skull and alcoholic. Collected by: Martin Theodor von Heuglin; collection date: 1861. Presented/Donated by: ?: Collector Unknown. See Dieterlen et al. (2013: 294), who confirm that the Lectotype status of this specimen was designated by Kock (1975: 6). - Etymology: From the scientific Latin feminine adjective ventràlis, meaning "ventral", referring to the presence of a wide, median, prominent orange ventral stripe (see Lanza et al., 2015: 324). Nyctinomus ventralis: Heuglin, Peterm. Geographische Mittheilungen, 8: 26. (Name Combination) Nyctinomus africanus Dobson, Ann. Mag. nat. Hist., ser. 4, 17 (101): 348. Publication date: 1 May 1876. Type locality: South Africa: Transvaal [Goto Description]. Holotype: BMNH 1875.11.19.1: ad ♂, skull and alcoholic. Presented/Donated by: Richard Browdler Sharp[e]. Dysopes ventralis: Heuglin, Reise in Nordost Afrika, 2: 26. (Name Combination) Nyctinomus [(Nyctinomus)] cestoni Dobson, Proc. zool. Soc. Lond., 1876, IV: 719. Publication date: April 1877. - Comments: Not of Savi, 1825. In part (see Kock, 1975: 6). N(yctinomus) taeniotis Thomas, Proc. zool. Soc. Lond., 1891, II: 183. Publication date: August 1891. - Comments: Not of Rafinesque, 1814. See Kock (1975: 7). N(yctinomus) midas de Winton, Ann. Mag. nat. Hist., ser. 7, 7 (37): 42. Publication date: 1 January 1901. - Comments: Not of Sundeval. In part. See Kock (1975: 7). Mops ventralis: Thomas, Ann. Mag. nat. Hist., ser. 10, 1 (2): 302. Publication date: 2 February 1928. (Name Combination) Mops rüppellii Allen, Bull. Mus. comp. Zool., 83: 108. - Comments: Not of Temminck, 1827. In part. See Kock (1975: 7). Tadarida africana: Roberts, Mammals of Southern Africa, 101. Tadarida (Tadarida) africana: Ellerman, Morrison-Scott and Hayman, Southern African mammals 1758 to 1951, 66. Tadarida (Tadarida) ventralis: Largen, Kock and Yalden, Mon. Zool. ital., (N.S.) 16 (Suppl. 5): 253. (Name Combination) Nyctinomus taeniatus: Lanza, Funaioli and Riccucci, The bats of Somalia and neighbouring areas, 323. (Lapsus) Tadarida ventralis africana: (Name Combination) Tadarida ventralis: (Name Combination, Current Combination) TAXONOMY: For use of this name in place of africana, see Kock (1975). COMMON NAMES: Afrikaans: Transvaalse losstertvlermuis, Reuse losstertvlermuis. Chinese: 非 洲 大 犬 吻 蝠 . Czech: morous africký, tadarida africká. English: Giant Guano Bat, Giant African Guano Bat, African Giant Free-tailed Bat, Giant African Free-tailed Bat, Transvaal Free-tailed Bat, Giant Free-tailed Bat. French: Tadaride géante d'Afrique, Grande tadaride africaine, Molosse géant. German: Große Bulldoggfledermaus, rothbraune Doggengrämler. Italian: Tadarìda ventràle. Portuguese: Morcego gigante de cauda livre. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Mickleburgh et al., 2008aj; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Mickleburgh et al., 2008aj; IUCN, 2009). 2004: NT ver 3.1 (2001) (Mickleburgh et al., 2004ao; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional South Africa:- 2016: VU D2 ver 3.1 (2001) (Taylor et al., 2016d). 2004: Considered a vagrant. Not assessed, known from a single locality (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: The threats to this species are poorly known. It may be threatened through the conversion of suitable habitat to agricultural land and the use of pesticides in these modified areas (Mickleburgh et al., 2008aj; IUCN, 2009). CONSERVATION ACTIONS: Mickleburgh et al. (2008aj) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. It is not known if the species is present in any protected areas. Further 580 ISSN 1990-6471 research is needed into the distribution, natural history, and threats to this little-known species. Trend:- 2008: Unknown (Mickleburgh et al., 2008aj; IUCN, 2009). GENERAL DISTRIBUTION: Tadarida ventralis is widely distributed in East and Southern Africa. It ranges from Eritrea in the north to Zimbabwe, and northern South Africa, in the south. The westernmost record is from the eastern Democratic Republic of the Congo, close to the border area with Uganda. Most records are from high altitude (1,500 m upwards to 2,500 m asl). REPRODUCTION AND ONTOGENY: In Malawi, young were born at the beginning of the wet season (Happold and Happold, 1990b: 568). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Eritrea, Ethiopia, Kenya, Malawi, South Africa, South Sudan, Tanzania, Zambia, Zimbabwe. Native: Congo (The Democratic Republic of the); Eritrea; Ethiopia; Kenya; Malawi (Ansell and Dowsett, 1988: 46; Monadjem et al., 2010d: 547); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 547); South Africa (Taylor et al., 2015: 50); Sudan; Tanzania; Zambia (Ansell, 1986; Monadjem et al., 2010d: 547); Zimbabwe (Cotterill, 2004a: 260; Monadjem et al., 2010d: 547). ECHOLOCATION: See Taylor et al. (2005). POPULATION: Structure and Density:- It has been rarely recorded and is largely known from single dead individuals (Mickleburgh et al., 2008aj; IUCN, 2009). Figure 206. Distribution of Tadarida ventralis Superfamily VESPERTILIONOIDEA Gray, 1821 *1821. Vespertilionoidea Gray, London Med. Repos., 15: 299. Publication date: 1 April 1821. (Current Combination) 1928. Vespertilionoidea: Weber, Die Säugetiere, 24. - Comments: Type genus: Vespertilio Linnaeus, 1758. Introduced as tribe (but actually similar to superfamily) and originally included the families Vespertilionidae J. Gray, 1821, Natalidae J. Gray, 1866, Myzopodidae Thomas, 1904 and Molossidae Gervais, 1855 (see Jackson and Groves, 2015: 265). TAXONOMY: Currently recognized families within the Superfamily VESPERTILIONOIDEA within Africa: CISTUGONIDAE Van Cakenberghe and Seamark, 2011; MINIOPTERIDAE Dobson, 1875; VESPERTILIONIDAE Gray, 1821. COMMON NAMES: Czech: netopýrov ci. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Ravel et al. (2011: 402) suggest that affinities such as the closure of the protofossa, the slender lophes, and a tiny metaconule on the upper molars (in the fossil genus Dizzya and an unnamed "Eochiropteran" from the El Kohol Formation in Algeria) could support an African endemic radiation of modern Vespertilionoidea. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Morocco, South Sudan. African Chiroptera Report 2020 581 Family CISTUGONIDAE Lack, Roehrs, Stanley, Ruedi and Van den Bussche, 2010 2008. 2010. 2010. Cistugoinae: Van Cakenberghe and Seamark, African Chiroptera Report, 213. Publication date: 1 July 2008. - Comments: Spelling used for subfamily. (Name Combination, Alternate Spelling) Cistugidae: Lack, Roehrs, Stanley, Ruedi and Van den Bussche, J. Mamm., 91(4): 980. (Alternate Spelling) Cistugonidae: Benda, Vespertilio, 18-19: 271. - Comments: Type genus: Cistugo Thomas, 1912. (Current Combination, Current Spelling) TAXONOMY: Stadelmann et al. (2004b) showed that the genus Cistugo was not placed with the genus Myotis, but was basal to the vespertilioid radiation, which is also supported by Horácek et al. (2006). For this reason, the ACR (2008, 2009, 2010) recognized representatives of the genus Cistugo as a separate distinct subfamily within the Vespertilionidae. Lack et al. (2010: 980) recognized that taxa within this group should be recognized at a family level and suggested the name “Cistugidae”. Benda (2010: 271) and ACR (2011) used a corrected form of the family (or subfamily) name: Cistugonidae (goninae) - according to the code (genitive from Cistugo is Cistugonis, thus Cistugonidae). Simmons and Cirranello (2020) refer to article 29.3.3 of the ICZN code to retain the usage of "Cistugidae" as family name, as this would indicate that "the stem for the formation of the family-group name is chosen by the author of the family level name". The article, however, explicitly mentions the "entire generic name", which would support the usage of "Cistugonidae". We therefore keep on using this name. Currently (Simmons and Cirranello, 2020) recognized genus within the family Cistugonidae: Cistugo Thomas, 1912. COMMON NAMES: Czech: žlázokřídlecovití. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The study by Amador et al. (2016: 26) supports the long-time separation from the Vespertilionidae, which probably occurred in the early middle Eocene (crown age 43 MYA). MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) mention the karyotype as 2n = 50 Genus Cistugo Thomas, 1912 *1912. Cistugo Thomas, Ann. Mag. nat. Hist., ser. 8, 10 (56): 205. Publication date: 1 August 1912 [Goto Description]. - Comments: Type species: Cistugo seabræ Thomas, 1912. (Current Combination) TAXONOMY: Meester et al. (1986) included Cistugo as a subgenus of Myotis. Baker et al. (1974) recorded the karyotype of Myotis as 2n = 44, AA = 50, but Dippenaar et al. (1983) gave it as 2n = 50, AA= 48 in both seabrai and lesueuri. Hayman and Hill (1971) doubt the validity of lesueuri and Corbet and Hill (1980) include it in seabrai, while Koopman (1982) retain it as a separate species. It is possible, particularly in view of their allopatric distribution, that lesueuri may best be treated as a subspecies of seabrai. Included as subgenus in Myotis by Koopman (1993a: 207), but considered a valid genus by Thomas (1915), Roberts (1919), Rautenbach et al. (1993), Bickham et al. (2004: 333), Stadelmann et al. (2004b: 185), and Simmons (2005). Stadelmann et al. (2004b: 185) state that it might be possible that the two genera share closer phylogenetic relationships with other families of bats than with the Vespertilionidae. Lack et al. (2010: 980) erected a new family: Cistugoidae (see family account above for information about corrected spelling of family name), containing only the genus Cistugo Thomas, 1912, which they separate from the Vespertilionidae by the presence of 2 to 4 glands in the wing membrane immediately behind the 582 ISSN 1990-6471 forearm (see also Seamark and Kearney, 2006), which are lacking in all other vespertilionoid bats. Currently (Simmons and Cirranello, 2020) recognized species of the genus Cistugo: lesueuri Thomas, 1912; seabrae Thomas 1912. COMMON NAMES: Czech: žlázokřídlecové. English: Wing-gland Bats. German: Mausohren. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Avery (2007: 619) reported on the presence of Cistugo sp. in Pleistocene deposits at Wonderwerk Cave, South Africa. GENERAL DISTRIBUTION: Seamark et al. (2012: 116-117) indicate that the ranges of the two species (C. seabrae and C. lesueuri) might not be allopatric as previously assumed, but might be overlapping. Cistugo lesueuri Roberts, 1919 *1919. Cistugo lesueuri Roberts, Ann. Transv. Mus., 6(3): 112-113. Type locality: South Africa: SW Cape province: Paarl district: near Paarl, Franschhoek [=Frenchhoek] valley: L'Ormarins [=Lormarins] [33 44 S 18 58 E] [Goto Description]. Holotype: TM 2286: ad ♂, skin and skull. Collected by: J.S. Le Sueur Esq. Collection date: 15 September 1917; original number: 154. - Comments: For details on the type locality, see Meester et al., 1986: 49). - Etymology: Named after J.S. le Sueur of L'Ormarins in the Paarl district, Western Cape, who recovered the original specimen from his cat (see Smithers, 1983: 92; Taylor, 2005). (Current Combination) ? Myotis (Cistugo) lesueuri: (Name Combination) ? Myotis lesueuri: (Name Combination) TAXONOMY: Hayman and Hill (1971) doubt the validity of lesueuri and Corbet and Hill (1980) include it in seabrae, while Koopman (1982) retain it as a separate species. It is possible, particularly in view of their allopatric distribution, that lesueuri may best be treated as a subspecies of seabrae. CONSERVATION STATUS: Global Justification Monadjem et al. (2017cg) report that this species is restricted to South Africa and Lesotho in areas with suitable rock crevices and water sources. It has a large range, with an estimated extent of occurrence of over 400,000 km 2 and there are more than 20 known locations. This species is highly likely to be under collected and many more subpopulations are suspected to occur, especially within the Nama and Succulent Karoo regions of South Africa. Wind farms represent an emerging threat, as its preferred habitat coincides with suitable wind farm sites. Although declines have been recorded these are not suspected to be at levels high enough to qualify the species for listing under a threat category. However, systematic long-term monitoring should be used to estimate rates of decline across its range, as this species may require reassessing in a threatened category (Jacobs, 2008b; IUCN, 2009; Monadjem et al., 2017cg). COMMON NAMES: Afrikaans: Lesueur se langhaarvlermuis, Lesueurlanghaarvlermuis, Lesueurse Vlerkkliervlermuis. Czech: žlázokřídlec Lesueurův. English: Lesueur's Wing-gland Bat, Lesueur's Hairy Bat, Lesueur's Myotis, Wing-gland bat. French: Murin de Lesueur. German: Leseuers Mausohr. Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cg). 2008: LC ver 3.1 (2001) (Jacobs, 2008b; IUCN, 2009). 2004: VU D2 ver 3.1 (2001) (Jacobs, 2004b; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Figure 207. A female Cistugo lesueuri (TM 48190) caught at the Farm Schaapplaats, Free State, South Africa. Regional South Africa:- 2016: LC ver 3.1 (2001) (Avenant et al., 2016b). 2004: NT ver 3.1 (2001) (Friedmann African Chiroptera Report 2020 and Daly, 2004). 1986: Indeterminate, assessed as Myotis lesueuri (Smithers, 1986). MAJOR THREATS: The species is locally threatened, in parts of its range, by conversion of land to agricultural use ( (Jacobs, 2008b; IUCN, 2009; Driver et al., 2012). However, as this species occurs mostly in highaltitude areas, this is not a severe threat. The growing trend of developing wind farms in the eastern parts of South Africa and in Lesotho is starting to pose a threat to this species. The degree of impact and levels of decline to the population are currently unknown and should be monitored (Monadjem et al., 2017cg). CONSERVATION ACTIONS: Monadjem et al. (2017cg) report that in the Western Cape, the species is recorded from the three protected areas, Cedarberg Wilderness Area, Gamkaberg Nature Reserve and Karoo National Park; in the Free State the species was recorded in the Golden Gate National Park; in Lesotho it is found in Sehlabathebe National Park, as well as in the Maloti-Drakensberg Transfrontier Conservation Area. No specific interventions are currently necessary, but conservation planning and engagement with the wind energy industry will be needed in future to mitigate subpopulation loss with wind farm construction (Monadjem et al., 2017cg). 583 these results were the same as those obtained for C. seabrae from Namibia (Rautenbach et al., 1993) . Protein / allozyme - Unknown. HABITAT: In the Free State Province of South Africa collected from broken country (e.g. koppies and cliffs), invariably away from human habitation and constructions, but close to open water (dams and streams). While in the Western Cape it inhabits fynbos and possibly the succulent karoo (Jacobs, 2008b; IUCN, 2009). Cooper-Bohannon et al. (2016: Table S2) mention its occurrence in the following habitat types: Afromontane (43 %), High veld (29 %), SW Cape (13 %) and SW arid (11 %). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: GENERAL DISTRIBUTION: This species is restricted to South Africa and Lesotho, where its distribution is best predicted by geology (Babiker Salata, 2012: 50). CooperBohannon et al. (2016: Table S2) calculated a potential distribution area of 673,792 km 2. Native: Lesotho (Lynch and Watson, 1990; Lynch, 1994: 192; Monadjem et al., 2010d: 562); South Africa (Free State - Watson, 1998; Monadjem et al., 2010d: 562; KwaZulu Natal - Monadjem et al., 2010d: 562; Western Cape - Herselman and Norton, 1985; Rautenbach et al., 1993; Seamark and Brand, 2005; Monadjem et al., 2010d: 562). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: For 4 specimens from the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) mention the following parameters: Fa: 36.0 ± 0.4 mm, Wing area: 93.3 ± 1.6 cm 2, Wing span: 23.8 ± 1.1 cm, Wing loading: 7.5 ± 1.4 N/m2, and aspect ratio: 6.1 ± 0.7. MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported 2n = 50, FN = 48, BA = 0, a submetacentric X chromosome, and an acrocentric Y chromosome, Figure 208. Clockwise from top lateral, ventral and dorsal skull images of TM 2286, holotype of Cistugo lesueuri Roberts, 1919. ECHOLOCATION: From Algeria Forest Station (RSA), Schoeman and Jacobs (2002: 157) reported the type of calls to be low duty echolocation dominated by frequency modulated calls, with a Fpeak: 36.8 ± 0.9 kHz. HABITS: In the Free State Province of South Africa specimens were netted in the early evening, between 18:30 and 20:00, with only two specimens being caught after 21:15 (Watson, 1998: 128). When distrubed in their day roost 584 ISSN 1990-6471 they flew to another roost 50 meters away (Watson, 1998: 128). IUCN, 2009). 2004: Decreasing (Jacobs, 2004b; IUCN, 2004). ROOST: Found in a crevice roughly four meters from the ground, on a west-facing dolorite rock face next to a river in the Free State Province of South Africa (Watson, 1998: 127). VIRUSES: Bunyaviridae Phlebovirus Rift Valley Fever Virus (RVFV) - Oelofsen and Van der Ryst (1999) tested a single individual from one locality in Lesotho for RVFV antigen using an enzyme linked immunosorbet assay (ELISA), none tested positive. DIET: From the Algeria Forestry Station (RSA) Schoeman and Jacobs (2002: 157) reported the following prey groups based on 39 faecal pellets from 4 bats (in volume percent): Diptera (47.3 ± 37.3), Hemiptera (32.5 ± 22.8), Hymenoptera (6.8 ± 13.5), Coleoptera (6.3 ± 4.9), Trichoptera (2.5 ± 5.0), Ephemenoptera (1.5 ± 3.0), and unknown (3.3 ± 6.5). POPULATION: Structure and Density:- Although endemic, the species has a wide distribution within the assessment region, despite not being common and very rarely recorded (Monadjem et al., 2017cg. In the Free State Province of South Africa, a group of approximately 40 individuals was located in a day roost (Watson, 1998: 127). Seamark and Brand (2005) showed that this species made up only 4.6% of the over all catch in the Cederberg, Western Cape. In inland Western Cape, near the border with the Northern Cape, a group of approximately 30 individuals was located in a day roost (T. Morgan unpubl. data in Monadjem et al. (2017cg)). Systematic long-term monitoring should be used to estimate rates of decline across its range, as this species may be increasingly threatened by wind farm expansion (Monadjem et al., 2017cg). Trend:- 2016: Decreasing (Monadjem et al., 2017cg). 2008: Decreasing (Jacobs, 2008b; Rhabdoviridae Lyssavirus - Rabies related viruses Rabies - Oelofsen and Smith (1993) tested 15 individual brains, from four localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Lesotho, South Africa. Figure 209. Distribution of Cistugo lesueuri Cistugo seabrae Thomas, 1912 *1912. Cistugo seabræ Thomas, Ann. Mag. nat. Hist., ser. 8, 10 (56): 205. Publication date: 1 August 1912. Type locality: Angola: Mossamedes [=Moçamedes] [ca. 15 56 S 12 10 E] [Goto Description]. Holotype: BMNH 1906.1.3.3: ad ♀, skull and alcoholic. Presented by the Lisbon Museum. (Current Combination) 2004. Myotis (Cistugo) sebrai: Bickham, Patton, Schlitter, Rautenbach and Honeycutt, Mol. Phylogenet. Evol., 33 (2): 333. (Name Combination, Lapsus) ? Cistugo seabrae: (Current Spelling) ? Cistugo seabrai: (Name Combination, Alternate Spelling) ? Myotis (Cistugo) seabrai: (Name Combination) ? Myotis seabrae: ? Myotis seabrai: (Name Combination) TAXONOMY: See Simmons (2005). African Chiroptera Report 2020 COMMON NAMES: Afrikaans: Angola-langhaarvlermuis, Angolase Vlerkkliervlermuis. Czech: žlázokřídlec Seabrův. English: Seabra's Wing-gland Bat, Angolan Winggland Bat, Angolan Hairy Bat. French: Murin d'Angola. German: Angola-Mausohr. ETYMOLOGY OF COMMON NAME: The colloquial name is derived from the fact that the original specimen originated from Moçâmedes on the south-western coast of Angola (Taylor, 2005). CONSERVATION STATUS: Global Justification Although this species is known mainly from isolated records from a large area, it is listed as Least Concern (LC ver 3.1 (2001)) in view of its Figure 210. A male Cistugo seabrae (TM 47582) caught at Khamkirri, Northern Cape, South Africa. Note the gland on the wing. wide distribution, presumed large population, and because there are no threats, making it unlikely to be declining fast enough to qualify for listing in a more threatened category (Griffin and Jacobs, 2008; IUCN, 2009; Monadjem et al., 2017bj). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bj). 2008: LC ver 3.1 (2001) (Griffin and Jacobs, 2008; IUCN, 2009). 2004: NT ver 3.1 (2001), assessed as Cistugo seabrai (Jacobs, 2004a; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional South Africa:- 2016: NT D1+D2 ver 3.1 (2001) (Jacobs et al., 2016e). 2004: VU D2 ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate, assessed as Myotis seabrai (Smithers, 1986). MAJOR THREATS: There are no major threats to this species. It may be locally threatened by mining operations in a small part of its range (Griffin and Jacobs, 2008; IUCN, 2009; Monadjem et al., 2017bj). 585 CONSERVATION ACTIONS: Griffin and Jacobs (2008) [in IUCN (2009)] report that it is not known if there are any direct conservation measures in place for this species, and it is unclear if it occurs within any protected areas. Further studies are needed into the distribution and natural history of this little-known species. GENERAL DISTRIBUTION: This southern African species ranges from the type locality of Moçâmedes in southwestern Angola, southwards through western Namibia to the Northern Cape of South Africa. Native: Angola (Crawford-Cabral, 1989; Thomas, 1912c; Monadjem et al., 2010d: 562); Namibia (Thomas and Hinton, 1925; Shortridge, 1934; Rautenbach et al., 1993; Kearney and Van Schalkwyk, 2009; Monadjem et al., 2010d: 562); South Africa (Herselman and Norton, 1985; Stadelmann et al., 2004b; Seamark and Kearney, 2006; Kearney and Van Schalkwyk, 2009; Monadjem et al., 2010d: 562). MOLECULAR BIOLOGY: DNA - Stadelmann et al. (2004b). Karyotype - Rautenbach et al. (1993) reported 2n = 50, FN = 48, BA = 0, a submetacentric X chromosome, and an acrocentric Y chromosome, these results were the same as those obtained for C. lesueuri from South Africa (Rautenbach et al., 1993). Protein / allozyme - Unknown. HABITAT: Little is known about the natural history of this species. All of the localities from which they have been collected are arid with a mean annual rainfall of less than 100 mm (Skinner and Chimimba, 2005). HABITS: Specimens have usually been caught close to open water and have been observed gleaning insects from orange trees (Roberts, 1951; Taylor, 2000; Skinner and Chimimba, 2005). POPULATION: Structure and Density:- It appears to be a rarely recorded species (Griffin and Jacobs, 2008; IUCN, 2009; Monadjem et al., 2017bj). Trend:- 2016: Unknown (Monadjem et al., 2017bj). 2008: Unknown (Griffin and Jacobs, 2008; IUCN, 2009). 2004: Decreasing (as Cistugo seabrai) (Jacobs, 2004a; IUCN, 2004). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Namibia, South Africa. 586 ISSN 1990-6471 Figure 211. Distribution of Cistugo seabrae †Family INDETERMINATE †Genus Chambinycteris Ravel, 2016 *2016. Chambinycteris Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 402. Publication date: 30 September 2016. - Etymology: Refers to Chambi, the locality where the specimens were collected (see Ravel et al., 2016: 402). TAXONOMY: Ravel et al. (2016: 358) indicate that this genus has a basal position within the Emballonuridae. †Chambinycteris pusilli Ravel, 2016 *2016. Chambinycteris pusilli Ravel, in: Ravel et al., Geodiversitas, 38 (3): 403. Publication date: 30 September 2016 [Goto Description]. - Etymology: From the Latin "pusilli" meaning puny or sickly, referring to the small size and fragility of the species (see Ravel et al., 2016: 403). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. †Genus Drakonycteris Ravel, 2016 *2016. Drakonycteris Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 399. Publication date: 30 September 2016. - Etymology: From the Greek "δράκων" meaning dragon and "νυκτερισ" meaning bat (see Ravel et al., 2016: 399). †Drakonycteris glibzegdouensis Ravel, 2016 *2016. Drakonycteris glibzegdouensis Ravel, in: Ravel et al., Geodiversitas, 38 (3): 356, 399, figs 23, 24. Publication date: 30 September 2016. Type locality: Algeria: Béchar province: Gour Lazib: African Chiroptera Report 2020 587 Sahara, Hammada du Dra: Glib Zegdou [29 42 49 N 05 45 58 W] [Goto Description]. - Etymology: Referring to the type locality of Glib Zegbou (see Ravel et al., 2016: 399). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Late lower Eocene (Ypresian - Brown et al., 2019: Suppl.) to early middle Eocene. Family MINIOPTERIDAE Dobson, 1875 1875. Miniopteri Dobson, Ann. Mag. nat. Hist., ser. 4, 16 (95): 349. Publication date: 1 November 1875. *1875. Miniopteridae Dobson, Ann. Mag. nat. Hist., ser. 4, 16 (95): 348. Publication date: 1 November 1875. - Comments: Type genus: Miniopterus Bonaparte, 1837. Originally included the genera Natalus J. Gray, 1838; Thyroptera Spix, 1823 and Miniopterus Bonaparte, 1837. (Current Combination) 1898. Miniopterae Trouessart, Cat. Mamm, p. 134. - Comments: See Taylor (1934). 1907. Miniopterinae: Miller, Bull. U.S. natl. Mus., 57: xi, 227. Publication date: 29 June 1907. Comments: Type genus: Miniopterus Bonapart, 1837. 1977. Miniopteridae: Mein and Tupinier, Mammalia, 41 (2): 207, 209. - Comments: Type genus: Miniopterus Bonaparte, 1837. TAXONOMY: Bannikova et al. (2002: 722) indicate that the Miniopterinae do not deserve the rank of independent family (as did Koopman, 1984c, 1993a, 1994; Yoshiyuki, 1989; Corbet and Hill (1992; McKenna and Bell, 1997), but see Hoofer and Van Den Bussche (2003: 13) who make a strong case for attributing family rank. They follow Mein and Tupinier (1977), Groombridge (1994), Gopalakrishna and Chari (1983), Tiunov (1989), Pierson (1986). See also Hawkins et al. (2019: 11355). Currently (Simmons and Cirranello, 2020) recognized genera of the family MINIOPTERIDAE: Miniopterus Bonaparte, 1837. In their study on rolling-circle transposons (Helitrons), Thomas et al. (2010: 55) found none of these in the Miniopteridae. They also indicated that these Helitrons colonized the Vespertilionidae some 30 - 36 mya, which is much later than the split between Vespertilionidae and Miniopteridae, which occurred some 43 million years ago. Patterson and Webala (2012: 32) indicate that the split between Vespertilionidae and Miniopterus occurred between 38 and 47 million years ago. Demos et al. (2019c: "6") indicate that the Miniopteridae diverged from the Vespertilionidae and Cistugidae some 48 mya. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Amador et al. (2016: 26) estimated the split from the Vespertilionidae at c. 48 MYA, and a stem age at 13 MYA. Herkt et al. (2017: Appendix 4S), investigating the IUCN distribution maps and species distribution models, left out the Miniopteridae because of their exceptionally high cryptic diversity and level of phenetic similarity, which leads to unreliably identification of large amounts of published records. COMMON NAMES: Czech: létavcovití, netopýři dlouhorucí. Dutch: Langvleugelvleermuizen. English: Long-fingered Bats. French: Minioptéridés. German: Langflügelfledermäuse. Italian: Miniottèridi, Minioptèridi. Russian: Длиннокрыловые. Ukrainian: Довгокрили [= Dovhokryly]. Vietnamese: Họ dơi cánh. MOLECULAR BIOLOGY: Capanna et al. (1968: 240) already pointed out that there was a cytotaxonomic difference between the Vespertilioninae and the Miniopterinae, especially in the average length of the aploid female set. Platt et al. (2016: 9) analyzed the genome data for M. natalensis and compared it with that of several other vesper bats and they concluded that Helitrons and hATs were introduced into the genome of the Vespertilionidae after their divergence from Miniopterus. Sotero-Caio et al. (2017: 5) indicate that the karyotype for all 25 species in the family has a 2n value of 46. 588 ISSN 1990-6471 Genus Miniopterus Bonaparte, 1837 *1837. Miniopterus Bonaparte, Iconografia Fauna italiana, 1, fasc. 20 [Goto Description]. Comments: Type species: Vespertilio ursinii Bonapate, 1837 (=Vespertilio schreibersii Kuhl, 1817). - Etymology: From the Greek "μινύς" or "μινός", meaning small and "πτέρυξ", meaning wing, referring to the very short first phalanx of the third and longest finger (see Palmer, 1904: 426; Flannery, 1995a: 374; Kozhurina, 2002: 16). Lanza et al. (2015: 328) mention the following: "According to Bonaparte (1837-41) who described the genus and Lanza (2012), Miniopterus is a scientific masculine Latin substantive made up of the proper feminine Latin noun Minyèias 'Minyad', i. e. a daughter of Minyas, king of Orchomenus, turned into a bat, and of the Greek neuter substantive πτερόν" (pteròn) 'wing', meaning 'flying Minyad'. The etymology proposed by Kozhurina (2002), meaning 'with small (narrow) wings', is incorrect, even if formally logical. Also partially erroneous is the etymology given by some Italian dictionaries, such as Battaglia (1978) and Devoto & Oli (1997), who interpret 'minio' as 'minimum, red-lead', the red colouring agent, which obviously has nothing to do with the greyish brown colour of Miniopterus' wings". (Current Combination) 1846. Minyopterus: Agassiz, Nomenclator. Zool., Mamm., 235. - Comments: Variant spelling (Meester et al., 1986: 45) or unjustified emendation (Jackson and Groves, 2015: 263). (Lapsus, Emendation) 1858. Miniopteris: Tomes, Proc. zool. Soc. Lond., 1858, XXVI (cccliv): 115. Publication date: 13 May 1858. - Comments: Variant spelling (Meester et al., 1986: 45) or unjustified emendation (Jackson and Groves, 2015: 262). Not Miniopteris J. Gray, 1849 (Mammalia, Chiroptera, Vespertilionidae) (used as genus name for Myotis macrotarsus (Waterhouse, 1845) (see Jackson and Groves, 2015: 263). (Alternate Spelling) 1900. Minneopterus: Lampe, Jb. nass. Ver. Naturkde, 53: Cat. Säugeth.-Sammlung, p. 12. Publication date: 1900. - Comments: Cat. Säugeth.-Sammlung, p. 12. Lapsus (see Allen, 1939a: 102; Meester et al., 1986: 45). Jackson and Groves (2015: 263) indicate it appeart to be a nomen nudum or an unjustified emendation. (Lapsus, Emendation) 1920. Mineopterus: Haagner, South African Mammals, 22. (Lapsus) 1980. Minioterus: Bernard, Ann. Transv. Mus., 32 (3): 55. Publication date: January 1980. (Lapsus) 2014. Mineropterus: Kohl and Kurth, Viruses, 6 (8): 3112. Publication date: 13 August 2014. (Lapsus) 2015. Minioptera: Golden and Comaroff, Ecol. Soc., 20 (2): 42: 5. (Lapsus) 2019. Minioptreus: Zeghbib, Herczeg, Kemenesi, Zana, Kurucz, Urbán, Madai, Földes, Papp, Somogyi and Jakab, Sci. Reps., 9 (15706): 1. Publication date: 31 October 2019. (Lapsus) ? Miniopterus sp. TAXONOMY: Reviewed (in part) by Peterson (1981). See Goodwin (1979: 119), Maeda (1982), and Hill (1983). All have come to very different conclusions. Molecular systematics and biogeography is discussed by Appleton et al. (2004); their DNA data (p. 437) suggest that the number of species currently recognized (11 or 13) is a gross underestimate. Phylogentic analyses performed by Christidis et al. (2014: 1) indicate that 18 clades are present within Malagasy Miniopterus of which only 11 have Latin binominals. Šrámek et al. (2012: 165) investigated the taxonomic status of the bent-winged bats in the western Palaearctic and distinguished four species: M. schreibersii (s.str.) from Europe, coastal Anatolia, Levant, Cyprus, western Transcaucasia, and North Africa; M. pallidus from inland Anatolia, Jordan, eastern Transcaucasia, Turkmenistan, Iran and southern Afghanistan (Kandahar); a Miniopterus sp., recorded from Nangarhar province in eastern Afghanistan, which was tentatively assigned to M. cf. fuliginosus; and another Miniopterus sp. With Afro-tropic affinities confirmed from south-western Arabia and Ethiopia, which tentatively named M. cf. arenarius. They also indicate that a possible new taxon (subspecies) within M. schreibersii exists in the Atlas Mountains of Morocco. Bilgin et al. (2013: 21) indicate that this Maghrebian form possibly even represents a new species, which they tentatively call: M. maghrebensis. See Taylor et al. (2019a: 318) for a discussion on the recent increase in Afro-Madagascan species. Demos et al. (2019c: "1") analysed cyt-b and African Chiroptera Report 2020 nuclear data from sub-Saharan Miniopterus specimens and found evidence for five undescribed species. Their data (p. "9") confirmed the species status of fraterculus, mossambicus, natalensis and newtoni and tentatively for africanus, arenarius, inflatus and minor. M. inflatus, minor and natalensis are probably paraphyletic. The 18 Malagasy clades recovery by Christidis et al. (2014) in their phylogenetic analysis were divided in into five primary lineages (representing sister species): (1) M. griveaudi; (2) M. mahafaliensis, M. sororculus and X3; (3) M. majori, M. gleni and M. griffithsi; (4) M. brachytragos; M. aelleni A, and M. aelleni B, and (5) M. manavi and M. petersoni. These are in turn linked to a group comprising M. egeri and five genetically distinct populations referred to as P3, P4, P5, P6 and P7. Currently (Simmons and Cirranello, 2020) recognized species of the genus Miniopterus:aelleni Goodman, Maminirina, Weyeneth, Bradman, Christidis, Ruedi and Appleton, 2009; africanus Sanborn, 1936; ambohitrensis Goodman, Ramasindrazana, Naughton and Appleton, 2015; australis Tomes, 1858 – Philippines, Borneo, Java, Timor, Moluccas, southeast to Vanuatu and eastern Australia (Simmons, 2005: 519); brachytragos Goodman, Maminirina, Bradman, Christidis and Appleton, 2011; egeri Goodman, Ramasindrazana, Maminirina, Schoeman and Appleton, 2011; egeri Goodman, Ramasindrazana, Maminirina, Schoeman and Appleton, 2011; fraterculus Thomas and Schwann, 1906;fuliginosus (Hodgson, 1835) – Afghanistan to Myanmar; Japan; Vietnam; Korea; fuscus Bonhote, 1902 – Ryuku Isls (Japan) (Simmons, 2005: 519); gleni Peterson, Eger and Mitchell, 1995; griffithsii Goodman, Maminirina, Bradman, Christidis and Appleton, 2009; griveaudi Harrison, 1959; inflatus Thomas, 1903; macrocneme Revilliod, 1914 – New Guinea to Vanuatu and New Caledonia (Simmons, 2005: 520); maghrebensis Puechmaille, Allegrini, Benda, Bilgin, Ibáñez and Juste, 2014; magnater Sanborn, 1931 – northeastern India, southeastern China, Burma, Thailand, Laos and Vietnam to Malaysia, Sumatra, Java, Timor (Indonesia), Borneo, Moluccas and New Guinea including the Bismarck Arch. (Simmons, 2005: 520); mahafaliensis Goodman, Bradman, Christides and Appleton, 2009; majori Thomas, 1906; manavi Thomas, 1906; medius Thomas and Wroughton,1909 – southeastern China, Thailand, western Malaysia, Borneo, Java, Sulawesi, Philippines, New Guinea (Simmons, 2005: 520); minor Peters, 1867; mossambicus Monadjem, Goodman, Stanley and Appleton, 2013; 589 natalensis (A. Smith, 1834); orianae Thomas, 1922 – Australia; pallidus Thomas, 1907 – N Arabia; Asian part of Turkey; Afghanistan; paululus Hollister, 1913 – Majuyod, Negros and Guimarás Isl (Philippines), Berneo, Selaru (Simmons, 2005: 521); petersoni Goodman, Bradman, Maminirina, Ryan, Christidis and Appleton, 2008; pusillus Dobson, 1876 – India, Nepal and Burma to Sumatra and Timor (Indonesia), Philippines and Moluccas (Simmons, 2005: 520); robustior Revilliod, 1914 – Loyalty Isls (east of New Caledonia) (Simmons, 2005: 521); schreibersii (Kuhl, 1817); shortridgei Laurie and Hill, 1957 – Java, Madura, Lombok, Sumbawa, Moyo, Alor, Wetar, Seralu, Timor, Semau, Roti and Savu Isls (Indonesia) (Simmons, 2005: 522); sororculus Goodman, Ryan, Maminirina, Fhar, Christidis and Appleton, 2007; tristis (Waterhouse, 1845) – Philippines, Sulawesi, Sanan Isl, New Guinea, Bismarck Arch., Solomon Isls, New Hebrides (Simmons, 2005: 522). Furthermore Monadjem et al. (2020: 242, 255) recognized arenarius Heller, 1912 and nimbae Monadjem, Shapiro, Richards, Karabulut, Crawley, Broman Nielsen, Hansen, Bohmann and Mourier, 2020 and villiersi Aellen, 1956 as additional, separate species. And there is also the extinct †horaceki Gunnell, Eiting, and Geraads, 1837. COMMON NAMES: Czech: létavci, netopýři dlouhorucí. English: Long-fingered Bats, Bent-winged Bats. French: Minioptères. German: Langflügelfledermäuse, Sackfledermaus. ETYMOLOGY OF COMMON NAME: The very long second phalanx of the third finger led to the name "long-fingered bats". The feature also allows these bats to fold or bent their wings back, hence "bent-winged bats" (see Demos et al., 2019c: "2"). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: The oldest fossil remains for Miniopterus date from some 13.65 million years ago (see Shi and Rabosky, 2015: 1532). Butler (1978) refers to African material of this genus, found from the early Pleistocene at Olduvai, and Avery (2007: 619) reported on the presence of Miniopterus sp. in Pleistocene deposits at Wonderwerk Cave, South Africa. Geraads et al. (2010: 279) reported Miniopterus in Late Cenozoic (ca. 2.5 MYA) deposits in Ahl al Oughlam, Morocco. Weyeneth (2010) suggested that Malagasy Miniopterus colonized the Comoros during the Pleistocene, favored by prevailing winds. 590 ISSN 1990-6471 BIOGEOGRAPHY: Weyeneth et al. (2008) suggest that the Comoros were colonized independently at least two or three times by ancestors from Madagascar. DETAILED MORPHOLOGY: Brain - Maseko and Manger (2007) describe the nuclear parcellation and neuronal morphology of the cholinergic, catecholaminergic and serotonergic systems within the brain of six adult female Miniopterus schreibersii [possibly M. natalensis or M. fraterculus, not Miniopterus schreibersii], captured from caves in northwest region of Gauteng Province, South Africa. SEXUAL DIMORPHISM: Goodman and Maminirina (2007) found no notable evidence for sexual dimorphism in Malagasy Miniopterus spp. ECHOLOCATION: Bioacoustic data of 11 Malagasy (Madagascar and the Comoros) has been collected and investigated by Ramasindrazana et al. (2011). PARASITES: BACTERIA Bartonellae Bartonella - Kosoy et al. (2010: 1877) reported a prelevence of Bartonella spp. in Miniopterus sp. from Kenya of 22/105 (21.0 %) cultured from blood samples. Fifty-one Bartonella spp. gltA sequences were obtained from 3 identified species of Miniopterus (africanus, minor, natalensis), were 24 unique gltA genotypes representing at least 3 genogroups (Kosoy et al., 2010: 1878). See also Kosoy (2010: 719) for further information. ACARI Myobiidae: Fain (1994: 1280) reported the genus Calcarmyobia to occur only on bats of the genus Miniopterus with 18 species and 8 subspecies. Trombiculidae: Trombigastia (Trombigastia) cadei (Vercammen-Grandjean and Brennan, 1957) was reported by Vercammen-Grandjean and Fain (1958: 26) and Stekolnikov (2018a: 118) on Miniopterus sp. specimens from N'Gong, S Kenya. Uchikawa (1985c: 110) reported the presence of Pteracarus miniopteri Uchikawa, 1978 from Miniopterus sp. specimens from Madagascar, Central Africa, Cameroon, Kenya (Mombasa), South Africa (Transvaal). DIPTERA Streblidae: Raymondiodes leleupi Jobling, 1954 described form a poorly identified bat (Hipposideros caffer or Miniopterus sp.) from Thysville, Congo (Haeselbarth et al., 1966: 104). Nycteribosca kollari Frauenfeld, 1855 reported by Vermeil (1960) from Tunisia. was Nycteribiidae: Nycteribia tecta Theodor, 1957 (Haeselbarth et al., 1966: 110). Duron et al. (2014: 2108) mention the presence of Nycteribia stylidiopsis (Speiser, 1908) on Miniopterus sp. from the Comoros, and of Penicillidia leptothrinax (Speiser, 1908) from Madagascar. The latter were carrying Enterobacteriales and Bartonella and Wolbachia bacteria (Szentiványi et al., 2019: Suppl.). Enterobacteriales were also found in Penicillidia cf. fulvida (Szentiványi et al., 2019: Suppl.). SIPHONAPTERA Ischnopsyllidae: Members of the genus Rhinolophopsylla Oudemans, 1909 are known to be parasites of Miniopterus and Rhinolophus (Haeselbarth et al., 1966: 191). VIRUSES: Adenoviridae These viruses were found by Waruhiu et al. (2017) in bats from Kenay. Coronaviridae 18 out of 300 Kenyan Miniopterus bats tested by Tao et al. (2017: Suppl.) was positive for CoV (6 %). Paramyxoviridae Mortlock et al. (2015: 1841) reported that 13 out of 77 examined Kenyan Miniopterus sp. specimens tested positive for Paramyxovirus sequences. This was also the case for two out of 41 specimens from the DRC. Retroviridae Farkašová et al. (2017: 3145) described an endogenous Deltaretrovirus, which only occurs in this family of bats and not in the closely related Vespertilionidae and Cistugonidae. This suggests that the endogenization occurred between 20 and 45 million years ago. Rhabdoviridae Lyssavirus - Rabies related viruses West Caucasian Bat virus (WCBV) - Kuzmin et al. (2008b) detected WCBV in four of five localities sampled in Kenya with a seroprevalence of 1726%. Horton et al. (2014: Table S1) tested 229 Kenyan Miniopterus sp. specimens, but failed to find neutralising antibodies to IKOV (Ikoma lyssavirus). Duvenhage lyssavirus (DUVV) - Banyard A. C. and Fooks A. R. (2017: 18) reported this virus from Miniopterus sp. specimens from South Africa and Kenya. African Chiroptera Report 2020 DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Angola, Comoros, Congo (Democratic Republic of the), Ethiopia, Gabon, 591 Kenya, Madagascar, Malawi, Mozambique, South Africa, South Sudan, Tanzania, Tunisia, Uganda, Zambia, Zimbabwe. †Miniopterus horaceki Gunnell, Eiting and Geraads, 2011 *2011. Miniopterus horaceki Gunnell, Eiting and Geraads, N. Jb. Geol. Paläont. Abh., 260 (1): 63, figs. 3B, 7C, 8D-F. Publication date: January 2011. Type locality: Morocco: Ahl al Oughlam [ca. 33 35 N 07 30 W] [Goto Description]. Holotype: INSAP AaO 4605: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Right dentary with p2-p4, m1 trigonid. - Etymology: In honour of Ivan Horácek, in recognition of his many contributions to the understanding of the evolutionary history of bats, and especially of vespertilionids (see Gunnell et al., 2011: 63). (Current Combination) TAXONOMY: Gunnell et al. (2011: 63) describe it as larger than nearly all extant African and European Miniopterus species (including M. gleni, M. inflatus, M. newtoni, M. fraterculus, M. natalensis, M. manavi, M. noreenae, M. minor, M. williamsi, and M. schreibersi). Generally similar in size to M. africanus but differs in having p34 relatively broader and more exodaenodont, p3 larger than p2, and in having relatively taller lower premolars. Larger than M. schreibersi with p2 smaller compared to p3, but M. schreibersi does possess relatively broad premolars similar to the Moroccan species. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pliocene (Brown et al., 2019: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Libya. Miniopterus aelleni Goodman, Maminirina, Weyeneth, Bradman, Christidis, Ruedi and Appleton, 2009 *2009. Miniopterus aelleni Goodman, Maminirina, Weyeneth, Bradman, Christidis, Ruedi and Appleton, Zool. Scr., 38 (4): 339, 353, figs. 3d, 3e, 4, 7 - 10. Publication date: 22 June 2009. Type locality: Madagascar: Antsiranana province: Ankarana special reserve, Canyon d'Antsiroandoa Centre,: 6.2 km NW Mahamasina [12 55.1 S 49 07.6 E, 120 m] [Goto Description]. Holotype: FMNH 173067: ad ♂, skull and alcoholic. Collected by: Vola Razakarivony and Scott G. Cardiff; collection date: 8 April 2002; original number: SMG 12809. See Goodman and Cardiff (2004: 230). Paratype: FMNH 173069: ad ♀, skull and alcoholic. Collected by: Vola Razakarivony and Scott G. Cardiff; collection date: 8 April 2002; original number: SMG. See Goodman and Cardiff (2004: 230). Paratype: FMNH 173070: ad ♀, skull and alcoholic. Collected by: Vola Razakarivony and Scott G. Cardiff; collection date: 8 April 2002; original number: SMG. See Goodman and Cardiff (2004: 230). Paratype: FMNH 173072: ad ♂, skull and alcoholic. Collected by: Vola Razakarivony and Scott G. Cardiff; collection date: 8 April 2002; original number: SMG. See Goodman and Cardiff (2004: 230). Paratype: FMNH 173075: ad ♂, skull and alcoholic. Collected by: Vola Razakarivony and Scott G. Cardiff; collection date: 8 April 2002; original number: SMG. See Goodman and Cardiff (2004: 230). Paratype: FMNH 173076: ad ♂, skull and alcoholic. Collected by: Vola Razakarivony and Scott G. Cardiff; collection date: 8 April 2002; original number: SMG. See Goodman and Cardiff (2004: 230). - Etymology: The name aelleni is a patronym for Villy Aellen (1926 - 2000) who was a mammalogist at the Muséum d’histoire naturelle de Genève from 1954 to his retirement in 1989, where he also served as curator, assistant director, and director (Mahnert, 2000; Goodman et al., 2009a: 353). (Current Combination) 2014. Miniopterus aelleni A: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. TAXONOMY: See Goodman et al. (2009a). Schoeman et al. (2014: 19) refer to Christidis et al. (2014) indicating that - from a phylogenetic species perspective - there is evidence that two sister species are present, which are tentatively named: M. aelleni A and M. aelleni B. 592 ISSN 1990-6471 COMMON NAMES: Czech: létavec Aellenův. English: Aellen's Longfingered Bat. German: Aellens Langflügelfledermaus. ETYMOLOGY OF COMMON NAME: In honour of Villy Aellen (1926 - 2000), who was a mammalogist at the Muséum d’histoire naturelle de Genève from 1954 to his retirement in 1989, where he also served as curator, assistant director, and director. He devoted much of his professional interests to the study of bats, particularly those of Africa (see Goodman et al., 2009a: 353). CONSERVATION STATUS: Global Justification This species occurs at several sites in the western half of Madagascar (Goodman and Ramasindrazana, 2013); it is also known from a few specimens obtained on Anjouan Island in the Comoros Archipelago (Goodman et al., 2010c). It is apparently not forest-dependant and may not be subjected to any important habitat degradation or other human pressures (Goodman et al., 2010c). Assessment History Global 2016: LC ver. 3.1 (2001) (Goodman, 2017g). Regional None known. CONSERVATION ACTIONS: Goodman (2017g) report that on Madagascar, this species is known from several protected areas, which include Ankarana, Namoroka and Bemaraha. While to their knowledge the sites this species occurs on Anjouan Island are not under any special protection. GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Goodman et al. (2009a). ECHOLOCATION: Kofoky et al. (2009: 382) reported the calls as M. manavi sen lat., these calls were broadband FM/QCF sweeps produced at low duty cycle with most energy at about 58.1 kHz in the fundamental, with pulses having a short duration of about 4 ms. Ramasindrazana et al. (2011: 294) recorded 19 calls from 7 Miniopterus aelleni specimens, and found the following echolocation call values: Frequency of maximum energy: 51.9 ± 1.57 (49.5 - 54.8) kHz, Maximum frequency: 99.8 ± 8.48 (75.0 - 119.0) kHz, Minimum frequency: 48.0 ± 1.43 (46.0 - 50.0) kHz, Call duration: 4.1 ± 0.65 (3.0 5.0) ms, Inter pulse interval: 87.3 ± 16.75 (61.0 118.5) ms. MOLECULAR BIOLOGY: DNA - See Goodman et al. (2009a). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: Dammhahn and Goodman (2013: 108) mention the lower portion of forest structure and partially open areas as the foraging habitat for this species. Also see Goodman et al., 2009a). ROOST: See Goodman et al. (2009a). POPULATION: Structure and Density:- See Goodman et al., 2009a). While Goodman (2017g) report that it is not known. Trend:- 2016: (Goodman, 2017g). GENERAL DISTRIBUTION: Lowland portions of northern and western Madagascar (Goodman et al., 2009a; Schoeman et al., 2014: 28). Schoeman et al. (2014: 31) assigned M. aelleni A to the western or dry deciduous region bioclimatic zone, whereas M. aelleni B was classified to the humid forest region. O'Brien (2011: 289) reports it also from the Comoros. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Goodman et al. (2009a). DENTAL FORMULA: 2123/ 3133 = 36. PARASITES: Tortosa et al. (2013: 4) report the presence of the Nycteribiid bat fly Nycteribia stylidiopsis Speiser, 1908 and Wilkinson et al. (2016) report Penicillidia leptothrinax Speiser, 1908, which in turn carried Bartonella bacteria and the blood parasite Polychromophilus melanipherus (Szentiványi et al., 2019: Suppl.). Ramasindrazana et al. (2017: Suppl.) report both flies. Ramasindrazana et al. (2016: 6) reported the presence of Litosoma Clade 3 (Nematoda: Onchocercidae) on this bat species. They also mention Litosoma goodmani Martin, Bain, Jouvenet, Raharimanga, Robert & Rousset, 2006 from a M. aelleni specimen, that was previously identified as "M. manavi". African Chiroptera Report 2020 593 DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, Madagascar. Figure 212. Distribution of Miniopterus aelleni Miniopterus africanus Sanborn, 1936 *1936. Miniopterus africanus Sanborn, Field Mus. Nat. Hist., Zool. Ser., (362) 20 (14): 111. Publication date: 15 August 1936. Type locality: Ethiopia: Shoa province: Mulo, Sanford's Ranch [ca. 09 00 N 39 00 E, 8000 ft] [Goto Description]. Holotype: FMNH 28769: ad ♀, skin and skull. Collected by: Alfred Marshall Bailey; collection date: 25 October 1926; original number: 73. (Current Combination) ? Miniopterus inflatus africanus: (Name Combination) TAXONOMY: Included in inflatus by Koopman (1993a: 230), but considered a valid species by Peterson et al. (1995: 120), Lavrenchenko et al. (2004b: 144), Simmons (2005). COMMON NAMES: Chinese: 非 洲 长 翼 蝠 . Czech: létavec východoafrický. English: African Long-fingered Bat. French: Minioptère d'Afrique orientale. German: Afrikanische Langflügelfledermaus. CONSERVATION STATUS: Global Justification Assessment History Global LC ver 3.1 (2001) (IUCN, 2004; Schlitter, 2004h). GENERAL DISTRIBUTION: Native: Botswana; Eritrea; Namibia; Tanzania. Ethiopia; Kenya; PARASITES: Lutz et al. (2016: 9) examined 20 M. africanus specimens from East Africa and found four of them infected with the haemosporidan parasite Polychromophilus melanipherus Dionisi, 1899. Uchikawa (1985a: 19) reported that the type specimen of Miniopterus africanus harboured the "atypical" form of Calcarmyobia congoensis Uchikawa, 1982 (Acari: Myobiidae). This species is also a host of Calcarmyobia kenyaensis Uchikawa, 1982 in Kenya (see Uchikawa (1985a: 22) as M. inflatus africanus). Possibly some other records mentioned by Uchikawa (1985a: 22) as collected from M. inflatus from Cameroon, Liberia, and The Democratic Republic of Congo should be included here. Uchikawa (1985c: 110) also reported on the presence of the mite Pteracarus miniopteri Uchikawa, 1978 on some specimens from Kenya, that were identified as M. inflatus africanus. Tortosa et al. (2013: 5) and Ramasindrazana et al. (2017: Suppl.) mention this species as host for the Nycteribiid bat flies Nycteribia schmidlii Schiner, 1853 and Pencillidia fulvida Bigot, 1885. DIPTERA: Streblidae: Raymondia intermedia Jobling, 1936 (Shapiro et al., 2016: 255). Raymondia seminuda Jobling, 1954 (Shapiro et al., 2016: 255). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Tong et al. (2009) tested 1 bat collected in Kenya during 2006 positive for the 594 ISSN 1990-6471 presence of coronavirus RNA in a fecal sample. One Kenyan bat tested by Tao et al. (2017: Suppl.) was negative for CoV. Polyomaviridae: Polyomavirus - Of the nine specimens tested from the Chyulu National Park (Kenya), one tested positive (Conrardy et al., 2014: 259). Rhabdoviridae: Rhabdovirus - One of the nine specimens tested from the Chyulu National Park (Kenya) tested positive (Conrardy et al., 2014: 259). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Eritrea, Ethiopia, Kenya, Somalia. Figure 213. Distribution of Miniopterus africanus Miniopterus ambohitrensis Goodman, Ramasindrazana, Naughton and Appleton, 2015 2014. Miniopterus aelleni B: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. *2015. Miniopterus ambohitrensis Goodman, Ramasindrazana, Naughton and Appleton, Zootaxa, 3936 (4): 538, 543, figs 2 - 5. Publication date: 23 March 2015. Type locality: Madagascar: Province d'Antsiranana: Parc National de la Montagne d'Ambre: Joffreville (Ambohitra), 5.5 km SW: Station Forestière des Roussettes [12 31 37.3 S 49 10 19.1 E, 1000 m] [Goto Description]. Holotype: FMNH 202450: ad ♂, skull and alcoholic. Collected by: Steven M. Goodman; collection date: 9 May 2009; original number: SMG 16169. Presented/Donated by: ?: Collector Unknown. - Etymology: The name ambohitrensis is derived from the geographical name of the type locality, which in Malagasy is Ambohitra and in French is Montagne d'Ambre. The nearby village of Joffreville is also referred to as Ambohitra. In the Malagasy language, the root word of Ambohitra is vohitra meaning mountain or highlands, giving an ecological context to the specific epithet of this bat, which occurs at higher elevations. COMMON NAMES: English: Montagne d'Ambre Long-fingered Bat. French: Minioptère de Montagne d'Ambre. German: Ambre-Langflügelfledermaus. CONSERVATION STATUS: Global Justification This species is known from several sites in the northern portion of Madagascar (Goodman et al., 2015b), most of which are montane and apparently not subjected to any important habitat degradation or other human pressures. It is notably common at several of these sites (Goodman, 2017h). Assessment History Global LC 3.1 (2001) (Goodman, 2017h). Regional Endemic to Madagascar, therefore assessment should be consulted. global CONSERVATION ACTIONS: Goodman (2017h) reports that this species is known from several protected areas, which include Marojejy, Montagne d’Ambre, Ambohitantely, and Bemanevika. POPULATION: Structure and Density:- Not known (Goodman, 2017h). Trends: 2016: Unknown (Goodman, 2017h). PARASITES: Wilkinson et al. (2016) report the presence of Penicillidia leptothrinax on "Miniopterus cf. ambohitrensis", which was carrying Enterobacteriales bacteria (Szentiványi et al., 2019: Suppl.). African Chiroptera Report 2020 595 VIRUSES: Paramyxoviridae Mélade et al. (2016b: 4) found one out of 19 M. cf. ambohitrensis specimens they tested to be infected with paramyxoviruses. Rhabdoviridae Four out of 15 M. cf. ambohitrensis specimens tested by Mélade et al. (2016a: 6) showed a reaction against Duvenhage Lyssavirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Figure 214. Distribution of Miniopterus ambohitrensis Miniopterus arenarius Heller, 1912 *1912. Miniopterus natalensis arenarius Heller, Smiths. Misc. Coll., 60 (12): 2. Publication date: 4 November 1912. Type locality: Kenya: Central province: Nanyuki district: Northern Guaso Nyiro: Nyuki (=Nanyuki) river [00 01 N 37 04 E] [Goto Description]. Holotype: USNM 181811: ad ♀. Collected by: Edmund Heller; collection date: 4 October 1911; original number: 4413. - Comments: Coordinates mentioned as 00 21 N 36 55 E by Thorn et al. (2009: 49). 2010. Miniopterus arenarius: Benda, Vespertilio, 13-14: 307. (Name Combination) ? Miniopterus schreibersi arenarius: (Name Combination) TAXONOMY: Formerly included in either Miniopterus schreibersii or Miniopterus natalensis, but based on genetic and biometrical data considered a valid species by Farkašová et al. (2017: Suppl.), Kruskop et al. (2014: 102), Kassahun et al. (2015: 168), Farkašová et al. (2017: Suppl.), Wilson and Mittermeier (2019: 704) and Monadjem et al. (2020: 242, 255), although the exact asignment still needs to be confirmed by material from the type locality. COMMON NAMES: Czech: létavec pískový. English: Sandy longfingered bat, Aequatorial Broad Winged Bat. GENERAL DISTRIBUTION: This species occurs in northeastern Africa, from the boundary between Tanzania and Kenya, through Uganda, South Sudan and Ethiopia to the Arabian Peninsula, where it has been reported from Saudi Arabia and Yemen (Harrison and Bates, 1991 [as M. schreibersii], Benda et al., 2011b: 45 [as M. natalensis]). HABITAT: Primarily occurring in Acacia - Commiphora bushlands and thickets Ibáñez and Juste (2019: 704). PARASITES: HAEMOSPORIDA Lutz et al. (2016: 9) examined 42 M. natalensis specimens from East Africa and found 20 of them infected with Polychromophilus melanipherus Dionisi, 1899. Kassahun et al. (2015: 168) examined two M. arenarius specimens from Masha (Ethiopia), and found one of them to be infected by Leishmania parasites. ACARI: Myobiidae: Uchikawa (1985a: 15) corrected the type host for Calcarmyobia rhinolophi from Rhinolophus lobatus to Miniopterus natalensis arenarius. Uchikawa (1985a: 22) mentioned Calcarmyobia kenyaensis Uchikawa, 1982 on "Miniopterus schreibersi" from Belgian Congo and on "Miniopterus schreibersi arenarius" (and M. natalensis arenarius) from Kenya. Uchikawa (1985c: 110) furthermore reported on the presence of Pteracarus miniopteri Uchikawa, 1978 on bats 596 ISSN 1990-6471 Kenya, Uganda (as either M. natalensis arenarius or M. schreibersi arenarius). SIPHONAPTERA: Ischnopsyllidae: Oxyparius isomalus (Waterston, 1915) from the foothills of Mt. Elgon, Kenya (Haeselbarth et al., 1966: 188). Rhinolophopsylla ectopa (Jordan, 1937) found in a cave in the foothills of Mt. Elgon, Kenya (Haeselbarth et al., 1966: 191). The hosts for these fleas might be M. arenarius. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia, Kenya, South Sudan, Sudan, Tanzania, Uganda. Figure 215. Distribution of Miniopterus arenarius Miniopterus brachytragos Goodman, Maminirina, Bradman, Christidis and Appleton, 2009 *2009. Miniopterus brachytragos Goodman, Maminirina, Bradman, Christidis and Appleton, Am. Mus. Novit., 3669: 1, 9, figs 3, 4A–C, 5. Publication date: 30 November 2009. Type locality: Madagascar: Province de Mahajanga: Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National]: Forêt d’Ambovonomby: 26 km NW Andranomavo [16 28.2 S 45 20.9 E, 200 m] [Goto Description]. Holotype: FMNH 175840: ad ♀, skull and alcoholic. Collected by: Steven M. Goodman; collection date: 9 October 2002; original number: SMG 13040. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 172685: Collected by: ?: Collector Unknown; collection date: 5 November 2001. Presented/Donated by: ?: Collector Unknown. Collected in Antsiranana Province, Forêt de Binara, near Analamazava River, 7.5 km SW Daraina (village), 13°15.3'S, 49°37.0'E, 325 - 600 m. Paratype: FMNH 172686: Collected by: ?: Collector Unknown; collection date: 5 November 2001. Presented/Donated by: ?: Collector Unknown. Collected in Antsiranana Province, Forêt de Binara, near Analamazava River, 7.5 km SW Daraina (village), 13°15.3'S, 49°37.0'E, 325 - 600 m. Paratype: FMNH 172867: Collected by: ?: Collector Unknown; collection date: 2 December 2001. Presented/Donated by: ?: Collector Unknown. Collected at the Parc National de Bemaraha, south bank Manambolo River, near Tombeau Vazimba, 3.5 km E Bekopaka, 19°08.4'S, 44°49.7'E, 100 m,. Paratype: FMNH 175846: Collected by: ?: Collector Unknown; collection date: 9 - 10 October 2002. Presented/Donated by: ?: Collector Unknown. Same locality as holotype. Paratype: FMNH 175847: Collected by: ?: Collector Unknown; collection date: 9 - 10 October 2002. Presented/Donated by: ?: Collector Unknown. Same locality as holotype. Paratype: FMNH 175850: Collected by: ?: Collector Unknown; collection date: 9 - 10 October 2002. Presented/Donated by: ?: Collector Unknown. Same locality as holotype. Paratype: FMNH 175851: Collected by: ?: Collector Unknown; collection date: 9 - 10 October 2002. Presented/Donated by: ?: Collector Unknown. Same locality as holotype. Paratype: FMNH 175852: Collected by: ?: Collector Unknown; collection date: 9 - 10 October 2002. Presented/Donated by: ?: Collector Unknown. Same locality as holotype. Paratype: FMNH 175856: Collected by: ?: Collector Unknown; collection date: 9 - 10 October 2002. Presented/Donated by: ?: Collector Unknown. Same locality as holotype. Paratype: FMNH 175864: Collected by: ?: Collector Unknown; collection date: 11 October 2002. Presented/Donated by: ?: Collector Unknown. Collected just outside limit of Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], along Ampandra River, 22 km NW Andranomavo, 16°26.4429'S, 45°24.723'E, 120 m. Paratype: FMNH 175865: Collected by: ?: Collector Unknown; collection date: 11 October 2002. Presented/Donated by: ?: Collector Unknown. Collected just outside limit of Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], along Ampandra River, 22 km NW Andranomavo, 16°26.4429'S, 45°24.723'E, 120 m. African Chiroptera Report 2020 597 Paratype: FMNH 175867: Collected by: ?: Collector Unknown; collection date: 11 October 2002. Presented/Donated by: ?: Collector Unknown. Collected just outside limit of Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], along Ampandra River, 22 km NW Andranomavo, 16°26.4429'S, 45°24.723'E, 120 m. Paratype: FMNH 175869: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175870: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175871: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175872: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175873: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175874: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175875: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175877: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175879: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175880: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175881: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175882: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175883: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 175884: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. 598 ISSN 1990-6471 Paratype: FMNH 175885: Collected by: ?: Collector Unknown; collection date: 14 - 15 October 2002. Presented/Donated by: ?: Collector Unknown. Collected at the Réserve Naturelle Intégrale de Namoroka [status subsequently changed to Parc National], near source of Mandevy River, 32 km NW Andranomavo, 16°22.8'S, 45°20.7'E, 100 m. Paratype: FMNH 188651: Collected by: ?: Collector Unknown; collection date: 14 February 2006. Presented/Donated by: ?: Collector Unknown. Collected in Antsiranana Province, Nosy Komba, 0.9 km SW Ampangorinana, on trail to Station Forestière d’Antanampoera, 13°26.835'S, 48°20.466'E, 100 m. Paratype: FMNH 202523: Collected by: ?: Collector Unknown; collection date: 26 November 2007. Presented/Donated by: ?: Collector Unknown. Collected in Toamasina Province, Masoala Peninsula, 1.8 km S Ambanizana (village), 15°38.291'S, 49°57.944'E, 5 m. Etymology: From the Greek brachys ("short") and tragos, meaning "goat," a word ultimately borrowed into New Latin to mean "tragus," and has been chosen as this taxon is easily distinguished from its congeners by its diminutive tragus (see Goodman et al., 2009c: 17).. (Current Combination) COMMON NAMES: German: Namoroka-Langflügelfledermaus. northwestern part of the island (western or dry deciduous region). CONSERVATION STATUS: Global Justification This species is known from seven sites in the northern and western portions of Madagascar (Goodman and Ramasindrazana, 2013; Goodman unpublished data in Goodman (2017i)), many of which are within existing protected areas. This species is apparently not forest-dependant and by extrapolation not subjected to any important habitat degradation or other human pressures. ECHOLOCATION: Ramasindrazana et al. (2011: 294) recorded 30 calls from 8 Miniopterus brachytragos specimens, and found the following echolocation call values: Frequency of maximum energy: 59.0 ± 1.06 (57.3 - 61.7) kHz, Maximum frequency: 105.8 ± 11.20 (85.0 - 128.0) kHz, Minimum frequency: 55.7 ± 0.84 (54.0 - 57.0) kHz, Call duration: 3.4 ± 0.49 (2.6 - 4.3) ms, Inter pulse interval: 84.6 ± 18.99 (56.1 - 122.5) ms. Assessment History Global 2016: LC ver. 3.1 (2001) (Goodman, 2017i). PARASITES: Rasoanoro et al. (2019: 67) refer to Raharimanga et al. (2003), who reported Trypanosoma sp. from "Miniopterus manavi", but one of these bats belonged to the current species. Regional Endemic to Madagascar, therefore assessment is the same as global. regional DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. MAJOR THREATS: Not known (Goodman, 2017i). Trend:- Unknown (Goodman, 2017i). CONSERVATION ACTIONS: Goodman (2017i) report that this species is known from several protected areas: Bemaraha, LokyManambato, Masoala and Namoroka. GENERAL DISTRIBUTION: Occur on Madagascar, from the Masoala Peninsula in the northeast, the Daraina region in the central northeast, and in the karstic zones of the Namoroka and Bemaraha massifs, as well as on the island of Nosy Komba (see Goodman et al., 2009c: 11). Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the Figure 216. Distribution of Miniopterus brachytragos African Chiroptera Report 2020 599 Miniopterus cf. inflatus Monadjem, Shapiro, Richards, Karabulut, Crawley, Broman Nielsen, Hansen, Bohmann and Mourier, 2020 2020. Miniopterus cf. inflatus Monadjem, Shapiro, Richards, Karabulut, Crawley, Broman Nielsen, Hansen, Bohmann and Mourier, Acta Chiropt., 21 (2): 239. TAXONOMY: Monadjem et al. (2020: 252) found that M. inflatus s.s. from the lower Guinea region isn't closely related fo M. cf. inflatus from eastern and southern Africa, which might indicate that these two forms represent different species, but further investigation remains necessary. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Monadjem et al. (2020: 246) indicate that M. cf. inflatus from eastern and southern Africa has a light pelage. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Malawi, Mozambique, South Africa, Tanzania. Figure 217. Distribution of Miniopterus cf. inflatus Miniopterus egeri Goodman, Ramasindrazana, Maminirina, Schoeman and Appleton, 2011 *2011. Miniopterus egeri Goodman, Ramasindrazana, Maminirina, Schoeman and Appleton, Zootaxa, 2880: 1, 9, figs 3, 4, 5A, 6, 7. Publication date: 17 May 2011. Type locality: Madagascar: Toamasina Province: Sahafina forest: Brickaville, 9.5 km W [18 48 37 S 48 58 48 E, 50 m] [Goto Description]. Holotype: FMNH 209160: ad ♀, skull and alcoholic. Collected by: Steven M. Goodman and Beza Ramasindrazana; collection date: 5 December 2009; original number: SMG 16602. Presented/Donated by: ?: Collector Unknown. - Etymology: in honor of Dr. Judith Eger, Senior Curator, Department of Mammalogy, Royal Ontario Museum, Toronto, for her contribution to taxonomic studies of Old World bats, including Madagascar (see Goodman et al., 2011: 13). (Current Combination) COMMON NAMES: English: Eger's long-fingered bat. Minioptère d'Eger. German: Langflügelfledermaus. French: Eger- Regional Endemic to Madagascar, therefore refer to global assessment. CONSERVATION STATUS: Global Justification This newly described species is known from numerous sites in the eastern portions of Madagascar (Goodman and Ramasindrazana, 2013), several of which are within existing protected areas. This species is apparently not forest-dependant and by extrapolation not subjected to any important habitat degradation or other human pressures (Goodman, 2017j). CONSERVATION ACTIONS: Goodman (2017j) report that this species is known from several protected areas, which include Masoala, Sahafina, and Andohahela. Assessment History Global 2016: LC ver 3.1 (2001) (Goodman, 2017j). Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the GENERAL DISTRIBUTION: Based on current knowledge it is known from scattered sites occuring across two-thirds of Madagascar's lowland area from near sea level to 550 m (Goodman et al., 2011: 15). 600 ISSN 1990-6471 eastern coastal part of the island (eastern humid forest or lowland forest region - humid). Native: Madagascar (Goodman et al., 2011: 15). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 63) could not find a baculum in the one specimen they examined. ECHOLOCATION: Low-duty cycle, frequency modulated/quasi constant (FM/QCF) echolocation call, with a peak frequency of 54.7 kHz (53.2 - 56.3 kHz) and a duration of 2.9 ms (2.5 - 3.4 ms) (Goodman et al., 2011: 8). Ramasindrazana et al. (2011: 294) recorded 16 calls from 2 Miniopterus egeri specimens, and found the following echolocation call values: Frequency of maximum energy: 54.7 ± 1.02 (53.2 - 56.3) kHz, Maximum frequency: 113.8 ± 3.62 (107.0 - 123.0) kHz, Minimum frequency: 49.0 ± 0.52 (48.0 - 50.0) kHz, Call duration: 2.9 ± 0.26 (2.5 - 3.4) ms, Inter pulse interval: 62.6 ± 12.57 (43.2 - 81.1) ms. DIET: Rasoanoro et al. (2015: 63, 64) found that the diet of M. egeri mainly consisted of Coleoptera (51.5 vol %), Hymentoptera (28 %), Lepidoptera (10.8 %), and Isoptera (9.7 %). POPULATION: Structure and Density:- Further information is needed (Goodman et al., 2011: 15). Not known (Goodman, 2017j). Trend:- 2016: Unknown (Goodman, 2017j). In the short term this species does not appear to face a risk of dramatic population decline (Goodman et al., 2011: 15). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. HABITAT: Often associated with disturbed forests with mixed native and introduced trees or at the ecotone between degraded anthropogenic habitats and native forest (Goodman et al., 2011: 15). ROOST: A day roost site of this species was found in a natural rock shelter in the Poste Forestier de Farankaraina surrounded by slightly disturbed natural lowland humid forest (Goodman et al., 2011: 15). Figure 218. Distribution of Miniopterus egeri Miniopterus fraterculus Thomas and Schwann, 1906 *1906. Miniopterus fraterculus Thomas and Schwann, Proc. zool. Soc. Lond., 1906, I: 162. Publication date: 7 June 1906. Type locality: South Africa: Southern Cape Province: Knysna [33 53 S 22 59 E] [Goto Description]. Holotype: BMNH 1905.5.7.18: ad ♂, skin and skull. Collected by: Captain Claude Henry Baxter Grant; collection date: 3 October 1905; original number: 1073. See Thomas and Schwann (1906a: 162). - Comments: Coordinates mentioned as 35 56 S 23 06 E by Thorn et al. (2009: 49). - Etymology: From the Latin meaning "little brother" (see Goodman et al., 2007b: 1222). (Current Combination) 1917. Miniopterus breyeri vicinior J.A. Allen, Bull. Am. Mus. Nat. Hist., 37 (18): 450. Publication date: 29 September 1917. Type locality: Congo (Democratic Republic of the): Oriental province: Uele district: Aba [03 53 N 30 17 E] [Goto Description]. Holotype: AMNH 49019: ad ♂, skin and skull. Collected by: Herbert Lang, James Paul Chapin and The American Museum Congo Expedition; collection date: 16 December 1911; original number: 1770. See Allen (1917: 450). - Comments: Considered a subspecies of fraterculus by Thorn et al. (2009: 49). 1978. M[iniopterus] fraterreulus: Herzit-Straschil and Robinson, Koedoe, 21: 104. (Lapsus) ? Miniopterus cf. fraterculus: ? Miniopterus fraterculus fraterculus: (Name Combination) African Chiroptera Report 2020 TAXONOMY: Meester et al. (1986) state that Harrison and Clancey (1952) and Harrison (1953) have demonstrated that Ellerman et al. (1953) are mistaken in regarding fraterculus as a synonym of M. schreibersii natalensis. Koopman (1966) regards fraterculus instead as a subspecies of the extralimital (to southern Africa) M. minor Peters, 1867, but Hayman and Hill (1971) disagree. Corbet and Hill (1980) retain fraterculus as species, as do Swanepoel et al. (1980) and Koopman (1982). Specimens from Madagascar assigned to M. fraterculus by Peterson et al. (1995) and Simmons (2005: 519) are now considered to be a distinct species: Miniopterus sororculus (Goodman et al., 2007b). COMMON NAMES: Afrikaans: Klein grotvlermuis. Chinese: 小长翼蝠 . Czech: létavec jihoafrický. English: Lesser Long-fingered Bat, Black Clinging Bat, Lesser bent-winged bat. French: Minioptère d'Afrique australe, Petit minioptère. German: Kleine Langflügelfledermaus. Portuguese: Morcego pequeno de dedos compridos. SiSwati: Lilulwane. CONSERVATION STATUS: Global Justification Known from a number of discrete ranges. This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread distribution on the African mainland and its ability to live in a wide variety of habitat types (Schlitter et al., 2008; IUCN, 2009; Monadjem et al., 2017bs). On Madagascar, it is quite rare and further surveys are required. Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bs). 2008: LC ver 3.1 (2001) (Schlitter et al., 2008; IUCN, 2009). 2004: LC ver 3.1 (2001) (Schlitter, 2004g; IUCN, 2004). 1996: EN (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (MacEwan et al., 2016c). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). Swaziland:- 2003: NT B2ab(iv); D2 ver 3.1 (2001) (Monadjem et al., 2003). 601 MAJOR THREATS: There are no major threats to this species on the African mainland (Schlitter et al., 2008; IUCN, 2009; Monadjem et al., 2017bs). CONSERVATION ACTIONS: Schlitter et al. (2008) [in IUCN (2009)] and Monadjem et al. (2017bs) report that this species is presumably present in a number of protected areas. In Tanzania it is present in the Manga Forest Reserve of Tanzania (Doggart et al., 1999b). Further studies are needed to better define the range of this species. GENERAL DISTRIBUTION: Traditionally, Miniopterus fraterculus was considered to be present in Central Africa, East Africa and southern Africa, where it was known from a number of discrete ranges, the largest of which stretched along south-eastern Africa, from Malawi, Mozambique and Zimbabwe, down to South Africa (where it extended along the south coast and its distribution was mostly affected by the annual temperature range (Babiker Salata, 2012: 50)). It was also known from eastern Democratic Republic of Congo near the borders with Uganda and Rwanda (although this population needed to be confirmed as Miniopterus fraterculus), and recorded from localities in Tanzania, Zambia, and Kenya. A small recorded range in Kenya near the border with Tanzania probably does not represent Miniopterus fraterculus. Taylor et al. (2018b: 62) mention Miniopterus cf. fraterculus from Angola, however based on acoustic evidence only. Beja et al. (2019: 388) believe the Angolan records to be misidentifications. ROM has 28 specimens from two localities in Cameroon, which need verification as they are well outside the presently known distribution of the species. This is also the case for the specimens from DRC, Kenya and Nigeria. Specimens from Madagascar assigned to M. fraterculus (Peterson et al. (1995; Russ et al., 2001; Simmons, 2005: 519) are now considered to be a distinct species: Miniopterus sororculus (Goodman et al., 2007b). However, recent changed in the taxonomy of the genus Miniopterus have led to the recognition of several new species, which reduced the distribution of M. inflatus to the southern and eastern parts of the Republic of South Africa, Swaziland and possibly extreme southern 602 ISSN 1990-6471 Mozambique (Ibáñez and Juste, 2019: 706, Monadjem et al., 2020: 239). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 297,363 km 2. In the RSA, its distribution is most affected by the annual temperature range (Babiker Salata, 2012: 50). Native: Mozambique (Monadjem et al., 2010d: 539; Monadjem et al., 2010c: 385 as M. cf. fraterculus); South Africa (Monadjem et al., 2010d: 539); Swaziland (Monadjem et al., 2010d: 539).. DENTAL FORMULA: van der Merwe (1985b: 251) examined 44 skulls and found 95 % of them vestigial teeth between the upper canine and the first premolar. In 44 % of the skulls, the teeth were present on both sides. ECHOLOCATION: Schoeman and Waddington (2011: 291) mention a peak frequency of 62.1 ± 1.7 kHz and a duration of 3.7 ± 0.6 msec for specimens from Durban, South Africa. The 135 calls analysed by Eisenring et al. (2016: SI 2) from bats in the Aberdares Range in Kenya had the following values: PF: 56.2 ± 1.4 (53.3 60.0), HF: 87.1 ± 15.9 (63.8 - 127.6), LF: 52.2 ± 1.6 (47.4 - 56.2), DT: 0.1 ± 0.0 (0.0 - 0.1), DF: 34.9 ± 15.9 (11.5 - 80.1), IPI: 0.8 ± 0.6 (0.3 - 4.8). For calls from specimens from Swaziland, Monadjem et al. (2017c: 179) recorded: Fmin: 58.5 ± 0.50 (58.1 - 58.8) kHz, Fknee: 63.6 ± 1.27 (62.7 64.4) kHz, Fc: 59.2 ± 0.40 (58.9 - 59.5) kHz and duration: 2.9 ± 0.32 (2.7 - 3.1) msec. Trend:- 2016: Unknown (Monadjem et al., 2017bs). 2008: Unknown (Schlitter et al., 2008; IUCN, 2009). REPRODUCTION AND ONTOGENY: Delayed implantation (lasting two-and-a-half months), insemination, ovulation and fertilization occur in the autumn. Embryogenesis proceeds to the blastocyst stage and development is then retarded until arousal in spring, when the blastocyst implants and embryogenisis resumes. The active fetal growth period takes about four months (Bernard, 1980a: 55; Bernard, 1980c). Bernard (1980b: 111) found that the ovarian and uterine function is asymmetrical as ovulations occurred primarily in the left ovary (94 %), whereas all implantations occurred in the right uterine horn. PARASITES: HAEMOSPORIDA Lutz et al. (2016: 9) examined 6 M. cf. fraterculus specimens from East Africa and found two of them infected with the haemosporidan parasite Polychromophilus melanipherus Dionisi, 1899. ACARI Myobiidae: Uchikawa (1985a: 16) reported mites belonging to Calcarmyobia rhinolophia (Radford, 1940) on bats identified as M. fuscus fraterculus from Natal, South Africa. Trombiculidae: Leptotrombidium lawrencei Vercammen-Grandjean and Langston, 1976 (see Stekolnikov, 2018a: 143). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Burundi, Cameroon, Congo (Democratic Republic of the), Eswatini, Kenya, Madagascar, Nigeria, Rwanda, South Africa, Tanzania. Weier et al. (2020: Suppl.) reported on 27 "M. cf. fraterculus calls from the Okavango River Basin with the following characteristics: Fmax: 70.82 ± 8.03 kHz, Fmin: 55.21 ± 14.04 kHz, Fknee: 62.13 ± 1.59 kHz, Fchar: 59.99 ± 1.53 kHz, slope: 36.47 ± 32.94 Sc, duration: 3.88 ± 1.45 msec. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Rautenbach et al. (1993) described the karyotype as having a diploid number 2n = 46, aFN = 50, BA = 6, X = SM, Y = A. Protein / allozyme - Unknown. POPULATION: Structure and Density:- It is common and widespread (Schlitter et al., 2008; IUCN, 2009; Monadjem et al., 2017bs). Figure 219. Distribution of Miniopterus fraterculus African Chiroptera Report 2020 603 Miniopterus gleni Peterson, Eger and Mitchell, 1995 *1995. Miniopterus gleni Peterson, Eger and Mitchell, Faune de Madagascar, Chiroptères, 84: 128, figs 60, 61. Type locality: Madagascar: 20 km S Tuléar: Sarodrano and SaintAugustin, in a marine cave [23 33 S 43 45 E] [Goto Description]. Holotype: ROM 42567: ad ♂, skin and skull. Collection date: 9 May 1967. (Current Combination) TAXONOMY: See Simmons (2005). the north-east (Eger and Mitchell, 2003) and many along the west coast (Goodman et al., 2005a). COMMON NAMES: Czech: létavec Glenův. English: Glen's Longfingered Bat. French: Minioptère malgache. German: Glens Langflügelfledermaus. GENERAL DISTRIBUTION: Miniopterus gleni is endemic to the island of Madagascar where it appears to be widely distributed (Eger and Mitchell, 2003; Goodman et al., 2005a). The areas from where it has yet to be recorded may reflect low sampling effort rather than a genuine absence. North of Onilahy river (incl. Ile Sainte-Marie; Rakotonandrasana and Goodman, 2007: 6). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Muldoon et al. (2009: 1114) report on subfossil remains from the Ankilitelo fauna (late Holocene, appr. 500 years ago). Gunnell et al. (2014: 3) refer to Goodman and Junker (2013) who also report fossils from the Andrahomana cave. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread distribution in Madagascar. It may be locally threatened in parts of its range by hunting and roost disturbance, but is not thought to be declining fast enough to place it in a higher category of threat (Andriafidison et al., 2008e; IUCN, 2009; Monadjem et al., 2017ce). Assessment History Global 2016: LC ver 3.1 (Monadjem et al., 2017ce). 2008: LC ver 3.1 (2001) (Andriafidison et al., 2008e; IUCN, 2009). 2000: LR/nt (IUCN, 2000). 1994: LR/nt ver 2.3 (1994) (Groombridge, 1994). Regional None known. MAJOR THREATS: It is threatened by roost site disturbance. This species is locally hunted in the south-west and north-west of Antananarivo (Goodman, 2006; Goodman et al., 2008d) and as a relatively large species may be subject to similar exploitation elsewhere. There is no evidence that hunting presents a major threat. It is unclear if this species is forest dependent (Andriafidison et al., 2008e; IUCN, 2009; Monadjem et al., 2017ce). CONSERVATION ACTIONS: Andriafidison et al. (2008e) [in IUCN (2009)] and Monadjem et al. (2017ce) report that this species is known to exist in a number of protected areas in Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the coastal areas of northeast, north, west, and the south of the island (western or dry deciduous region). Native: Madagascar (Eger and Mitchell, 2003; Goodman et al., 2005a; Rakotonandrasana and Goodman, 2007: 6) DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 63) could not find a baculum in the three males they examined. ECHOLOCATION: Kofoky et al. (2009: 382) reported the calls of eight individuals, which were characterized by broadband FM/QCF pulses produced at low duty cycle with a frequency of maximum energy at about 44.8 kHz, while the fundamental was always the most intense call and pulses are short duration. Ramasindrazana et al. (2011: 294) recorded 16 calls from 6 Miniopterus gleni specimens, and found the following echolocation call values: Frequency of maximum energy: 42.3 ± 1.29 (40.1 - 44.6) kHz, Maximum frequency: 82.6 ± 8.22 (70.0 - 93.2) kHz, Minimum frequency: 37.4 ± 0.94 (36.0 - 38.9) kHz, Call duration: 3.7 ± 0.49 (3.0 - 4.4) ms, Inter pulse interval: 88.9 ± 17.75 (66.7 - 124.1) ms. HABITAT: Dammhahn and Goodman (2013: 108) mention the lower portion of forest structure and partially open areas as the foraging habitat for this species. 604 ISSN 1990-6471 ROOST: On Madagascar, Wilkinson et al. (2012: 160) reported the typical roosting site to be caves. POPULATION: Structure and Density:- There was a maximum of 90 individuals recorded at a roost in Ankarana (S. G. Cardiff pers. comm.), but otherwise this species always appears to roost in rather small colonies (Robinson et al., 2006). Its relative abundance, as determined by mist netting, varies from rare (Kofoky et al., 2007) to locally common (Robinson et al., 2006). VIRUSES: Astroviridae One of two M. gleni specimens tested by Lebarbenchon et al. (2017: Suppl.) was positive for Astroviridae. Paramyxoviridae Wilkinson et al. (2012: 160) tested 7 individuals from the Mauritius using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Trend:- 2016: Unknown (Monadjem et al., 2017ce). 2008: Unknown (Andriafidison et al., 2008e; IUCN, 2009). Mélade et al. (2016b: 4) found four out of 22 M. gleni specimens they tested to be infected with paramyxoviruses. PARASITES: Lagadec et al. (2012: 1696) and Dietrich et al. (2014: Suppl.) report the presence of spirochaetes bacteria of the genus Leptospira, and which were identified as L. borgpetersenii by Dietrich et al. (2018a: 3). UTILISATION: Goodman (2006) describes the hunting method used by local inhabitants. See also Goodman et al. (2008d) and Jenkins and Racey (2008). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Ramasindrazana et al. (2016: 2) indicate that Litomosa goodmani Martin, Bain, Jouvenet, Raharimanga, Robert & Rousset, 2006 (Onchocercidae, Nematoda) was described from a M. gleni from northern Madagascar. They also recovered Litomosa Clade 1 from this bat species. Perkins and Schaer (2016: Suppl.) report the presence of the haemosporidan Polychromophilus sp. Ramasindrazana et al. (2018: Suppl.) identified this parasite as Polychromophilus melanipherus Dionisi, 1899. Tortosa et al. (2013: 4) and Ramasindrazana et al. (2017: Suppl.) reported the following Nycteribiid bat flies: Nycteribia stylidiopsis, Penicillidia leptothrinax, and Penicillidia sp. The first one was also mentioned by Wilkinson et al. (2016), which was found to carry Enterobacteriales and Bartonella bacteria, as well as the blood parasite Polychromophilus melanipherus (Szentiványi et al., 2019: Suppl.). Figure 220. Distribution of Miniopterus gleni Miniopterus griffithsi Goodman, Maminirina, Bradman, Christidis and Appleton, 2009 *2009. Miniopterus griffithsi Goodman, Maminirina, Bradman, Christidis and Appleton, J. Syst. Evol. Res., 38: 75, 81, figs 4, 6. Publication date: prior to 25 October 2009. Type locality: Madagascar: Province de Toliara: Grotte d'Androimpano: 4.2 km NE Itampolo (village), on old road to Ejeda [24 39.012 S 43 57.797 E, 110m] [Goto Description]. Holotype: FMNH 184214: ad ♂, skull and alcoholic. Collected by: Steven M. Goodman; collection date: 23 February 2005; original number: SMG 14593. Paratype: FMNH 184153: ad ♂. Grotte de Vitane (Vitany), 4.1 km SE Itampolo, 244208.5S 435749.4E, 25m. Paratype: FMNH 184154: ad ♂. Grotte de Vitane (Vitany), 4.1 km SE Itampolo, 244208.5S 435749.4E, 25m. Paratype: FMNH 184167: ad ♂. Grotte de Vitane (Vitany), 4.1 km SE Itampolo, 244208.5S 435749.4E, 25m. Paratype: FMNH 184215: ad ♂. same locatlity as the African Chiroptera Report 2020 605 holotype. Paratype: FMNH 184216: ad ♂. Same locatlity as the holotype. Paratype: FMNH 184236: ad ♀. Same locatlity as the holotype. Paratype: USNM 577076:. Fivondronana de Tolagnaro, 20 km WNW Ranopiso, 12 km ENE Amboasary, near Itaranta River, 2501S 4630E, 40m. Paratype: USNM 577077:. Fivondronana de Tolagnaro, 20 km WNW Ranopiso, 12 km ENE Amboasary, near Itaranta River, 2501S 4630E, 40m. (Current Combination) TAXONOMY: Goodman et al. (2009b) examining the phylogeography of the M. gleni complex, found that the Onilahy River is a major barrier to gene flow. Subsequently, morphological characters (tragus shape, pelage coloration and skull proportions) were identified, that supported the separation of the populations found on either side of the Onilahy River (Goodman et al., 2009b). COMMON NAMES: German: Griffiths Langflügelfledermaus. ETYMOLOGY OF COMMON NAME: In honour of Owen Griffiths, founder of Biodiversity Conservation Madagascar, who is a specialist on the terrestrial mollusc fauna of the western Indian Ocean and has been instrumental in preserving key important areas of habitat on Mauritius and Madagascar. He has generously supported many initiatives for documenting the rich Malagasy fauna (including genetic studies on Miniopterus) (see Goodman et al., 2009a: 346). CONSERVATION STATUS: Global Justification This species is known from few localities in the southern portion of Madagascar (Goodman and Ramasindrazana, 2013). Research is needed to better define the geographical distribution of Miniopterus griffithsi and new field data are necessary to assess properly this taxon (Goodman, 2017k). Assessment History Global 2016: DD ver 3.1 (2001) (Goodman, 2017k). 2009: EN (Goodman et al., 2009b). Regional Endemic to Madagascar assessment should be used. therefore global MAJOR THREATS: Threats to this species are not known (Goodman, 2017k). CONSERVATION ACTIONS: Goodman (2017k) report that this species is not currently known from any protected area. Two of the three sites it has been documented are in degraded forested ecosystems (Goodman et al., 2009b). If indeed it is not forest dependent, these human pressures may not directly impact this species. GENERAL DISTRIBUTION: Miniopterus griffithsii is endemic to Madagascar, where it seems to be restricted to the region between the southern banks of the Onilahy River and the Mandrare River (Itampolo east to near Ranopiso). Schoeman et al. (2014: 28,31) found the most suitable area for this species to be located in the south/southwestern coastal part of the island (south-west sub-arid or spiny bush). Native: Madagascar. ECHOLOCATION: Ramasindrazana et al. (2011: 294) recorded 8 calls from 2 Miniopterus griffithsi specimens, and found the following echolocation call values: Frequency of maximum energy: 44.1 ± 0.67 (43.5 - 45.3) kHz, Maximum frequency: 80.4 ± 15.66 (61.0 - 99.0) kHz, Minimum frequency: 40.0 ± 0.0 (40.0 - 40.0) kHz, Call duration: 3.2 ± 0.29 (2.9 3.6) ms, Inter pulse interval: 89.2 ± 27.42 (56.4 128.5) ms. ROOST: Wilkinson et al. (2012: 160) indicate that caves are the typical roosting sites for these bats on Madagascar. MIGRATION: Reher et al. (2019: 121) found that M. griffithsi was only present in the Vintany Cave (southwestern Madagascar) during the wet season, whereas other species such as Triaenops menamena and Miniopterus mahafaliensis were present in both the dry and wet season. POPULATION: Structure and Density:- Population size and trends are not known for this species (Goodman, 2017k). Trends:- 2016: Unknown (Goodman et al., 2009b). PARASITES: Lagadec et al. (2012: 1696), Dietrich et al. (2014: Suppl.) and Gomard et al. (2016: 5) report the presence of spirochaetes bacteria of the genus 606 ISSN 1990-6471 Leptospira, and which were further identified as L. borgpetersenii by Dietrich et al. (2018a: 3). The haemosporidian Polychromophilus melanipherus Dionisi, 1899 was reported from this species by Ramasindrazana et al. (2018: Suppl.). Ramasindrazana et al. (2016: 6) recovered Litomosa Clade 2 (Nematoda: Onchocercidae) from this bat species. VIRUSES: Paramyxoviridae Wilkinson et al. (2012: 160) tested 2 individuals from the Mauritius using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Figure 221. Distribution of Miniopterus griffithsi DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Miniopterus griveaudi Harrison, 1959 *1959. Miniopterus minor griveaudi Harrison, Durban Mus. Novit., 5 (15): 192. Publication date: 30 April 1959. Type locality: Comoros: Grand Comoro [11 35 S 43 20 E] [Goto Description]. Holotype: BMNH 1967.1231: ad ♀, alcoholic (skull not removed). Collected by: Paul Griveaud; collection date: 10 August 1958. Formerly HZM 3.2803, see Harrison (1959a: 192). - Comments: The museum labels of the two other specimens, on which Harrison (1959a) based his description (HZM 4.2804 and 5.2805), have the following note by P. Griveaud: "couloir souterrain et grotte sous coulée de lave Niombadjée". Niombadjée is a toponym for Niombadjou (also spelled as Niumbadju), located on the western lower flank of Mt. Karthala at 11° 47' 57" S, 43° 17' 50" E (approximately 600 m) and where members of the expedition were based from 8 August 1958 to 23 August 1958: see Goodman et al. (2009a: 342). - Etymology: In honour of Mr. Paul Griveaud, the collector of the type specimen. ? Miniopterus griveaudi (Clade 3): Goodman, Maminirina, Weyeneth, Bradman, Christidis, Ruedi and Appleton, Zool. Scr., 38: 351 - 353, 362 - 363. ? Miniopterus griveaudi: (Current Combination) TAXONOMY: Originally synonomised with Miniopterus minor. Then moved as a synonym of Miniopterus manavi (Peterson et al., 1995: 135; Simmons, 2005: 520). Peterson et al. (1995: 119) suggest that it may be a vaild species, which Simmons (2005: 520) recognises as a valid subspecies. Juste [B.] et al. (2007: 32) consider griveaudi a valid species based on mtDNA data, a view supported by Weyeneth et al. (2008). ETYMOLOGY OF COMMON NAME: In honour of Mr. Paul Griveaud, the collector of the type specimen. COMMON NAMES: Czech: létavec komorský. English: Griveaud`s Long-fingered bat. French: Minioptère des Comores. German: KomorenLangflügelfledermaus. Assessment History Global 2008: DD ver 3.1 (2001) (Juste, 2008a; IUCN, 2009). CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Juste, 2008a; IUCN, 2009). Regional None known. African Chiroptera Report 2020 MAJOR THREATS: The threats to this species are not known (Juste, 2008a; IUCN, 2009). CONSERVATION ACTIONS: Juste (2008a) [in IUCN (2009)] reports that it is not known if the species is present in any protected areas. Further studies are needed into the distribution, abundance, natural history, and threats to this species. GENERAL DISTRIBUTION: Miniopterus griveaudi is endemic to Grand Comore Island in the Comoros. O'Brien (2011: 289) also reports it from Madagascar. Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the northwestern part of the island (western or dry deciduous region). Native: Comoros (Koopman, 1993a: 231 [as minor], Simmons, 2005: 521 [as minor], Juste [B.] et al., 2007: 32). ECHOLOCATION: Kofoky et al. (2009: 382) reported the calls as M. manavi s.l., these calls were broadband FM/QCF sweeps produced at low duty cycle with most energy at about 58.1 kHz in the fundamental, with pulses having a short duration of about 4 ms. Ramasindrazana et al. (2011: 294) recorded 49 calls from 17 Miniopterus griveaudi specimens from Madagascar, and found the following echolocation call values: Frequency of maximum energy: 59.9 ± 1.73 (56.4 - 62.4) kHz, Maximum frequency: 105.6 ± 12.05 (82.0 - 130.0) kHz, Minimum frequency: 55.9 ± 1.36 (53.0 - 58.0) kHz, Call duration: 3.5 ± 0.54 (2.7 - 4.4) ms, Inter pulse interval: 85.7 ± 19.80 (40.0 - 123.6) ms. For 27 calls of 8 specimens from the same species from Anjoun, Ramasindrazana et al. (2011: 294) reported the following echolocation call values: Frequency of maximum energy: 59.2 ± 0.73 (58.2 - 60.8) kHz, Maximum frequency: 104.3 ± 8.86 (86.0 - 123.0) kHz, Minimum frequency: 54.5 ± 1.22 (53.0 - 57.0) kHz, Call duration: 3.2 ± 0.53 (2.7 - 4.6) ms, Inter pulse interval: 83.8 ± 15.60 (57.6 - 128.1) ms. And 22 calls of 6 specimens from Grande Comore gave the following echolocation call values: Frequency of maximum energy: 59.4 ± 0.78 (58.2 - 60.9) kHz, Maximum frequency: 110.0 ± 11.10 (74.0 - 124.0) kHz, Minimum frequency: 55.0 ± 0.76 (53.0 - 56.0) kHz, Call duration: 3.1 ± 0.34 (2.6 - 3.6) ms, Inter pulse interval: 83.5 ± 13.18 (66.1 - 123.9) ms. MOLECULAR BIOLOGY: DNA - Unknown 607 Karyotype - Richards et al. (2010) reported one female with a 2n=46 and a FNa=50. The karyotype consists of two large metacentric, one medium metacentric and 19 acrocentric chromosomal pairs. As FISH with the complete set of Myotis probes was applied, the X chromosome could be identified as a submetacentric element. Volleth and Eick (2012: 167) also mention a segment number of 23. HABITAT: Dammhahn and Goodman (2013: 108) mention the lower portion of forest structure and partially open areas as the foraging habitat for this species. ROOST: Wilkinson et al. (2012: 160) indicated that caves are the typical roosting site for this species on the Comoros. POPULATION: Structure and Density:- The abundance and population size of this species are not known (Juste, 2008a; IUCN, 2009). Trend:- 2008: Unknown (Juste, 2008a; IUCN, 2009). PARASITES: Lagadec et al. (2012: 1696), Lei and Olival (2014: Suppl.) and Gomard et al. (2016: 5) found this species to be infected by Leptospira bacteria on the Comoro Islands. These were further identified as L. borgpetersenii by Dietrich et al. (2018a: 3). The haemosporidian Polychromophilus melanipherus Dionisi, 1899 was reported from this species by Ramasindrazana et al. (2018: Suppl.), as wel as an unidentified Haemosporida sp. Tortosa et al. (2013: 4) reported these bats to be parasitized by Nycteribia stylidiopsis on both the Comoro Islands and on Madagascar. On the latter island, two additional Nycteribiid bat flies were found: Penicillidia leptothrinax and Penicillidia sp. N. stylidiopsis and P. leptothrinax were also reported by Wilkinson et al. (2016) and Ramasindrazana et al. (2017: Suppl.). Wilkinson et al. (2016) furthermore reported the presence of "Penicillidia sp. (cf. fulvida)". The N. stylidiopsis flies were found to carry Enterobacteriales, and the P. cf. fulvida the blood parasite Polychromophilus murinus as well as Enterobacteriales and Bartonella bacteria, whereas P. leptothrinax was carrying Enterobacteriales and Bartonella and Wolbachia bacteria (Szentiványi et al., 2019: Suppl.). 608 ISSN 1990-6471 Ramasindrazana et al. (2016: 6) recovered Litomosa Clades 1, 2 and 3 (Nematoda: Onchocercidae) from this bat species, as well as an unnamed filariod. VIRUSES: Astroviridae 26 bats were tested by Lebarbenchon et al. (2017: Suppl.) of which 15 were positive for Astroviridae. seroreactivity. This was also the case for 10 out of 29 specimens they tested for Duvenhage lyssavirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, Madagascar. Paramyxoviridae Wilkinson et al. (2012: 160) tested 20 individuals from the Comoros using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 3 positive results for viral nucleic acids. Wilkinson et al. (2014) tested 17 individuals from Amabhibe in Madagascar with an RT-PCR specific for the Respiro-, Morbilli- and Henipavirus genera, four of the 17 individuals tested positive for paramyxovirus RNA. 18 out of 116 Madagascan specimens tested by Mélade et al. (2016b: 4) were positive for paramyxoviruses. Figure 222. Distribution of Miniopterus griveaudi Rhabdoviridae Mélade et al. (2016a: 6) tested 33 specimens for Lagos bat lyssavirus and two of them showed Miniopterus inflatus Thomas, 1903 *1903. Miniopterus inflatus Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (72): 634. Publication date: 1 December 1903. Type locality: Cameroon: Efulen [02 46 N 10 42 E] [Goto Description]. Holotype: BMNH 1903.2.4.8: ad ♂, skin only. Collected by: George Latimer Bates Esq. Collection date: 24 July 1901. - Etymology: From the masculine Latin adjective inflatus, meaning "inflated, swollen", referring to its inflated skull (see Lanza et al., 2015: 329). (Current Combination) ? Miniopterus infatus: (Lapsus) TAXONOMY: Koopman (1975: 230) included africanus, but see Peterson et al. (1995: 120), Simmons (2005: 520) and Juste [B.] et al. (2007: 28). Simmons (2005: 520) recognises rufus Sanborn, 1936 as a valid subspecies. Based on cytochrome b and morphometric analyses, Monadjem et al. (2020: 237) found that M. inflatus is paraphyletic with the populations from the central African rainforest not being closely related to the savanna forms from eastern and southern Africa. COMMON NAMES: Afrikaans: Groot grotvlermuis. Castilian (Spain): Murciélago de Ala Larga. Chinese: 大 长 翼 蝠 . Czech: létavec větší. English: Greater Long- fingered Bat, High-crowned bat. French: Grand minioptère africain, Grand minioptère, Minioptère à couronne. German: Aufgeblasene Langflügelfledermaus. Italian: Miniòttero maggióre di Thòmas, Miniòptero maggióre di Thòmas. Portuguese: Morcego grande de dedos compridos. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Schlitter, 2008h; IUCN, 2009; Monadjem and Schlitter, 2017d). African Chiroptera Report 2020 609 Assessment History Global 2016: LC ver 3.1 (2001) [includes africanus Sanborn, 1936] (Monadjem et al., 2017cm). 2008: LC ver 3.1 (2001) [includes africanus Sanborn, 1936] (Schlitter, 2008h; IUCN, 2009). 2004: LC ver 3.1 (2001) (Schlitter, 2004j; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Monadjem et al., 2010d: 539); Zimbabwe (Monadjem et al., 2010d: 539). Presence uncertain: Nigeria (Simmons, 2005: 520). Regional South Africa:- 2016: NT D1 ver 3.1 (2001) (Richards et al., 2016c). MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Ruedas et al. (1990) described the karyotype as having a diploid number of 2n = 46, aFN = 50, BA = 6, X = M, Y = A. Protein / allozyme - Unknown. MAJOR THREATS: There do not appear to be any major threats to this species. Some roosting caves are disturbed by tourism activities (Schlitter, 2008h; IUCN, 2009; Monadjem and Schlitter, 2017d). CONSERVATION ACTIONS: Schlitter (2008h) [in IUCN (2009)] and Monadjem and Schlitter (2017d) report that this species has been recorded from a number of protected areas. There is a need to limit disturbance of important roosting caves. Further studies are needed into the distribution of this species in West Africa. GENERAL DISTRIBUTION: Patchily recorded over much of sub-Saharan Africa. It has been reported from Liberia and Guinea in West Africa; from Cameroon, Equatorial Guinea, Gabon, Central African Republic and the Democratic Republic of the Congo in Central Africa; from Ethiopia, Uganda, Kenya and Tanzania in East Africa; and from Namibia, Zimbabwe and Mozambique in southern Africa. The distribution of this species is somewhat unclear in West Africa due to confusion with records of Miniopterus schreibersii (Simmons, 2005:520). Native: Burundi (Simmons, 2005: 520); Cameroon (Simmons, 2005: 520); Central African Republic; Congo (Bates et al., 2013: 329 - 1st authenticated records); Congo (The Democratic Republic of the) (Schouteden, 1944; Hayman et al., 1966; Simmons, 2005: 520; Monadjem et al., 2010d: 539); Equatorial Guinea; Ethiopia; Gabon (Simmons, 2005: 520); Guinea; Kenya (Simmons, 2005: 520); Liberia (Simmons, 2005: 520); Malawi (Monadjem et al., 2010d: 539); Mozambique (Smithers and Lobão Tello, 1976; Simmons, 2005: 520; Monadjem et al., 2010d: 539; Monadjem et al., 2010c: 385); Namibia (Monadjem et al., 2010d: 539); South Africa (Monadjem et al., 2010d: 539); Tanzania; Uganda (Kityo and Kerbis, 1996: 62; Simmons, 2005: 520); Zambia (Ansell, 1974; DENTAL FORMULA: van der Merwe (1985b: 251) examined three skulls and found vestigial teeth between the upper canine and the first premolar in all of them. HABITAT: In the Mount Nimba area, Monadjem et al. (2016y: 371) recorded this species from a variety of forested and disturbed habitats between 500 and 1,000 m. ROOST: Monadjem et al. (2016y: 371) found a roost of this species in an old mine adit. POPULATION: Structure and Density:- This is generally considered to be a locally rare species, although it can be common in some areas (Schlitter, 2008h; IUCN, 2009; Monadjem and Schlitter, 2017d). Trend:- 2016: Unknown (Monadjem and Schlitter, 2017d). 2008: Unknown (Schlitter, 2008h; IUCN, 2009). REPRODUCTION AND ONTOGENY: Brosset and Saint Girons (1980: 227) indicate that in Gabon all females were pregnant between August and September, and births were taking place in October. Between January and July, none of the females were either having any fetuses or young or were lactating. The males were nonreproductive between 9 October and 10 February. PARASITES: Fain (1959a: 342) described Nycteridocoptes miniopteri (Acari: Sarcoptidae) from a M. inflatus specimen collected in Mulungu, Kivu, DRC. Uchikawa (1985a: 22) reported Calcarmyobia kenyaensis Uchikawa, 1982 (Acari: Myobiidae) from Miniopterus inflatus specimens from Cameroon, Liberia, The Democratic Republic of Congo, Uganda and Kenya. Some Kenyan bats were identified as belonging to Miniopterus inflatus africanus, which might indicate that the host should actually be M. africanus. 610 ISSN 1990-6471 Uchikawa (1985b: 52) further collected two paratypes of Calcarmyobia producta from a M. inflatus specimen from N Chyulu Hills, Kenya. Uchikawa (1985c: 110) furthermore reported on several specimens from Kenya and The Democratic Republic of Congo, infected with four female Pteracarus miniopteri Uchikawa mites. Justine (1989b: 555) described a new stomach parasite: Capillaria magnova from a M. inflatus from Belinga (Gabon). Duval et al. (2012: 1559) indicate that the haemosporidan Polychromophilus (Polychromophilus) corradetti Landau et al., 1980 was described from M. inflatus specimens from Gabon (see Landau et al., 1980b: 22). They also report (p. 1561) on five haplotypes belonging to the Polychromophilus melanipherus group and isolated during their study in Gabon. ACARI Laelapidae: Neospinolaelaps Illiniopteri Zumpt and Patterson, 1952 was reported by Taufflieb (1962: 112). Sarcoptidae: Fain (1959d: 157) described Notoedres (Metanotoedres) miniopteri from M. inflatus from Thysville (=Mbanza-Ngungu), DRC. Spinturnicidae: Taufflieb (1962: 112) reported Spinturnix seIllilunaris de Meillon and Lavolplerre, 1944. DIPTERA: Streblidae: Brachytarsina allaudi (Falcoz, 1923) in Gabon (Obame-Nkoghe et al., 2016: 5). Raymondia huberi Frauenfeld 1856 in GuineaBissau (Haeselbarth et al., 1966: 102). Raymondia seminuda Jobling, 1954 (Shapiro et al., 2016: 255). Raymondia tauffliebi Theodor, 1968 in Congo (Shapiro et al., 2016: 256). Nycteribiidae: Nycteribia exacuta Theodor, 1957 from Dalaba, Guinea (Haeselbarth et al., 1966: 108). Nycteribia schmidlii Schiner, 1853 (Haeselbarth et al., 1966: 108; Tortosa et al., 2013: 5; Duron et al., 2014: 2108; Obame-Nkoghe et al., 2016: 5; Ramasindrazana et al., 2017: Suppl.), which was in Gabon carrying the blood parasite Polychromophilus melanipherus (Szentiványi et al., 2019: Suppl.). Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114; Duron et al., 2014: 2109; Obame-Nkoghe et al., 2016: 5), which in Gabon was also carrying the same blood parasite as above (Szentiványi et al., 2019: Suppl.). Eucampsipoda africana Theodor, 1955 (Obame-Nkoghe et al., 2016: 5). VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibody to SARS-CoV in sera in 34 individuals from Oriental Province, DRC, one tested positive (1/34, 2.9 %). Tong et al. (2009) tested 7/12 bats collected in Kenya during 2006 positive for the presence of coronavirus RNA in a fecal sample. Anthony et al. (2017b: Suppl.) mention the alphacoronaviruses: Kenya_Cov_KY33 and Predict_CoV_28. One out of two Kenyan specimens tested by Tao et al. (2017: Suppl.) was positive for CoV. One out of 249 Gabonese bats (0.4 %) tested by Maganga et al. (2020: 2) was found to be positive for an infection by Alphacoronavirus. Filoviridae - Filoviruses Marburgvirus This virus is reported by Becquart et al. (2010: 1), Kuzmin et al. (2010b: 353), Luis et al. (2013: Suppl), Changula et al. (2014: 486), Simons et al. (2014: 2090), and Willoughby et al. (2017: Suppl.). Markotter et al. (2020: 6) mentoin the bat species as "Miniopterus cf. inflatus" (see also below). Ebolavirus On 24 January 2019, a series of newspaper articles was published, where several scientists from EcoHealth Alliance were quoted (Simon J. Anthony, Jonathan Epstein) stating that in a mouth swab one Liberian Miniopterus inflatus (out of 150 tested) RNA material of the Zaire Ebolavirus was found. Epstein, however, pointed out that if the bat was a natural host for the virus, it would have been found in more than one bat. He also pointed out that the bat could have become infected by another bat species living in the same habitat. Fabian Leendertz (veterinary epidemiologist at the Robert Koch Institute - not involved in the study) indicated that the virus itself wasn't isolated but only about one-fifth of its genome (Grady, 2019; Kupferschmidt, 2019; Su, 2019; Brainard, 2019). Demos et al. (2019c: "11") pointed out that M. inflatus - as currently understood - has a very wide distribution, ad might be paraphyletic, indicating that further research is needed to confirm the true identity of the bat involved. Markotter et al. (2020: 6) therefore listed the bat as "Miniopterus cf. ". Nairoviridae Orthonairovirus 9 out of 51 M. inflatus specimens from Gabon tested by Müller et al. (2016: 3) were positive for Crimean Congo hemorrhagic fever virus (CCHFV). African Chiroptera Report 2020 Paramyxoviridae Orthorubulavirus Drexler et al. (2012a: Suppl. Table S1) indicated that two of the 125 specimens they examined from Gabon tested positive for Rubulavirus (Human orthorubulavirus-related viruses according to Markotter et al., 2020: 6). 611 Madagascar, Namibia, Nigeria, Rwanda, Sierra Leone, South Africa, Uganda, Zambia, Zimbabwe. Polyomaviridae Nieto-Rabiela et al. (2019: Suppl.) mentioned the presence of Bat Polyomavirus. In their overview table, Maganga et al. (2014a: 8) reported the following viruses were already found on M. inflatus: Marburg virus (MBGV), Coronavirus, Rubulavirus. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Cape Verde, Central African Republic, Congo (Democratic Republic of the), Ethiopia, Gabon, Guinea, Kenya, Liberia, Figure 223. Distribution of Miniopterus inflatus Miniopterus inflatus inflatus Thomas, 1903 *1903. Miniopterus inflatus Thomas, Ann. Mag. nat. Hist., ser. 7, 12 (72): 634. Publication date: 1 December 1903. Type locality: Cameroon: Efulen [02 46 N 10 42 E] [Goto Description]. - Etymology: From the masculine Latin adjective inflatus, meaning "inflated, swollen", referring to its inflated skull (see Lanza et al., 2015: 329). (Current Combination) ? Miniopterus inflatus inflatus: (Name Combination, Current Combination) DISTRIBUTION BASED ON VOUCHER SPECIMENS: Burundi, Congo, Congo (Democratic Republic of the), Equatorial Guinea, Kenya, Liberia, Madagascar, Uganda. Miniopterus inflatus rufus Sanborn, 1936 *1936. Miniopterus rufus Sanborn, Field Mus. Nat. Hist., Zool. Ser., (362) 20 (14): 112. Publication date: 15 August 1936. Type locality: Congo (Democratic Republic of the): Tanganyika-Moero: Lualaba River, 40 mi (64 km) below Bukama: Katobwe [08 51 S 26 05 E] [Goto Description]. Holotype: FMNH 29416: ad ♂, skin and skull. Collected by: John Todd Zimmer; collection date: 28 November 1926; original number: 566. ? Miniopterus inflatus rufus: (Name Combination, Current Combination) PARASITES: Lutz et al. (2016: 9) examined 31 "M. rufus" specimens from East Africa and found 22 of them infected with the haemosporidan parasite Polychromophilus melanipherus Dionisi, 1899. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Kenya, Uganda. Miniopterus maghrebensis Puechmaille, Allegrini, Benda, Bilgin, Ibañez and Juste, 2014 *2014. Miniopterus maghrebensis Puechmaille, Allegrini, Benda, Bilgin, Ibañez and Juste, in: Puechmaille, Allegrini, Benda, Gürün, Šrámek, Ibáñez, Juste and Bilgin, Zootaxa, 3794 (1): 108, 112, figs 2, 3a-b, 4a-b, 5a-b. Publication date: 5 May 2014. Type locality: Morocco: Er Rachidiyah Province: 5.7 km S of Ksar Tazougart, 19 km W-NW of Boudenib: Kef Azigza Cave (or Tazzouguert Cave) [32 01 46.6 N 03 47 16.7 W, 1060 m] [Goto 612 ISSN 1990-6471 Description]. Holotype: NMP 94426: ad ♂, skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 28 April 2008; original number: pb3907. Presented/Donated by: ?: Collector Unknown. Paratype: EBD MAM 25780: ad ♂, alcoholic (skull not removed). Collected by: Carlos J. Ibáñez Uargui, Javier Juste, J.A. Garrido and Juan Quetglas; collection date: 30 April 2000. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 90047: ♂, alcoholic (skull not removed). Collected by: Petr Benda; collection date: 30 August 2003. Presented/Donated by: ?: Collector Unknown. Talknout, Oued Tessaoud. Paratype: NMP 90051: ♂, skull and alcoholic. Collected by: Petr Benda; collection date: 30 August 2003. Presented/Donated by: ?: Collector Unknown. Talknout, Oued Tessaoud. Paratype: NMP 90103: ♀, skull and alcoholic. Collected by: Petr Benda; collection date: 9 September 2003. Presented/Donated by: ?: Collector Unknown. Sebt-es-ÂïtSerhrouchèn, Oued El Ammar. Paratype: NMP 94505: alcoholic (skull not removed). Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94506: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94507: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94508: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94509: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94510: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94511: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94512: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94513: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94514: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Paratype: NMP 94515: skull and alcoholic. Collected by: Petr Benda, Jaroslav Cervený, Adam Konecný and Peter Vallo; collection date: 26 April 2008. Presented/Donated by: ?: Collector Unknown. Kef Azigza Cave. Etymology: The name maghrebensis refers to the region (the Maghreb; the region of northern Africa located between the Atlantic Ocean, the Mediterranean Sea, and the Sahara) where the new species was discovered (see Puechmaille et al., 2014a: 118). (Current Combination) TAXONOMY: Bilgin et al. (2016: 330) calculated that the split between M. maghrebensis and M. schreibersii took place some 72,041 years ago (95 % HDP Interval = 5,637 - 139,570 years). COMMON NAMES: Castilian (Spain): Murrcielago de cueva magrebí. Czech: létavec maghrebský. Dutch: Maghrebijns langvleugelvleermuis. English: Maghrebian bentwing bat. French: Minioptere du Maghreb. German: Maghreb-Langflügelfledermaus. CONSERVATION STATUS: Global Justification This species is known from very few sites only, all representing underground spaces threatened by human activities. Listed as Near Threatened (NT ver 3.1 (2001)) because actual levels of disturbance are likely to lead the habitat of this species to decline of at least 30% in the next 15 years (Benda and Piraccini, 2017). Assessment History Global 2016: NT ver 3.1 (2001) (Benda and Piraccini, 2017). African Chiroptera Report 2020 Regional None known. MAJOR THREATS: Threats to the species are mainly human disturbance of roost sites, and perhaps the use of pesticides against insects (Benda and Piraccini, 2017). CONSERVATION ACTIONS: Benda and Piraccini (2017) report that no specific measures are known to be in place for this species, but it occurs in protected areas across its range. GENERAL DISTRIBUTION: Northern Morocco to south of the High Atlas Mountains and northern Tunisia. 613 MOLECULAR BIOLOGY: The GenBank sequence (partial Cytochrome b, tRNA-Threonine, tRNA-Proline, HV1) of the holotype corresponds to the accession numbers KJ535784 & KJ535824 (see Puechmaille et al., 2014a: 117). POPULATION: This species has been found in the Mediterranean and steppe habitats of the Maghreb (Benda and Piraccini, 2017). Trend:- 2016: Unknown (Benda and Piraccini, 2017). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Morocco. SEXUAL DIMORPHISM: Šrámek and Benda (2014: 222) found that most of the cranio-dental measurements in males were larger than in females, except for the dimensions associated with skull and rostral width. In dental characters, on the other hand, females were larger in 38 of 57 variables and the sexes highly diverged (P < 0.001) in the length of the lower canine; slightly diverged in widths of the second lower incisor, second upper molar (in central part) and third lower molar. ECHOLOCATION: Puechmaille et al. (2014a: 113) provide the following data [(avg ± sd (min - max)]: Peak frequency: 52.89 ± 0.74 (51.00 - 54.78) kHz; Start frequency: 74.69 ± 7.66 (63.57 - 101.07) kHz; End frequency: 50.36 ± 0.81 (48.92 - 51.85) kHz; Duration: 4.48 ± 0.67 (3.20 - 5.60) msec; Interval between pulses: 107.14 ± 27.35 (67.20 - 239.20) msec. Figure 224. Distribution of Miniopterus maghrebensis Miniopterus mahafaliensis Goodman, Bradman, Christides and Appleton, 2009 *2009. Miniopterus mahafaliensis Goodman, Bradman, Christides and Appleton, Am. Mus. Novit., 3669: 1, 17, figs 4D-E, 6, 7. Publication date: 30 November 2009. Type locality: Madagascar: Province de Toliara: Parc National de Tsimanampetsotsa: 6.5 km NE Efoetse: near Mitoho Cave [24 03.0 S 43 45.0 E, 50 m]. Holotype: FMNH 173179: ad ♂. Collected by: Steven M. Goodman; collection date: 2 March 2002; original number: SMG 12649. Paratype: FMNH 172918:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172919:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172920:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172921:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172922:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172923:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172924:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172925:; collection date: 7 May 2002. Grotte d’Ambanilia, 614 ISSN 1990-6471 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172926:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172927:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172928:; collection date: 7 May 2002. Grotte d’Ambanilia, 3.7 km SSE Sarodrano, 23°32.397' S, 43°44.763' E, sea level. Paratype: FMNH 172929:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 172930:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 172931:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 172932:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 172933:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 173154:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173189:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173190:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173191:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173192:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173193:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173194:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173195:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173196:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173198:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173199:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173200:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173201:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173202:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173203:. Parc National de Tsimanampetsotsa, 6.5 km NE Efoetse, near Mitoho Cave, 24°03.0' S, 43°45.0' E, 50 m. Paratype: FMNH 173204:; collection date: 4 March 2002. Parc National de Tsimanampetsotsa, Malaza Manga Aven, 24°1.827' S, 43°45.283' E, 80 m. Paratype: FMNH 173252:; collection date: 24 July 2001. Antafiky, 23°29'16.0" S, 44°04'39.1" E, 50 m. Paratype: FMNH 173253:; collection date: 4 February 2002. Antafiky, 23°29'16.0" S, 44°04'39.1" E, 50 m. Paratype: FMNH 175991:; collection date: 9 December 2002. Province de Fianarantsoa, Parc National de l’Isalo, along Sahanafa River, at foot of Bevato, 28 km SE Berenty-Betsileo, 22°19.0' S, 45°17.6' E, 550 m. Paratype: FMNH 176096:; collection date: 11 November 2002. Province de Toliara, Parc National de Kirindy-Mite, 13 km W Marofihitsa, 20°47.4' S, 44°08.8' E, 30 m. Paratype: FMNH 176097:; collection date: 11 November 2002. Province de Toliara, Parc National de Kirindy-Mite, 13 km W Marofihitsa, 20°47.4' S, 44°08.8' E, 30 m. Paratype: FMNH 176098:; collection date: 11 November 2002. Province de Toliara, Parc National de Kirindy-Mite, 13 km W Marofihitsa, 20°47.4' S, 44°08.8' E, 30 m. Paratype: FMNH 176099:; collection date: 11 November 2002. Province de Toliara, Parc National de Kirindy-Mite, 13 km W Marofihitsa, 20°47.4' S, 44°08.8' E, 30 m. Paratype: FMNH 176100:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176101:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176102:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176103:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176104:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: African Chiroptera Report 2020 615 FMNH 176105:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176106:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176107:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176108:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176109:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176110:; collection date: 16-17 November 2002. Parc National de Kirindy-Mite, near village of Betakilotra, 11 km SE Marofihitsa, 20°53.2' S, 44°04.8' E, 35 m. Paratype: FMNH 176167:; collection date: 14, 15, and 18 February 2003. Forêt des Mikea, 9.5 km W Ankililoaka, 22°46.7' S, 43°31.4' E, 80 m. Paratype: FMNH 176168:; collection date: 14, 15, and 18 February 2003. Forêt des Mikea, 9.5 km W Ankililoaka, 22°46.7' S, 43°31.4' E, 80 m. Paratype: FMNH 176169:; collection date: 14, 15, and 18 February 2003. Forêt des Mikea, 9.5 km W Ankililoaka, 22°46.7' S, 43°31.4' E, 80 m. Paratype: FMNH 176170:; collection date: 14, 15, and 18 February 2003. Forêt des Mikea, 9.5 km W Ankililoaka, 22°46.7' S, 43°31.4' E, 80 m. Paratype: FMNH 176508:; collection date: 4 June 2002. Mahaleotse, 23°31'38.8" S, 44°05'16.1" E, 70 m. Paratype: FMNH 176509:; collection date: 27 November 2002. Fiherenana, 23°14'17" S, 43°52'23" E. Paratype: FMNH 176510:; collection date: 27 November 2002. Fiherenana, 23°14'17" S, 43°52'23" E. Paratype: FMNH 177376:; collection date: 16 August 2003. Ranobe, 23°14.033' S, 43°52.493' E, approximately 50 m. Paratype: FMNH 184155:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184156:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184157:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184158:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184159:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184160:; collection date: 29 May 2005. Grotte d’Amborombe, 4.0 km Itampolo, 24°37'42.7" S, 43°58'56.9" E, 70 m. Paratype: FMNH 184168:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184217:; collection date: 18 February 2005. 10.5 km SE Itampolo (village), 24°44.2' S, 44°1.79' E, 120 m. Paratype: FMNH 184218:; collection date: 18 February 2005. 10.5 km SE Itampolo (village), 24°44.2' S, 44°1.79' E, 120 m. Paratype: FMNH 184219:; collection date: 18 February 2005. 10.5 km SE Itampolo (village), 24°44.2' S, 44°1.79' E, 120 m. Paratype: FMNH 184220:; collection date: 18 February 2005. 10.5 km SE Itampolo (village), 24°44.2' S, 44°1.79' E, 120 m. Paratype: FMNH 184221:; collection date: 18 February 2005. 10.5 km SE Itampolo (village), 24°44.2' S, 44°1.79' E, 120 m. Paratype: FMNH 184222:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184223:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184224:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184225:. Grotte d’Andraimpano, 4.2 km NE Itampolo, 24°39.012' S, 43°57.797' E.m,. Paratype: FMNH 184226:; collection date: 26 February 2005. Grotte d’Antagneotsy, 5.0 km NE Vohombe, 24°23.001' S, 43°50.742' E, 100 m. Paratype: FMNH 184470:; collection date: 1 November 2004. Ihosy, commune rurale d’Akily, Grotte d’Andranomilitry, 22°23.111' S, 46° 03.355' E, 950 m. Paratype: FMNH 184471:; collection date: 1 November 2004. Ihosy, commune rurale d’Akily, Grotte d’Andranomilitry, 22°23.111' S, 46° 03.355' E, 950 m. Paratype: FMNH 184472:; collection date: 1 November 2004. Ihosy, commune rurale d’Akily, Grotte d’Andranomilitry, 22°23.111' S, 46° 03.355' E, 950 m. Paratype: FMNH 184473:; collection date: 1 November 2004. Ihosy, commune rurale d’Akily, Grotte d’Andranomilitry, 22°23.111' S, 46° 03.355' E, 950 m. Paratype: FMNH 202484:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 202485:; collection date: 8 May 2002. Grotte de Bisihiko, 0.75 km E St. Augustin, 23°32.933' S, 43°46.044' E, 10 m. Paratype: FMNH 75774:; collection date: 18 June 1953. Betroka, 23°15'50" S, 46°05'30" E, 800 m. Etymology: The name mahafaliensis is derived from the Malagasy word mahafaly, meaning ‘‘to make taboos,’’ but here specifically referring to one of the local cultural groups 616 ISSN 1990-6471 in southwestern Madagascar, the Mahafaly, and the Mahafaly Plateau, a limestone karst area, from whence many of the specimens of this taxon were collected. In the region of the Forêt des Mikea (see Goodman et al., 2009c: 28). (Current Combination) COMMON NAMES: German: Mahafaly-Langflügelfledermaus. Malagasy: kitrotroke. CONSERVATION STATUS: Global Justification This species is known from numerous sites in the dry southern portion of Madagascar (Goodman and Ramasindrazana, 2013); a few of these localities are within existing protected areas. This species is apparently not forest-dependant and may not be very sensitive to any important habitat degradation or other human pressures (Goodman, 2017l). Assessment History Global 2016: LC ver. 3.1 (2001) (Goodman, 2017l). Regional Endemic to Madagascar assessment should be used. therefore global CONSERVATION ACTIONS: Goodman (2017l) this species is known from several protected areas, which include: Isalo, Kirindy Mité, Mikea, and Tsimanampetsotsa. GENERAL DISTRIBUTION: Madagascar (known from numerous sites on or in close proximity to the limestone Mahafaly Plateau in the southwestern portion of the island, including the zone from Itampolo north across the Onilahy River to Sarodrano spanning the elevational range from sea level to 120 m. Its distribution then continues further north to the Forêt des Mikea (70 - 80 m) and based on current data ends in the Kirindy-Mite region (30 - 35 m). Further, it is known from the more inland and upland sites of Ihosy and near Betroka from 800 - 950 m and within the Isalo formation at 550 m.). Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the southwestern quarter of the island (south-west sub-arid or spiny bush). DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 63) could not find a baculum in the one specimen they examined. ECHOLOCATION: Ramasindrazana et al. (2011: 294) recorded 24 calls from 9 Miniopterus mahafaliensis specimens, and found the following echolocation call values: Frequency of maximum energy: 59.6 ± 1.46 (57.3 - 62.2) kHz, Maximum frequency: 113.7 ± 8.08 (95.0 - 123.0) kHz, Minimum frequency: 55.1 ± 1.47 (53.0 - 57.0) kHz, Call duration: 3.3 ± 0.25 (2.9 - 3.8) ms, Inter pulse interval: 68.7 ± 14.04 (43.5 - 95.3) ms. ROOST: Wilkinson et al. (2012: 160) reported caves to be the typical roosting site for these bats on Madagascar. DIET: Ramasindrazana et al. (2012: 120-121) examined feacal pellets from bats in the Tsimanampetsotsa NP, and found them to contain 47.5 ± 5.5 volume percent Lepidoptera, 39.9 ± 5.5 Coleoptera, 6.9 ± 2.9 Hymenoptera, 3.4 ± 0.6 Psocoptera, 1.1 ± 0.4 Diptera, 1.0 ± 0.2 Aranea, and 0.1 ± 0.0 Homoptera/Hemiptera. During the dry season, 52.9 ± 6.1 volume percent Coleoptera were found and 35.2 ± 5.7 Lepidoptera. The latter order was represented by 82.3 ± 7.7 percent in the wet season, whereas the Coleoptera dropped to 3.2 ± 0.8 percent. POPULATION: Structure and Density:- Not known (Goodman, 2017l). Trends:- Unknown (Goodman, 2017l). PARASITES: Lagadec et al. (2012: 4696), Dietrich et al. (2014: Suppl.), Lei and Olival (2014: Suppl.) and Gomard et al. (2016: 5) report the presence of spirochaetes bacteria of the genus Leptospira, which were identified as L. borgpetersenii by Dietrich et al. (2018a: 3). The haemosporidian Polychromophilus melanipherus Dionisi, 1899 was reported from this species by Ramasindrazana et al. (2018: Suppl.), as wel as an unidentified Haemosporida sp. Rasoanoro et al. (2019: 67) refer to Raharimanga et al. (2003) who reported Trypanomsoma sp. from a "Miniopterus manavi". Ramasindrazana et al. (2016: 6) recovered Litomosa Clade 2 (Nematoda: Onchocercidae) from M. mahafaliensis. African Chiroptera Report 2020 617 On Madagascar, Tortosa et al. (2013: 4) found Nycteribiid bat flies belonging to Penicillidia leptothrinax. VIRUSES: Paramyxoviridae Wilkinson et al. (2012: 160) tested 11 individuals from the Mauritius using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. In their extensive study, Mélade et al. (2016b: 3) found that M. mahafaliensis was the least infected species of Madagascan bat, with only 4 out of 89 specimens (4.5 %) infected. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. Figure 225. Distribution of Miniopterus mahafaliensis Miniopterus majori Thomas, 1906 *1906. Miniopterus Majori Thomas, Ann. Mag. nat. Hist., ser. 7, 17 (98): 175. Publication date: 1 February 1906. Type locality: Madagascar: SE Betsileo: Imasindrary [=Sahamananina] [20 17 S 47 31 E, 1 600 m] [Goto Description]. Holotype: BMNH 1897.9.1.38: ad ♀, skin and skull. Collected by: Dr. Charles Immanuel Forsyth Major; collection date: 3 July 1895; original number: 457. - Etymology: In honour of Dr. C.I. Forsyth Major, collector of the type specimen (see Thomas, 1906a: 176). (Current Combination) ? Miniopterus majori majori: (Name Combination) ? Miniopterus majori: (Current Spelling) TAXONOMY: Included in schreibersi by Koopman (1993a: 231), but considered a valid species by Peterson et al. (1995: 131), Hutson et al. (2001: 32), Russ et al. (2001), Simmons (2005: 520). COMMON NAMES: Czech: létavec východomadagaskarský. English: Major Long-fingered Bat, Major's Longfingered Bat. French: Grand minioptère malgache. German: Majors Langflügelfledermaus. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its relatively wide range, ability to live in a wide variety of habitats from intact forests to heavily degraded areas, and because there is no evidence of a decline that would warrant listing in a higher category of threat (Jenkins and Rakotoarivelo, 2008a; IUCN, 2009; Monadjem et al., 2017br). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017br). 2008: LC ver 3.1 (2001) (Jenkins and Rakotoarivelo, 2008a; IUCN, 2009). 2003: DD (IUCN, 2003). 2001: DD (IUCN/SSC Action Plan, 2001). Regional None known. MAJOR THREATS: The major threats to this species are not known. It could be susceptible to cave disturbance in some sites and it could perhaps be hunted in some areas (Jenkins and Rakotoarivelo, 2008a; IUCN, 2009; Monadjem et al., 2017br). CONSERVATION ACTIONS: Jenkins and Rakotoarivelo (2008a) [in IUCN (2009)] and Monadjem et al. (2017br) report that this species is reported from a few protected areas, such as Parc National de Masoala and Parc National de Mantadia (Russ and Bennet, 1999; Randrianandriananina et al., 2006). Some of these records may need to be reassessed in view 618 ISSN 1990-6471 of the ongoing taxonomy of this genus in Madagascar (Goodman and Maminirina, 2007). GENERAL DISTRIBUTION: Miniopterus majori is endemic to Madagascar and has a sympatric distribution with Miniopterus sororculus in the highlands of Madagascar (Goodman et al., 2007b) and is widely, but patchily, distributed across the island (Peterson et al., 1995; Eger and Mitchell, 2003). Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the eastern part of the island (higher elevations of the Central Highlands - sub-humid region). Goodman and Maminirina (2007) suggest that the only specimens known form the Comoro Islands were collected by M. Humbolt and may have been incorrectly provenanced. Until new material is available from the island to confirm that M. majori exists, it is removed from the known distribution of the species, and is restricted at this time to Madagascar. Native: Madagascar (Peterson et al., 1995; Eger and Mitchell, 2003; Simmons, 2005: 520; Goodman and Maminirina, 2007). Presence incorrectly assigned: Comoro Islands (Simmons, 2005: 520). ECHOLOCATION: Kofoky et al. (2009: 382) reported the calls as Miniopterus majori/sororculus, which produced broadband FM/QCF echolocation calls at low duty cycle, but with a lower maximum energy, at about 48.5 kHz. The fundamental being most intense with short duration pulses of about 4.5 ms. Ramasindrazana et al. (2011: 294) recorded 39 calls from 12 Miniopterus majori specimens, and found the following echolocation call values: Frequency of maximum energy: 48.5 ± 1.15 (46.1 - 52.0) kHz, Maximum frequency: 82.5 ± 11.78 (64.0 - 102.0) kHz, Minimum frequency: 44.4 ± 0.97 (43.0 - 46.0) kHz, Call duration: 3.6 ± 0.42 (2.8 - 4.5) ms, Inter pulse interval: 85.0 ± 16.34 (59.5 - 134.8) ms. DIET: Kemp et al. (2018: Suppl.) used DNA metacarcoding to detect insect pest species in the diet of these bats and found the following prey orders (in descending order): Lepidoptera (Ericeia inangulata (Guenée, 1852), Pandemis sp., Celsumaria elongata Brown, 2017, Meyrickiella homosema (Meyrick, 1887), Emmalocera sp., Palpita sp., Pandemis retroflua Diakonoff, 1960, Achaea euryplaga (Hampson, 1913), Eublemma viettei (Berio, 1954), Herpetogramma licarsisalis (Walker, 1859), Chloroclystis androgyna Herbulot, 1957, Idiodes oberthuri Dognin 1911, Scopula AH01Md, Ectropis AH27Md), Hemiptera, Blattodea, Trombidiformes, Diptera (Simulium lineatum (Meigen, 1804)), Coleoptera, Hymenoptera, Ephemeroptera, Trichoptera, Sarcoptiformes, Astigmata POPULATION: Structure and Density:- There is no information on its population status (Jenkins and Rakotoarivelo, 2008a; IUCN, 2009; Monadjem et al., 2017br). Trend:- 2016: Unknown (Monadjem et al., 2017br). 2008: Unknown (Jenkins and Rakotoarivelo, 2008a; IUCN, 2009). PARASITES: Dietrich et al. (2014: Suppl.) and Gomard et al. (2016: 5) report the presence of spirochaetes of the genus Leptospira in bats from Fianarantsoa, Madagascar. These were identified as L. borgpetersenii by Dietrich et al. (2018a: 3). Uchikawa (1985b: 45) reports Calcarmyobia comoresensis mites of bat specimens from Vobima and Bersroha, Madagascar. Uchikawa (1985c: 111) mentions the presence of Pteracarus miniopteri Uchikawa, 1978 mites on M. schreibersi majori specimens from Madagascar. Tortosa et al. (2013: 4), furthermore, reported the presence of Nycteribiid bat flies: Nycteribia stylidiopsis and Penicillidia leptothrinax. The Nycteribia is also reported by Ramasindrazana et al. (2017: Suppl.). Ramasindrazana et al. (2016: 6) recovered Litomosa Clade 1 (Nematoda: Onchocercidae) from this bat species. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar. Figure 226. Distribution of Miniopterus majori African Chiroptera Report 2020 619 Miniopterus manavi Thomas, 1906 *1906. Miniopterus manavi Thomas, Ann. Mag. nat. Hist., ser. 7, 17 (98): 176. Publication date: 1 February 1906. Type locality: Madagascar: Eastern/Northeastern Betsileo: Fandriana region [20 17 S 47 31 E] [Goto Description]. Holotype: BMNH 1897.9.1.37: ad ♂, skin and skull. Collected by: Dr. Charles Immanuel Forsyth Major; collection date: 3 July 1895; original number: 453. See Peterson et al. (1995: 135). Topotype: MCZ 45101: ♀, complete skeleton. Collected by: The Grandidier Collection; collection date: 3 July 1895. Presented/Donated by: ?: Collector Unknown. - Comments: [Imansindrary = ? Masindrary, 15 km ESE Fandriana, see Jenkins and Carleton, 2005 (in Goodman et al., 2009a: 340)]. (Current Combination) 1993. Miniopterus minor grivaudi: Koopman, in: Wilson and Reeder, Mammal species of the World (2nd Edition): Chiroptera, 231. (Lapsus) 2007. Miniopterus (griveaudi) manavi: Goodman and Maminirina, Mammalia, 71 (4): 151. (Name Combination) 2009. Miniopterus manavi (Clade 1): Goodman, Maminirina, Weyeneth, Bradman, Christidis, Ruedi and Appleton, Zool. Scr., 38: 346-351, 362. ? Miniopterus manavi manavi: (Name Combination) TAXONOMY: Included in minor by Koopman (1993a: 231), but considered a valid species by Peterson et al. (1995: 135), Hutson et al. (2001: 32), Russ et al. (2001), and Simmons (2005: 520). (IUCN, 2003). 2001: DD [as M. menavi] (IUCN/SSC Action Plan, 2001). 2000: DD. Simmons (2005: 520) recognised griveaudi Harrison, 1959 as a vaild subspecies of M. manavi, Juste [B.] et al. (2007: 33) elevated griveaudi as a separate species, which is followed in this account. MAJOR THREATS: There are no major threats to the species. In Makira, this species is eaten by people (Golden, 2005), but it is not known if this species is hunted in other parts of its range. It is probably not forest dependent, but is rarely netted in areas without vegetation stands (Andriafidison et al., 2008f; IUCN, 2009; Monadjem et al., 2017cf). Weyeneth et al. (2008) used mitochondrial DNA markers to demonstrate that the specimens from the Comoro islands are not closely related to M. minor from the African mainland, and that these islands were probably populated as a result of two or three invasions originating from Madagascar. COMMON NAMES: Czech: létavec manavijský. English: Manavi Long-fingered Bat, Manavil Long-fingered Bat. French: Petit minioptère malgache. German: Manavi-Langflügelfledermaus. CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread distribution (near suitable caves) across Madagascar. While it is locally threatened by hunting, this is not thought to be a major threat of this species across its range (Andriafidison et al., 2008f; IUCN, 2009; Monadjem et al., 2017cf). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017cf). 2008: LC ver 3.1 (2001) (Andriafidison et al., 2008f; IUCN, 2009). 2003: DD [as M. menavi] Regional None known. CONSERVATION ACTIONS: Andriafidison et al. (2008f) [in IUCN (2009)] and Monadjem et al. (2017cf) report that this species occurs in many of the forest protected areas in western and eastern Madagascar and as such does not require specific conservation measures. Additional information on its ecology would be helpful to better understand how it responds to deforestation and roost disturbance. There is some uncertainty about the taxonomy of M. manavi and that this taxon is actually a complex of new species (F. Ratromomanarivo pers. comm.). Further studies on morphology, acoustics and genetics are therefore needed. GENERAL DISTRIBUTION: Miniopterus manavi is found on Madagascar including the islands of Nosy Be and Nosy Komba (Rakotonandrasana and Goodman, 2007: 6), where it is widely distributed (Eger and Mitchell, 2003; Goodman et al., 2005a). It has a wide elevational range, from 20 to 1,500 m above sealevel. 620 ISSN 1990-6471 Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the eastern part of the island, but not along the coast (higher elevations of the Central Highlands - subhumid region). POPULATION: Structure and Density:- It is locally abundant in cavities or rocky overhangs that provide suitable roosting features. A colony of approximately 4,000 was found in Makira (Bayliss and Hayes, 1999). Grand Comoro (Peterson et al., 1995; Simmons, 2005: 520). Removed by Juste [B.] et al. (2007: 33), due to the removal of griveaudi as a separate species. Trend:- 2016: Unknown (Monadjem et al., 2017cf). 2008: Unknown (Andriafidison et al., 2008f; IUCN, 2009). Native: Madagascar (Peterson et al., 1995; Eger and Mitchell, 2003; Goodman et al., 2005a; Simmons, 2005: 520; Rakotonandrasana and Goodman, 2007: 6). PARASITES: Gomard et al. (2016: 5) reported the presence of Leptospira bacteria in 5 out of 7 tested bats (73.7 %). ECHOLOCATION: Russ et al. (2003) describe the echolocation as a freqency-modulated sweep that terminates in a constant-frequency portion with maximum energy around 57 kHz. Kofoky et al. (2009: 382) reported the calls as M. manavi s.l., these calls were broadband FM/QCF sweeps produced at low duty cycle with most energy at about 58.1 kHz in the fundamental, with pulses having a short duration of about 4 ms. Low-duty cycle, frequency modulated/quasi constant (FM/QCF) echolocation call, with a peak frequency of 57.2 kHz (55.5 - 58.2 kHz) and a duration of 2.5 ms (2.1 - 3.0 ms) (Goodman et al., 2011: 9). Megali et al. (2010: 1042) and Duval et al. (2012: 1563) reported Genbank data for Polychromophilus sp. (Haemosporida) from M. manavi. Ramasindrazana et al. (2018: Suppl.) identified the parasite as Polychromophilus melanipherus Dionisi, 1899, and the bat as "M. manavi s.l.". Raharimanga et al. (2003: 72) reported Trypanosoma sp. from M. manavi, but these bats were later re-identified as M. mahafaliensis and M. brachytragos (see Rasoanoro et al., 2019: 67). Ramasindrazana et al. (2011: 294) recorded 9 calls from 2 specimens tentatively assigned to Miniopterus 'manavi', and found the following echolocation call values: Frequency of maximum energy: 57.2 ± 0.77 (55.5 - 58.2) kHz, Maximum frequency: 98.4 ± 7.60 (89.0 - 110.0) kHz, Minimum frequency: 53.0 ± 0.0 (53.0 - 53.0) kHz, Call duration: 2.5 ± 0.32 (2.1 - 3.0) ms, Inter pulse interval: 66.0 ± 8.70 (54.1 - 84.7) ms. DIET: Razakarivony et al. (2005) found Araneae in 2 % of stomachs examined. Rakotoarivelo et al. (2007), in Madagascar, found M. manavi to mainly consume Hemiptera. Kemp et al. (2018: Suppl.) used DNA metacarcoding to detect insect pest species in the diet of these bats and found the following prey orders (in descending order): Lepidoptera (Pandemis retroflua Diakonoff, 1960), Hymenoptera, Coleoptera, Hemiptera, Blattodea, Diptera (Simulium lineatum (Meigen, 1804), Anopheles squamosus Theobald, 1901, Coquillettidia sp.), Ephemeroptera, Mesostigmata, Sarcoptiformes, Trombidiformes, Astigmata, Neuroptera. Uchikawa (1985b: 45) reported a mite belonging to Calcarmyobia comoresensis from the type specimen of M. manavi. From the same specimen, Uchikawa (1985c: 110) also reported mites belonging to Pteracarus miniopteri Uchikawa, 1978. Furtermore, he also described two mite subspecies: Calcarmyobia steatosetae steatosetae and Calcarmyobia steatosetae rectipenis from several other bat specimens. The presence of the Nycteribiid bat fly Penicillidia leptothrinax was reported by Tortosa et al. (2013: 4) and Ramasindrazana et al. (2017: Suppl.). Additionally, Ramasindrazana et al. (2017: Suppl.) reported both "Penicillidia sp. and P. leptothrinax from one and the same bat identified as "Miniopterus cf. manavi". Ramasindrazana et al. (2016: 6) reported the presence of Litosoma Clade 1 (Nematoda: Onchocercidae) on this bat species. VIRUSES: Zeghbib et al. (2019: Suppl.) refer to the presence of a Hepatovirus. UTILISATION: In Makira, this species is eaten by people (Golden, 2005), also see Goodman et al. (2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Comoros, Madagascar. African Chiroptera Report 2020 621 Figure 227. Distribution of Miniopterus manavi Miniopterus minor Peters, 1867 *1867. Miniopterus minor Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 870 (for 1866). Publication date: 1867. Type locality: Tanzania: Coast opposite Zanzibar Island [Goto Description]. Holotype: ZMB 3268: ♂, skin and skull. Collected by: Baron Karl Klaus von der Decken. See Turni and Kock (2008: 52). - Comments: Peters (1867a: 885) mentions "Ein einziges ausgewachsenes Männchen von der Küste von Zanzibar". - Etymology: From the masculine Latin adjective minor, meaning "smaller", used as comparative of parvus, meaning "small", referring to the relatively small size of the species (see Lanza et al., 2015: 334). (Current Combination) 1992. Miniopterus minor occidentalis Juste and Ibáñez, Bonn. zool. Beitr., 43 (3): 358, 362. Publication date: 21 October 1992. Type locality: Congo: Koilou, Meya-Nzouari Cave [03 53 S 14 31 E]. Holotype: MHNG 1074.013: ad ♀, skull and alcoholic. Collected by: M. Taufflieb; collection date: 1 July 1961. - Comments: (see Simmons, 2005). - Etymology: From the Latin for western. (Name Combination) ? Miniopterus minor minor: (Name Combination) TAXONOMY: Koopman (1993a: 231) included griveaudi Harrison, 1959 and manavi Thomas, 1906, but Peterson et al. (1995: 119, 135) and Simmons (2005: 520) recognised manavi as a distinct species, with griveaudi as a subspecies. Juste [B.] et al. (2007: 33) elevated griveaudi as a separate species. Simmons (2005: 521) recognised newtoni Bocage, 1889 and occidentalis Juste and Ibáñez, 1992 as vaild subspecies of M. minor. Juste [B.] et al. (2007: 33) elevated newtoni as a separate species. Reviewed by Juste and Ibáñez (1992). COMMON NAMES: Chinese: 侏 长 翼 蝠 . Czech: létavec nejmenší. English: Least Long-fingered Bat. French: Petit minioptère africain, Minioptère minuscule. German: Zwerg-Langflügelfledermaus. Italian: Miniòttero minóre, Miniòptere minóre. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of continuing problems with its taxonomy as well as absence of recent information on its extent of occurrence, ecological requirements and threats (Jacobs et al., 2008n; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) [includes occidentalis Juste and Ibáñez, 1992] (Jacobs et al., 2008n; IUCN, 2009). 2004: NT ver 3.1 (2001) (Jacobs et al., 2004o; IUCN, 2004). 2003: LR/nt (IUCN, 2003). 2001: LR/nt (IUCN/SSC Action Plan, 2001). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. 622 ISSN 1990-6471 MAJOR THREATS: This species is presumably threatened in parts of its range by habitat loss resulting from logging operations and the conversion of land to agricultural use. CONSERVATION ACTIONS: Jacobs et al. (2008n) [in IUCN (2009)] report that there appear to be no direct conservation measures in place. It has been recorded from the Kambai Forest Reserve in Tanzania by Cunneyworth (1996b). Further studies are needed into the taxonomy, distribution, natural history and threats to this poorly known species. GENERAL DISTRIBUTION: Miniopterus minor recorded from very few locations over a wide area. It has been reported from a small area on the border between Congo and the Democratic Republic of the Congo, a single locality in southwestern Democratic Republic of the Congo, and from a small area of coastal Kenya and Tanzania (including the island of Unguja [see O'Brien, 2011: 289]). São Tomé (Koopman, 1993a: 231; Simmons, 2005: 521) removed by Juste [B.] et al. (2007: 33) due to the removal of newtoni as a separate species. Native: Angola; Congo (Juste and Ibáñez, 1992: 363; Simmons, 2005: 521; Bates et al., 2013: 337); Congo (The Democratic Republic of the) (Hayman et al., 1966; Juste and Ibáñez, 1992: 363; Van Cakenberghe et al., 1999; Simmons, 2005: 521; Monadjem et al., 2010d: 539); Kenya (Simmons, 2005: 521); Tanzania (Simmons, 2005: 521). ROOST: McWilliam (1982: 176) found this bat to be roosting in caves. MIGRATION: In Kenya, McWilliam (1982: 175) found that most long distance movements were made by females, whereas males showed a larger site specificity. POPULATION: Structure and Density:- There are very few records of this species (Jacobs et al., 2008n; IUCN, 2009). Trend:- 2008: Unknown (Jacobs et al., 2008n; IUCN, 2009). ACTIVITY AND BEHAVIOUR: In Kenya, M. minor exhibits seasonal torpor between April and September, when the temperatures are lower (McWilliam, 1982: 184). MATING: McWilliam (1982: 176, 177) described a small "mating dome" within the Homa Lime cave in Kenya, which was almost exclusively occupied by mature males, which returned the following years. Immature males were excluded from the dome. During the mating period (May to July), the males produced a pungent odour (resembling rotting cocoa) in their urine (McWilliam, 1982: 181). PARASITES: Uchikawa (1985b: 50) described the mite Calcarmyobia minoris from Miniopterus minor minor from Similani Cave, 16 km S. of Mombasa, Coast Province, Kenya. Uchikawa (1985c: 110) also reported three female mites belonging to Pteracarus miniopteri Uchikawa, 1978 from bats from Kenya. Adam and Landau (1973a: 5) and Landau and Adam (1973: 6) report on the presence of a protozoan parasite of the genus Polychromophilus (Haemoproteidae) in M. minor minor from the Republic of Congo. About 50 % of the specimens they examined, were found to be infected. Gametocytes were found year-round. The bats were also infested by the nycteribiids Penicillidia fulvida Bigot, 1885 and Nycteribia schmidlii scotti Falcoz, 1923, of which the prior was a major carrier of Polychromophilus sporozoites. Garnham (1973: 237) reports the parasite as Polychromophilus melanipherus. Polychromophilus sp. was also reported by Rosskopf et al. (2018: 31) from Gabonese M. minor occidentalis, where both of the examined specimens were infected. Landau and Adam (1973: 6) also report on another prozotoan parasite: Hepatocystis, of which schizonts were found in the liver and lungs. Duval et al. (2012: 1559) and Landau et al. (2012: 142) indicate that the haemosporidan Polychromophilus (Polychromophilus) adami Landau et al., 1980 was described from M. minor minor specimens in the Republic of Congo (see Landau et al., 1980b: 27). Landau et al. (2012: 142) also mention this bat as the type host for Bioccala murinus (Dionisi, 1899). Lutz et al. (2016: 9) examined 13 M. minor specimens from East Africa and found five of them infected with Polychromophilus melanipherus Dionisi, 1899. Nycteribiidae: Nycteribia schmidlii Schiner, 1853 (Haeselbarth et al., 1966: 108). VIRUSES: In their country-wide survey of Kenyan bats, Waruhiu et al. (2017) found the following viruses in African Chiroptera Report 2020 Miniopterus minor: Astroviruses, Coronaviruses, Paramyxoviruses and Polyomaviruses. 623 Kenya, Madagascar, São Tomé and Principé, Tanzania. Coronaviridae - Coronaviruses SARS-CoV - Tong et al. (2009) tested 1 bat collected in Kenya during 2006 positive for the presence of coronavirus RNA in a fecal sample. 22.6 % (66 out of 292) of the Kenyan bats tested by Tao et al. (2017: 3) were positive for CoV. Paramyxoviridae: Paramyxovirus - Conrardy et al. (2014: 259) tested 16 bats from the Three Caves (Kenya) and found one of them to be positive. Mortlock et al. (2015: 1841) tested 151 Kenyan bats and found 14 to be positive. Nieto-Rabiela et al. (2019: Suppl.) mention Bat paramyxovirus and Miniopterus paramyxovirus to be present. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Central African Republic, Comoros, Congo, Congo (Democratic Republic of the), Figure 228. Distribution of Miniopterus minor Miniopterus mossambicus Monadjem, Goodman, Stanley and Appleton, 2013 *2013. Miniopterus mossambicus Monadjem, Goodman, Stanley and Appleton, Zootaxa, 3746 (1): 123, 129, figs 3, 6, 7. Publication date: 10 December 2013. Type locality: Mozambique: Nampula province: outskirts of Nampula town: Bamboo Inn [15.10306 S 39.21748 E, 420 m] [Goto Description]. Holotype: FMNH 213651: ad ♂, skull and alcoholic. Collected by: Steven M. Goodman and Ara Monadjem; collection date: 9 October 2010; original number: SMG-16875. Presented/Donated by: ?: Collector Unknown. Paratype: FMNH 213652: ad ♂, alcoholic (skull not removed). Collected by: Steven M. Goodman and Ara Monadjem; collection date: 9 October 2010; original number: SMG-16876. Presented/Donated by: ?: Collector Unknown. - Etymology: Refers to the country from which the species was collected (see Monadjem et al., 2013a: 129). (Current Combination) COMMON NAMES: Chinese: 莫桑比克长翼蝠. English: Mozambique long-fingered bat. German: MossambiqueLangflügelfledermaus. GENERAL DISTRIBUTION: Native: Kenya (López-Baucells et al., 2017: 20); Malawi; Mozambique; Zambia; Zimbabwe. ECHOLOCATION: Monadjem et al. (2013a: 139) mention a knee frequency of 55 kHz. PARASITES: BACTERIA: Dietrich et al. (2018a: 4) report the presence of Leptospira borgpetersenii. DIPTERA: Streblidae: Raymondia aspera Maa, 1968 from Mozambique (Shapiro et al., 2016: 254). VIRUSES: Astroviridae Two out of 21 bats tested positive in the study by Hoarau et al. (2018: 2). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Kenya, Malawi, Mozambique, Tanzania, Zambia, Zimbabwe. 624 ISSN 1990-6471 Figure 229. Distribution of Miniopterus mossambicus Miniopterus natalensis (A. Smith, 1833) *1833. Vespertilio Natalensis A. Smith, S. Afr. Quart. J., ser. 2, 1 (2): 59. Publication date: November 1833. Type locality: South Africa: KwaZulu-Natal: Durban [29 52 S 31 00 E] [Goto Description]. Cotype: BMNH 1846.6.2.19:. Shared cotype for Vespertilio natalensis and Vespertilio scotinus - this needs further investigation as it is unlikely that Sundeval examined material described by Andrew Smith. It is possible that this co-type should only be assigned to one of these species. Cotype: BMNH 1848.6.12.19:. Comments: Concerning the type locality, Meester et al. (1986: 46) remark: "Durban nominated by Roberts (1951: 73)"). 1840. Vespertilio d'asythrix Temminck, Monographies de mammalogie ou description de quelques genres de mammifères, dont les espèces ont été observées dans les différens musées de l'Europe. Tome Seconde, 2: 268. Publication date: 1840. Type locality: South Africa: "Interior of Caffraria" [Goto Description]. Syntype: BMNH 1849.8.16.25:. Syntype: RMNH MAM.25093:. - Comments: . 1846. Vespertilio scotinus Sundevall, Öfvers. kongl. Sv. Vet.-Akad. Förhandl., 3 (4): 119. Publication date: 1846. Type locality: South Africa: KwaZulu-Natal: Durban [29 52 S 31 00 E]. Cotype: BMNH 1846.6.2.19:. Shared cotype for Vespertilio natalensis and Vespertilio scotinus - this needs further investigation as it is unlikely that Sundeval examined material described by Andrew Smith. It is possible that this co-type should only be assigned to one of these species. - Comments: Type locality originally mentioned as "variis Caffrariæ locis allata"; Meester et al. (1986: 46) state: "Durban nominated by Roberts (1951: 74)". 1909. Miniopterus breyeri Jameson, Ann. Mag. nat. Hist., ser. 8, 4 (23): 471. Publication date: 1 November 1909. Type locality: South Africa: N Transvaal province: Waterberg district: Gat(s)koppies [Goto Description]. Holotype: BMNH 1909.7.2.6: imm ♀. Collected by: Henry Paul William Lyster Jameson; original number: 398. 1927. Miniopterus smitianus Thomas, Proc. zool. Soc. Lond., 373. Publication date: 12 July 1927. Type locality: Namibia: E Damaraland, 40 mi (64 km) W of Gobabis: Witvlei. Holotype: BMNH 1926.12.7.15:. 2014. Miniopterus 'natalensis': Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: Suppl.. Publication date: 18 March 2014. 2018. Miniopteris schriebersii: Adams and Kwiecinski, Diversity, 10 (3) 103: 7. Publication date: 18 September 2018. (Lapsus) ? Miniopterus africanus natalensis: (Name Combination) ? Miniopterus breyeri breyeri: ? Miniopterus natalensis dasythrix: (Name Combination) ? Miniopterus natalensis natalensis: (Name Combination) ? Miniopterus natalensis smitianus: (Name Combination) ? Miniopterus natalensis: (Name Combination, Current Combination) ? Miniopterus schreibersi dasythrix: African Chiroptera Report 2020 ? ? ? ? ? ? ? 625 Miniopterus schreibersi natalensis: (Name Combination) Miniopterus schreibersi: Miniopterus schreibersii natalensis A: Miniopterus schreibersii natalensis B: Miniopterus schreibersii natalensis: (Name Combination) Miniopterus schreibersii: Vesperugo scotinus: TAXONOMY: Figure 230. Miniopterus natalensis. Considered a synonym of schreibersi by Koopman (1993a: 231). O'Shea and Vaughan (1980), Koopman (1994), Peterson et al. (1995) and Horácek et al. (2006: 105) suggested that natalensis may be distinct from schreibersi. Considered a valid species by Simmons (2005: 521). Simmons (2005: 521) recognised arenarius Heller, 1912 as valid subspecies. Based on mitochondrial data, Demos et al. (2019c: "9") indicate that natalensis is sister to all of the other African and Madagascan forms, although this was not confirmed by the nuclear data. COMMON NAMES: Chinese: 纳 塔 尔 长 翼 蝠 . Czech: létavec natalský. English: Natal Long-fingered Bat, Natal Clinging Bat. French: Minioptère du Natal. German: Natal-Langflügelfledermaus, Südafrikanischen Sackfledermaus, KaffernSackfledermaus. SiSwati: Lilulwane. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: This species is present in the later half of the upper Pleistocene sequence and in the late Holocene at Border Cave, and probably at Nkupe during the middle and late Holocene (Avery, 1991: 6). Further Holocene material is reported by Avery and Avery (2011: 20) from Blinkklipkop, Limerock, and Zoovoorbij (Northern Cape province, South Africa), who also reported Pleistocene remains from Wonderwerk. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008aj; IUCN, 2009; Monadjem et al., 2017bg). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bg). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008aj; IUCN, 2009). 2004: NT ver 3.1 (2001) (Jacobs et al., 2004ai; IUCN, 2004). Regional South Africa:- 2016: LC ver 3.1 (2001) (MacEwan et al., 2016b). 2004: NT ver 3.1 (2001) [assessed as M. schreibersii] (Friedmann and Daly, 2004:263). Swaziland:- 2003: NT B2ab(iv); D2 ver 3.1 (2001) [assessed as M. schreibersii] (Monadjem et al., 2003: 184). MAJOR THREATS: In view of its wide range, there appear to be no major threats to this species. In parts of the species distribution it is locally threatened by habitat loss resulting from conversion of land to agricultural use, incidental poisoning with insecticides, and the disturbance of roosting and maternity caves by tourist activities (Jacobs et al., 2008aj; IUCN, 2009; Monadjem et al., 2017bg). CONSERVATION ACTIONS: Jacobs et al. (2008aj) [in IUCN (2009)] and Monadjem et al. (2017bg) report that there appear to be no conservation measures in place for this species. There is a need to identify and protect important roost sites (especially maternity caves). In view of the species wide range, it is presumably present in some protected areas. There is a need to better determine the range of this species when compared to that of Miniopterus schreibersii. GENERAL DISTRIBUTION: Miniopterus natalensis is a widely distributed species, recorded from southern and East Africa, with some records from Central Africa. It ranges south from Angola and southern Democratic 626 ISSN 1990-6471 Republic of the Congo, into Namibia, Botswana, South Africa, Lesotho, Mozambique, Malawi, Zimbabwe and Zambia. Because of frequent misidentification between this species and Miniopterus schreibersii, there is a need to carefully review the distribution of Miniopterus natalensis. Records from the Arabian Peninsula, Ethiopia, South Sudan and Uganda are currently assigned to Miniopterus arenarius (see Monadjem et al., 2020: 239; Ibáñez and Juste, 2019: 704). We also tentatively include the records from Kenya and Tanzania to this species. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 1,227,870 km2. In the RSA, its distribution is best predicted by geology (Babiker Salata, 2012: 50). Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 539; Taylor et al., 2018b: 62); Botswana (Smithers, 1971; Monadjem et al., 2010d: 539); Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 539); Kenya?; Lesotho (Lynch, 1994: 191 [as "M. schreibersii"]; Monadjem et al., 2010d: 539); Malawi (Happold et al., 1988; Ansell and Dowsett, 1988: 35; Monadjem et al., 2010d: 539); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 539; Monadjem et al., 2010c: 385); Namibia (Monadjem et al., 2010d: 340); South Africa (Simmons, 2005: 521, Seamark and Kearney, 2007: 4; Monadjem et al., 2010d: 540); Swaziland (Monadjem et al., 2010d: 540); Tanzania? (Stanley and Goodman, 2011: 47 as M. schreibersii); Zambia (Ansell, 1974; Ansell, 1978; Monadjem et al., 2010d: 540); Zimbabwe (Monadjem et al., 2010d: 540). Presence uncertain: Uganda, Kenya, Tanzania. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Schoeman and Jacobs (2002: 157) reported the following wing parameters for one "M. schreibersii" specimen from the Algeria Forestry Station (RSA): Fa: 46.6 mm, Wing area: 137.7 cm 2, Wing span: 30.9 cm, Wing loading: 7.8 N/m 2, and aspect ratio: 6.9. DENTAL FORMULA: I 2/3 C 1/1 P 2/3 M 3/3 = 36 (van der Merwe, 1985a: 73). van der Merwe (1985a: 73) also indicates that the deciduous tooth replacement starts in the lower jaw and is completed prior to that of the upper jaw. The replacement starts at an age of one and a half weeks. In general the replacement sequence is: i3, i1, c1, i2, p4, p3, and all teeth are replaced at week 5. In the upper jaw, the replacement starts two and a half weeks (p4, p3, i2, i3, c1) and is generally completed in week 6 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: Camacho et al. (2019: "5") describe the facial profile of M. natalensis as convex ("airorhynchy"), with an extreme upward orientation in its snout. FUNCTIONAL MORPHOLOGY: Moussy et al. (2012: 189), referring to MillerButterworth et al. (2003), indicate that isolation of different populations in South Africa has led to local adaptation in Miniopterus schreibersii natalensis, which is strongly associated with four biomes and results in differences in wing morphology and migratory behaviour. The animals from the northeastern population have a higher wing aspect ratio, which forms an advantage for long-distance flight, allowing for migrations over longer distances. Brown (1999: 170) found that "M. schreibersii" is able to thermoregulate in temperatures between 5 and 40°C. ECHOLOCATION: See Jacobs (1999) and Taylor (1999b). Schoeman and Waddington (2011: 291) mention a peak frequency of 51.1 ± 0.9 kHz and a duration of 6.8 ± 0.4 msec for specimens from Durban, South Africa. For Miniopterus natalensis specimens from Swaziland and Namibia, Taylor et al. (2013b: 17) recorded the following parameters for 21 calls: Fmax: 65.9 ± 5.76 (59.0 - 80.9) kHz, Fmin: 53.7 ± 0.58 (52.5 - 54.9) kHz, Fknee: 56.5 ± 0.31 (55.7 57.0) kHz, Fchar: 54.1 ± 0.78 (52.7 - 55.5) kHz, duration: 2.6 ± 0.48 (1.9 - 3.8) msec. Values for a further 11 specimens from Swaziland were reported by Monadjem et al. (2017c: 179): Fmin: 53.2 ± 0.87 (51.3 - 54.8) khz, Fknee: 57.0 ± 0.96 (55.5 - 58.5) kHz, Fc: 53.6 ± 0.83 (51.6, 55.0) kHz and duration: 2.8 ± 0.37(2.2 - 3.7) msec. They also indicated that the maximum detection distance for this species is 5 m. At Farm Welgevonden, South Africa, Taylor et al. (2013a: 556) report a Fknee value of 56 (56 - 57) kHz. Schoeman and Jacobs (2002: 157) reported a Fpeak of 53.6 kHz for one "M. schreibersii" from the Algeria Forestry Station (RSA). Linden et al. (2014: 40) reported the following parameters from the Soutpansberg area (RSA): Fmin: 52 - 55 kHz, African Chiroptera Report 2020 Fchar: 45 - 62 kHz, Fknee: 56 - 57 kHz, Slope: 235 622 OPS, duration: 2 - 4 msec. In the Mapungubwe National Park (RSA), Parker and Bernard (2018: 57) recorded 15 calls with Fchar: 49.91 ± 0.95 kHz, Fmax: 67.84 ± 4.75 kHz, Fmin: 49.39 ± 0.88 kHz, Fknee: 53.61 ± 2.27 kHz, duration: 4.96 ± 1.31 msec and 7.30 ± 2.65 calls/sec. Eisenring et al. (2016: SI 2) reported for 17 calls of bats from the Aberdares Range in Kenya the follow values: PF: 44.1 ± 3.5 (41.5 - 54.8), HF: 66.6 ± 17.7 (51.3 - 104.4), LF: 40.4 ± 1.5 (38.5 - 44.0), DT: 0.1 ± 0.0 (0.1 - 0.1), DF: 26.1 ± 17.5 (12.5 63.9), IPI: 1.0 ± 0.5 (0.5 - 2.5). 10 calls from the Okavango River Basin reported by Weier et al. (2020: Suppl.) had the following characteristics: Fmax: 58.77 ± 2.45 kHz, Fmin: 55.48 ± 1.10 kHz, Fknee: 57.07 ± 1.50 kHz, Fchar: 55.90 ± 1.29 kHz, slope: 13.28 ± 20.65 Sc, duration: 3.92 ± 1.48 msec. Luo et al. (2019a: Supp.) reported the following data (Hand released bats): Fpeak: 51.4 kHz and duration: 3.4 msec. MOLECULAR BIOLOGY: DNA - Teeling et al. (2017: 32) mention the assembly size for M. natalensis to be 1.80 Gb. Karyotype - M. schreibersii from Namibia and South Africa now assigned to M. natalensis, were karyotyped by Rautenbach et al. (1993) who described the karyotype as having a diploid number of 2n = 46, aFN = 50, BA = 6, X = SM, Y = A. Banerjee et al. (2017: 10) detected potential c-Rel binding sites in M. natalensis. This protein has a potential role in the DNA repair pathway as a regulator of inflammatory gene expression. HABITAT: Sirami et al. (2013: 34) recorded in the Western Cape Province, South Africa that M. natalensis activity was not significantly, nor positively influenced by size of wetland, while habitats 100 m surrounding wetlands were also significantly and positively influenced by the water body. M. natalensis prefered trees and orchards surrounding wetlands (Sirami et al., 2013: 35). Voigt et al. (2013: 751) investigated stable isotope ratios of carbon, nitrogen and hydrogen in the fur keratin of M. natalensis on the slopes of Mount Kilimandjaro and found that this species migrates seasonally between low and high elevations. They found that M. natalensis captured before and after the cold period at around 1,800 m above sea 627 level originated from around 1,400 m a.s.l. or lower, which they suggest might be the result of a search for colder hibernacula at higher elevations. HABITS: van der Merwe (1973b: 380) indicates that in the Transvaal M. natalensis forms hibernating (winter) and maternity (summer) colonies, and he reports on maternity caves housing 50,000 to 100,000 animals. The bats usually form closely packed clusters, hanging from the roofs of caves, although some might be hanging against the walls. Hibernation started at the beginning of May and lasted until the end of July. At De Hoop Nature Reserve (Western Cape Province, South Africa), Thomas and Jacobs (2013: 124) found that M. natalensis emerged significantly later during early summer than in late summer. Norton and van der Merwe (1978: 219) already found that juveniles returned later in the night than adults, probably because they needed more food as their fat reserves would be insufficient to get them over the winter. ROOST: van der Merwe (1973b: 380) reports "M. schreibersi natalensis" to occur in colonies of hundreds or thousands of animals occupying a wide range of caves. At Peppercorn's cave and at Sandspruit Cave No. 1 clusters of 50,000 and 100,000 individuals were reported. van der Merwe (1973a: 133) found that the bats preferred caves with lower temperature for hibernation. Norton and van der Merwe (1978: 220) also found that temperatures in the Long One Cave were probably too high and variable for prolonged hibernation, which resulted in fluctuating numbers of bats. By the end of the winter, however, temperatures decreased and the proportion of M. natalensis bats staying in the cave increased. Bernard and Bester (1988: 921) describe the annual movements of "M. schreibersi" between three roosts in the Eastern Cape region (RSA). The maternity roost was warmer than the hibernation site. One of the sites was used as maternity and hibernaculum, but as these functions were at different times of the year, the temperatures were also different. The higher temperature at the maternity roost might increase prenatal and postnatal development rates. Brown (1999: 171) reported the mean temperature in tunnel roosts is on average 19.5°C in summer (max: 22°C) and 14°C in winter. 628 ISSN 1990-6471 DIET: At the Umbilo River (KwaZulu Natal, RSA), Naidoo et al. (2011: 26) found the following percentage volume diet composition: Lepidoptera (35.7 ± 10.3), Coleoptera (11 ± 9.1), Hemiptera (28.3 ± 12.9), and Trichoptera (25.0 ± 13.9). No data were provided for the winter. Schoeman and Jacobs (2002: 157) reported the following volume percentages found in 20 faecal pellets from one "M. schreibersii" from the Algeria Forestry Station (RSA): Diptera (49), Hemiptera (33), Lepidoptera (6), Coleoptera (6), and Trichoptera (6). For 21 "M. schreibersii" specimens from Sudwala (RSA), Jacobs (2000: 201) reported the following prey proportions: Lepidoptera (23.1 ± 31.2), Coleoptera (4.5 ± 16.7), Isoptera (70.8 ± 32.2), Diptera (0.6 ± 2.3), Hemiptera (0.6 ± 2.3), and Mantodea (0.4 ± 0.3). PREDATORS: Avery et al. (2005: 1054) found this species (reported as M. schreibersii) to be present in pellets from Tyto alba in South Africa. van der Merwe (1980a: 15) already reported the barn owl as a predator, but indicated that these bats do not rank high on its diet, although there was an increase when other terrestrial and subterrestrial preys are descreasing in availability. POPULATION: Structure and Density:- This species can be found in colonies of more than 2,500 animals (Jacobs et al., 2008aj; IUCN, 2009; Monadjem et al., 2017bg). Mason et al. (2010: 165) observed that the number of bats in the De Hoop maternity chamber increased from a few hundred to several thousand from 18th August to the 25 September 2006. Trend:- 2016: Unknown (Monadjem et al., 2017bg). 2008: Unknown (Jacobs et al., 2008aj; IUCN, 2009). LIFESPAN: Based on mark-recapture techniques, van der Merwe (1989b: 87) determined the maximum life expectancy for this species in the wild to be 13 years. REPRODUCTION AND ONTOGENY: Delayed implantation, insemination, ovulation and fertilization occur in the autumn (April to May in South Africa, see Bernard et al. (1991: 31), as M. schreibersii). Embryogenesis proceeds to the blastocyst stage and development is then retarded until arousal in spring, when the blastocyst implants and embryogenisis resumes (van der Merwe, 1980b). van der Merwe (1982b: 319) also mentions that implantation is centric and superficial in the right uterine horn, as was already noticed by Ginther (1967: 580) and van der Merwe (1996: 154), who also indicated [referring to Matthews (1941)], that the implantation occurs in the uterine horn opposite to the ovary containing the corpus luteum. van der Merwe (1979a: 17) and Bernard (1980a: 55) indicated that the delayed implantation period for "M. schreibersi natalensis" lasted for four months (120 days), which is as long as the active fetal growth period, giving a gestation period of 240 days. van der Merwe (1981: 172) presented the following formula to calculate the embryo's age in days (t) based on its body mass (W): t = W 1/3/0.0145 + 144. Bernard and Davison (1996: 218) investigated the relation between calcium availability (from the poor source that insects are) and the reproduction of "M. schreibersii" and found that seasonal changes in insect abundance could not explain the delayed implantation, but that lactating females nevertheless had the lowest bone calcium concentrations. This is probably the reason why parturition occurs in seasons with maximum insect availabitiy. Mason et al. (2010: 165) found at De Hoop that in mid August 2006 the bats sampled were either not pregnant or at a very early stage of pregnancy and no embryos were present. However, on the 8th September the first bat sampled was pregnant and carried an embryo at the limb bud stage (CS13), this is found approximately the same time of year as those of the KwaZulu Natal and Eastern Cape populations. No twins were observed and pregnant bats collected on the same day never had embryos from only one stage of development (Mason et al., 2010). Mason et al. (2010: 168) found that the period of early to mid-embryonic development (CS11 to CS21) occurred over a period of about two weeks in both years sampled, however this period appeared to occur about two weeks later in 2008 than in 2006, this delay in pregnancy may be due to environmental factors. Also the embryonic stages were not the same among individuals sampled on the same day, indicating that although reproductive events may be synchronous (within the period of a month) in this species, exact developmental events were not highly synchronized between individuals. At the Gatkop cave (RSA), Dietrich et al. (2018b: 4) were only able to trap non-scrotal males in September 2015. In November 2015, 90 % of the females they trapped had given birth very recently, and in January 2016, large amounts of juveniles were present, together with post-lactating females. African Chiroptera Report 2020 The mass of pregant females varied greatly and was not correlated to the developmental stage of the embryos that they carried (Mason et al., 2010: 165). Pretorius et al. (2019: 319), however, found that lactating females with sclerotized nipples were 6 % heavier than non-reproductive females and pregnant females were 25 % heavier than nonreproductive females, a difference - which at least in part - was attributed to foetus mass. Over three successive years (2015 - 2017), the proportion of active breeding females was found to be 31 %, 33 % and 70 %. Mason et al. (2015) studied the embryological development of the fore- and hindlimbs, and found that that the transcription factor Meis2 has a significantly higher expression in bat forelimb autopods compared to hindlimbs. Eckalbar et al. (2016: 528) found over 7,000 genes and noncoding RNAs (incl. Tbx5-as1 and Hottip), that were differentially expressed between forelimb and hindlimb, and across different stages of development. MATING: van der Merwe (1973b: 384) reported that the period of mating was still uncertain, although he assumed that it occurred at the wintering caves prior to hibernation. PARASITES: BACTERIA: Dietrich et al. (2016b: 3) tested 17 bats from Irene (RSA) and found two testing positive for Bartonella and one for Rickettsia. Five out of 8 bats from Vanderkloof Dam (RSA) also tested positive for Bartonella. Dietrich et al. (2015a: 3) reported the presence of the Enterobacteria Hafnia and Nitrococcus. They also found Leptospira in the urine sample, and Bartonella in urine and saliva samples. Dietrich et al. (2018a: 4;2018b: 4) reported the presence of bacteria related to Leptospira borgpetersenii and a second group closely related to L. interrogans and L. kirschneri. Clément et al. (2019: 5) reported on 97 southern African M. natalensis specimens of which 12 were infected by either Trypanosoma sp. 1_AF (2), Trypanosoma sp. 2_A (2), Trypanosoma sp. 2_B (3), Trypanosoma dionisii_AF1-AF4 (4) and Trypanosoma cf. livingstonei_likeH_G1 (1). Gastro-intestinal Helminthes - See Ortlepp (1932) and Junker et al. (2008a). NEMATODA: Genov et al. (1992) reported Molinostrongylus omatus (Mönnig, 1927) from M. natalensis from 629 South Africa. Ramasindrazana et al. (2016: 2) indicate that Litomosa chiropterorum Ortlepp, 1932 (Onchocercidae) was redescribed from a M. natalensis population from South Africa. This Nematode was already reported from a bat from Katanga (DRC) by Durette-Desset and Chabaud (1975: 304). ACARI: Myobiidae: Uchikawa (1985a: 19) also reported that the type specimens of Miniopterus breyeri, M. dasythrix, and M. scotinus harboured the "atypical" form of Calcarmyobia congoensis Uchikawa, 1982, and he also (Uchikawa, 1985a: 22) mentioned Calcarmyobia kenyaensis Uchikawa, 1982 on "Miniopterus schreibersi" from Belgian Congo. Uchikawa (1985c: 110) furthermore reported on the presence of Pteracarus miniopteri Uchikawa, 1978 on bats from South Africa, Zambia, Namibia (as M. schreibersi natalensis or M. smitianus). Trombiculidae: Stekolnikov (2018a: 147) mentioned Microtrombicula armata VercammenGrandjean, 1965 from a South African "M. schreibersii". DIPTERA Streblidae: Ascodipteron africanum Jobling, 1939 from Kapretwa and Mt. Kenya, Kenya, other records referred to this species are misidentification (Haeselbarth et al., 1966: 106). Ascodipteron theodori Maa 1965 from Rooiberg, South Africa (Haeselbarth et al., 1966: 106). Raymondi waterstoni Jobling 1931 which may be accidental (Haeselbarth et al., 1966: 104) (see also Shapiro et al., 2016: 256). Nycteribiidae: Nycteribia capensis Karaman, 1939 from the Cape and Pietermaritzburg, South Africa (Haeselbarth et al., 1966: 108). Nycteribia latiterga Theodor, 1957 from Mt. Menengai, Kenya (Haeselbarth et al., 1966: 108). Nycteribia schmidlii Schiner, 1853 (Zumpt, 1950: 97; Haeselbarth et al., 1966: 108; Blackwell, 1980: 146 - infected by the fungus Arthrorhynchus eucampsipodae Thaxter, 1901 [see also Haelewaters et al., 2018: 794; Szentiványi et al., 2019: Suppl.]). Penicillidia fulvida (Bigot, 1885) (Haeselbarth et al., 1966: 114; Blackwell (1980: 144 - infected by the fungus Arthrorhynchus nycteribiae (Peyritsch) Thaxter [see also Haelewaters et al., 2018: 794; Szentiványi et al., 2019: Suppl.]). SIPHONAPTERA: Ischnopsyllidae: Oxyparius isomalus (Waterston, 1915) known from several localities in southern Africa (Transvaal, Natal of South Africa, Botswana and Namibia) (Haeselbarth et al., 1966: 188). 630 ISSN 1990-6471 ACARI: Argasidae: Argas vespertilionis (Latreille, 1802) from a "M. schreibersi" from Pretoria (see Howard, 1908: 168). Dermanyssidae: Zumpt and Till (1954: 49) report Steatonyssus natalensis Zumpt and Patterson, 1951 from bats from Natal. Zumpt (1950 92) found a new mite of the genus Bdellonyssus. Macronyssidae: Zumpt (1950: 89) described Hirstesia transvaalensis from bats from the Sterkfontein cave (RSA). Myobiidae: Myobia miniopterus Womersley, 1941 (see Zumpt, 1950: 92). Spinturnicidae: Spinturnix semilunaris De Meillon and Lavoipierre, 1944 (see Zumpt, 1950: 92). VIRUSES: Bunyaviridae Phlebovirus Rift Valley Fever Virus (RVFV) - Oelofsen and Van der Ryst (1999) tested [as M. schreibersi] 49 individuals from one locality in the Free State Province, South Africa for RVFV antigen using an enzyme linked immunosorbet assay (ELISA), none tested positive. This virus does not group with the known paramyxoviruses and remains an unclassified paramyxovirus. Rhabdoviridae Lyssavirus - Rabies related viruses Duvenhage - van der Merwe (1982a) implicated Miniopterus schreibersii, as an isolate of the virus but the bat was never positively identified. This record was also referred to by Hayman et al. (2011a: 88), who indicated that the bas was possibly a M. natalensis. Rabies - Oelofsen and Smith (1993) tested [as M. schreibersi] 137 individual brains, from 5 localities by means of the "Rapid rabies enzyme immunodiagnosis" (RREID) test (Diagnostic Pasteur), none were tested positive. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Botswana, Congo (Democratic Republic of the), Eswatini, Lesotho, Malawi, Mozambique, Namibia, South Africa, Zambia, Zimbabwe. Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested [as M. schreibersi] between 1986 and 1999, for antibody to SARS-CoV in sera in one individual from Limpopo Province, South Africa, none were tested positive (0/1). Tong et al. (2009) tested 1 bat collected in Kenya during 2006 positive for the presence of coronavirus RNA in a fecal sample. Nine out of 53 (17 %) Kenyan bats tested by Tao et al. (2017: Suppl.) was positive for CoV. One out of 14 specimens tested by Geldenhuys et al. (2013: 517) from Irene Caves, Gauteng, RSA tested positive for CoV. Paramyxoviridae Conrardy et al. (2014) detected a paramyxovirus sequence in one of eight bats sampled in Kenya. Figure 231. Distribution of Miniopterus natalensis Miniopterus newtoni Bocage, 1889 *1889. Miniopterus Newtoni Bocage, J. Sci. mat. phys. nat., ser. 2, 1 (3): 198. Publication date: December 1889. Type locality: São Tomé and Principé: São Tomé Island [Goto Description]. Neotype: EBD MAM 17350: ad ♂, skull and alcoholic. Collected by: Javier Juste and Carlos J. Ibáñez Uargui; collection date: 4 April 1988. São Tomé Island, Santa Catarina, in a cave near the seashore (00 16 N 06 29 E), designated as neotype by Juste and Ibáñez (1992: 361). (Current Combination) ? Miniopterus newtoni: (Current Spelling) TAXONOMY: Recognised as a synonym (Koopman, 1993a: 231) or a subspecies (Simmons, 2005: 521) of M. minor Peters, 1867. Removed from minor by Juste [B.] et al. (2007: 33), based on mtDNA analyses. Juste and Ibáñez (1992: 361) designated a neotype for newtoni Bocage, 1889. African Chiroptera Report 2020 COMMON NAMES: Czech: létavec tomášský. English: Sao Thome Long-fingered bat. French: Minioptère de São Tomé. German: Sao Thome Langflügelfledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, natural history, threats and conservation status (Juste, 2008b; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Juste, 2008b; IUCN, 2009). 631 Native: São Tomé and Principe (Koopman, 1993a: 231 [as minor]; Simmons, 2005: 521 [as minor]; Juste [B.] et al., 2007; Rainho et al., 2010a: 33). ECHOLOCATION: Rainho et al. (2010a: 21) reports the calls of 14 individuals from São Tomé. POPULATION: Structure and Density:- It is a common species (Juste B. and Ibáñez, 1994b). Trend:- 2008: Unknown (Juste, 2008b; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: São Tomé and Principé. Regional None known. MAJOR THREATS: The threats to this species are not well known (Juste, 2008b; IUCN, 2009). CONSERVATION ACTIONS: Juste (2008b) [in IUCN (2009)] reports that it is not known if the species is present in any protected areas. Further studies are needed into the natural history and threats to this species. GENERAL DISTRIBUTION: Miniopterus newtoni is endemic to the island of São Tomé (São Tomé and Príncipe) where it is widespread (Juste [B.] et al., 2007). Figure 232. Distribution of Miniopterus newtoni Miniopterus nimbae Monadjem, Shapiro, Richards, Karabulut, Crawley, Nielsen, Hansen, Bohmann and Mourier, 2020 *2020. Miniopterus nimbae Monadjem, Shapiro, Richards, Karabulut, Crawley, Nielsen, Hansen, Bohmann and Mourier, Acta Chiropt., 21 (2): 239, 245, figs 6, 7A, 8, 9A. Type locality: Liberia: Nimba county: Mount Gangra: 10 km W Mount Nimba [07 33 16 N 08 37 44 W, 720 m] [Goto Description]. Holotype: DNSM 12621: ad ♂, skull and alcoholic. Collected by: Dr. Ara Monadjem; collection date: 18 December 2010; original number: AM2010_12_18_1. Presented/Donated by: Dr. Ara Monadjem. Paratype: DNSM 12614: ad ♀. Collected by: Dr. Ara Monadjem; collection date: 17 December 2010. Presented/Donated by: Dr. Ara Monadjem. - Etymology: Referring to the type locality on Mount Nimba. (Current Combination) COMMON NAMES: English: Nimba long-fingered bat CONSERVATION STATUS: Monadjem et al. (2020: 252) suggest this species might be assigned to the IUCN category Vulnerable or even Endangered. GENERAL DISTRIBUTION: Only known from Liberia (Mount Nimba and surrounding uplands, Wonegizi mountains) and Guinea (Mount Béro) (see Monadjem et al., 2020: 247). 632 ISSN 1990-6471 ECHOLOCATION: Monadjem et al. (2020: 248) found that the knee frequency of the echolocation calls from four handreleased M. nimbae specimens ranged between 47.2 and 48.9 kHz (average 48.4 kHz). REPRODUCTION AND ONTOGENY: Monadjem et al. (2020: 248) found that five out of 13 females captured between late December and the end of March were pregnant, and three out of 10 males had scrotal testes. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Liberia. Figure 233. Distribution of Miniopterus nimbae Miniopterus petersoni Goodman, Bradman, Maminirina, Ryan, Christidis & Appleton, 2008 2007. Miniopterus cf. sororculus: Goodman, Ryan, Maminirina, Fahr, Christidis and Appleton, J. Mamm., 88 (5): 1229. 2008. Miniopterus cf. petersoni: Goodman, Bradman, Maminirina, Ryan, Christidis and Appleton, Mamm. biol., 73: 202. *2008. Miniopterus petersoni Goodman, Bradman, Maminirina, Ryan, Christidis and Appleton, Mamm. biol., 73: 199, 200, 201, figs 2 - 5. Type locality: Madagascar: Province de Toliara: Cascade de Manantantely: Tolagnaro, 5.2km NW [2459.343S 4655.370E, 65m]. Holotype: FMNH 194136: ad ♂. Collected by: Steven M. Goodman; collection date: 20 February 2007; original number: SMG 15833. Paratype: USNM 577096:. Paratype: USNM 577097:. Paratype: USNM 577098: Collected by: G. Kenneth Creighton and E. Raholimavo; collection date: 19-30 October 1990. Presented/Donated by: ?: Collector Unknown. Paratype: USNM 577099:. Paratype: USNM 577100:. Paratype: USNM 577101:. - Etymology: The name petersoni is a patronym in honour of Randolph L. Peterson of the Royal Ontario Museum, Toronto, Canada, who made pioneering contributions to the study of Malagasy bats, based on a 1967 collecting trip to the island, as well as other fields in mammalogy (Eger and Mitchell, 1990; Goodman et al., 2008a). (Current Combination) TAXONOMY: See Goodman et al. (2008a). COMMON NAMES: Czech: létavec Petersonův. English: Peterson's Long-fingered Bat. French: Minioptère de Peterson. German: Petersons Langflügelfledermaus. SIMILAR SPECIES: See Goodman et al. (2008a). CONSERVATION STATUS: Global Justification This species is listed as Data Deficient (DD ver 3.1 (2001)) because although it appears to have a restricted range in the southeast of Madagascar and is associated with forest, there is taxonomic uncertainty about specimens from the north of Madagascar (which if proved valid would greatly extend the known range). There is also limited information on the population status and threats to this species (Jenkins and Rakotoarivelo, 2008b; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Jenkins Rakotoarivelo, 2008b; IUCN, 2009). and Regional None known. MAJOR THREATS: Although robust data are lacking and there are still some taxonomic uncertainties of specimens from the north of Madagascar, the information on M. petersoni from the southeast suggests that this African Chiroptera Report 2020 species is closely associated with natural forest formations and is therefore threatened by the loss or degradation of these habitats (Goodman et al., 2008a). Threats to roosting colonies have not yet been documented (Jenkins and Rakotoarivelo, 2008b; IUCN, 2009). CONSERVATION ACTIONS: Jenkins and Rakotoarivelo (2008b) [in IUCN (2009)] report that none of the known collecting localities for verified specimens of this species are from within protected areas (Goodman et al., 2008a). The taxonomic status of specimens from north Madagascar requires further study. GENERAL DISTRIBUTION: Miniopterus petersoni is endemic to the island of Madagascar and appears to be restricted to lowland (< 550 m elevation) habitats in the southeast (Goodman et al., 2008a). Schoeman et al. (2014: 28, 31) found the most suitable area for this species to be located in the southeastern part of the island, but reaching along the coast to the northeast, where the suitability seems to increase again (eastern humid forest or lowland forest region - humid). The taxonomic status of specimens from the north of Madagascar remain unresolved (Goodman et al., 2008a). Native: Madagascar (Goodman et al., 2008a). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: See Goodman et al. (2008a). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Goodman et al. (2008a). 633 MOLECULAR BIOLOGY: DNA - See Goodman et al. (2008a). Karyotype - Unknown. Protein / allozyme - Unknown. HABITAT: See Goodman et al., 2008a). HABITS: See Goodman et al. (2008a). ROOST: See Goodman et al. (2008a). MIGRATION: May be involved in seasonal migrations between cave roost sites (Goodman et al., 2008a). POPULATION: Structure and Density:- This species was only formally described in 2008 and there is no information on its population (Goodman et al., 2008a). Trend:2008: Unknown (Jenkins Rakotoarivelo, 2008b; IUCN, 2009). and PARASITES: Tortosa et al. (2013: 4) and Ramasindrazana et al. (2017: Suppl.) reported the presence of the Nycteribiid bat flies Nycteribia stylidiopsis and Penicillidia leptothrinax. The latter is also reported by Wilkinson et al. (2016), and was found to be carrying Rickettsia bacteria Szentiványi et al., 2019: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. ECHOLOCATION: Low-duty cycle, frequency modulated/quasi constant (FM/QCF) echolocation call, with a peak frequency of 53.2 kHz (52.0 - 53.9 kHz) and a duration of 2.9 ms (2.5 - 3.3 ms) (Goodman et al., 2011: 9). Ramasindrazana et al. (2011: 294) recorded 6 calls from 1 Miniopterus petersoni specimen, and found the following echolocation call values: Frequency of maximum energy: 53.2 ± 0.75 (52.0 - 53.9) kHz, Maximum frequency: 106.5 ± 6.66 (95.0 - 115.0) kHz, Minimum frequency: 49.0 ± 0.63 (48.0 - 50.0) kHz, Call duration: 2.9 ± 0.32 (2.5 - 3.3) ms, Inter pulse interval: 71.0 ± 4.63 (63.8 - 76.7) ms. Figure 234. Distribution of Miniopterus petersoni 634 ISSN 1990-6471 Miniopterus schreibersii (Kuhl, 1817) *1817. Vespertilio schreibersii Kuhl, Die Deutschen Fledermäuse, Hanau, 14. Type locality: Romania: S Ban(n)at Mnts, near Coronini, left bank Danube: Kolumbács Cave [=Kulmbazer cave] [ca. 44 37 N 21 40 E]. - Comments: Turni and Kock (2008) mention that the type and three "doublets" should have been present in the NMW, but are no longer found. - Etymology: In honour of Karl Franz Anton, Ritter von Schreibers (1775 - 1852), then director of the Vienna Museum (Flannery, 1995a: 378; 1995b: 452; Kozhurina, 2002: 17). 1827. Vespertilio Screbersii: Lesson, Manuel de Mammalogie, 89. (Lapsus) 1855. V[espertilio] Schreibersi: Giebel, Die Säugethiere, 950. (Name Combination) 1953. Miniopterus schrebersii: Harrison, Durban Mus. Novit., 4 (5): 65. Publication date: 30 June 1953. (Lapsus) 1995. M[iniopterus] screibersi: Pavlinov, Borissenko, Kruskop and Jahonton, Arch. Zool. Mus., Moscow State Univ., 133: 119. (Lapsus) 2013. Miniopterus schreibersiias: Shi, Sci. China Life Sci., 56 (8): 679. (Lapsus) ? Miniopterus schreibersi schreibersi: (Name Combination) ? Miniopterus schreibersii dasythrix: (Name Combination) ? Miniopterus schreibersii schreibersii: (Name Combination) ? Miniopterus schreibersii villiersi: (Name Combination) ? Miniopterus schreibersii: (Name Combination, Current Combination) TAXONOMY: Molecular systematics and biogeography are discussed by Appleton et al. (2004: 436 - 437), who indicate that based on their data M. schreibersii would be restricted to Europe and North Africa; the Ethiopian representatives would belong to a second species and the OrientalAustralian a third (see also Stadelmann et al., 2004b: 188). Tian et al. (2004: 303) confirm the views presented by Appleton et al. (2004), and suggest M. schreibersii for the European specimens, M. fuliginosus for the Asian representatives and M. oceanensis for the Australian ones (which was already suggested by Maeda (1982), although he separated the specimens from Hainan from the other Asian representatives, which is no longer supported by Appleton et al. (2004). Reviewed by Crucitti (1976) and Hill (1983). If Miniopterus schreibersii is restricted to northern Africa, then Miniopterus inflatus villiersi Aellen, 1956 should possibly be assigned to Miniopterus natalensis, but as already indicated by Fahr et al. (2006a: 74) and Monadjem et al. (2016y: 371), villiersi represents a distinct species, restricted to the Upper Guinea forest zone. COMMON NAMES: Afrikaans: Schreibers se grotvlermuis, Schreibergrotvlermuis. Albanian: Miniopteri i Schreibers-it, Lakuriq nate I Schreiber-it. Arabian: Khaffash. Armenian: Սովորական երկարաթև չղջիկ. Azerbaijani: Adi uzunqanad yarasa. Basque: Schreibers saguzar, Schreiber saguzarra. Belarusian: Даўгакрыл звычайны. Bosnian: Šrajberov šišmiš, Dugokrili slijepi miš. Breton: Miniopter Schreibers. Bulgarian: Пещерен дългокрил. Castilian (Spain): Murciélago de cueva. Catalan (Spain): Ratpenat de cova, Rat penat de cova, Ratapinyada de cova. Croatian: Dugokrili pršnjak, Šrajberov šišmiš. Czech: Létavec stěhovavý, nedopír širokouchý. Danish: Schreibers' langfingerflagermus. Dutch: Langvleugelvleermuis, Schreibers' vleermuis, Grote Schreiber's vleermuis. English: Schreiber's Bent-winged Bat, Schreiber's Bat, Long-wing Bat, Schreibers's Long-fingered Bat, Common Bentwinged Bat, Greater Bent-winged Bat, Large Bentwing Bat, Bent-winged bat, Common Longfingered Bat, large bent-winged bat. Estonian: Euroopa pikktiib. Finnish: Pitkäsiipiyökkö. French: Minioptère de Schreibers, Minioptère à longues ailes, Chauve-souris de Schreiber, Minioptère. Frisian: Schreibers'langwjukflearmûs. Galician (Spain): Morcego das covas. Georgian: ჩვეულებრივი ფრთაგრძელი. German: Langflügelfledermaus, Gemeine Langflügelfledermaus, Schreibers Langflügelfledermaus, Europäische Sackfledermaus, Schreibers Fledermaus. Greek: Πτερυγονυχτερίδα. Hebrew: ‫כנפן‬, Kanfan. Hungarian: Európai hosszúszárnyú-denevér, Törpe denevér, Hosszúszárnyú denevér, Nycteris i macropterys. Indonesian: Kelelawar mini schreiber, Tomosu Biasa, Kelelawar mini biasa. Irish Gaelic: Ialtóg mhéarfhada Schreiber. Italian: Miniottero, Miniòttero di Schrèibers, Miniòptero di Schrèibers. Latvian: Šreibersa siksparnis. Lithuanian: Šreiberio ilgapirštis. Luxembourgish: Laangflillekefliedermaus. Macedonian: Долгокрилест лилјак, Dolgokrilest Liljak. Maltese: Farfett il-Lejl ta' Xrajber. Montenegrin: African Chiroptera Report 2020 Dugorili prstenjak. Nepali: Bange Chamero. Norwegian: Middelhavslangvinge. Polish: Podkasaniec Schreibersa. Portuguese: Morcego-de-peluche, Morcego de schreiber. Rhaeto-Romance: Miniopter da l'ala lunga, Minopter ad ala lunga. Romanian: Liliacul cu aripi lungi, Liliac-cu-aripi-lungi. Russian: Длиннокрыл обыкновенный, Обыкновенный длиннокрыл [= Obyknovennyj dlinnokryl]. Serbian: Европски дугокрилаш. Serbian Latin: Evropski dugokrilaš. Scottish Gaelic: Ialtag mheur-fhada Schreiber. Sinhalese: Schreibersge dik-angeli wawula. Slovak: Lietavec stahovavý. Slovenian: Dolgokrili netopir. Swedish: Europeisk vikvinge. Tamil: Sraipars-ṉ Nīṇṭa Viraluḷḷa Veḷavāl, ஸ்றரப ர்ஸ்ன் நீ ண்ட விரலுள் ள வெளொல் . Turkish: Uzunkanatlı Yarasa. Ukrainian: Довгокрил звичайний. Welsh: Ystlum adain-gam Schreiber. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Gunnell et al. (2015a: 22) report skeletal remains from a Pleistocene deposit at Olduvai Gorge (Tanzania), which they assigned to Miniopterus cf. M. schreibersi. CONSERVATION STATUS: Global Justification Listed as Near Threatened NT ver 3.1 (2001). Significant population declines and range contractions have been recorded in a number of range states and although it is stable in the Balkans and Turkey, overall the rate of population decline may approach 30 % (almost qualifies as VU under A2a) (Hutson et al., 2008c; IUCN, 2009). Assessment History Global 2008: NT ver 3.1 (2001) (Hutson et al., 2008c; IUCN, 2009). 2004: LC ver 3.1 (2001) [included natalensis] (Schlitter, 2004i; IUCN, 2004). 1996: LR/nt [included M. natalensis] (Baillie and Groombridge, 1996). Regional Mediterranean:- 2008: NT ver 3.1 (2001) (IUCN, 2009). MAJOR THREATS: The main threat in Africa is disturbance in roosts. In Europe, the disturbance and loss of underground habitats and pesticide use may threaten this species. In the Caucasus, disturbance caused by tourism in caves is a problem (K. Tsytsulina pers. comm., 2005). The cause of recent mass mortality events is unknown. In 2002 mass mortalities of this species were reported for populations in France, Spain and Portugal. There are also historical records for such mortalities in Italy, Australia and a 635 possible incidence in Iran. A meeting was held at the 9th European Bat Research Symposium to discuss these incidents. Veterinary investigations in Spain did not identify any disease as the cause of the die offs, and there is increasing belief that the die offs are caused by bad weather in late winter/early spring (Hutson et al., 2008c; IUCN, 2009). The species's small range, cave/tree roosting and aerial hawking feeding style are believed to be the most important risk factors in view of climatic change (Sherwin et al., 2012: 174). CONSERVATION ACTIONS: Hutson et al. (2008c) [in IUCN (2009)] report that in Europe, it is protected by national legislation in most range states. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention in parts of the range where these apply. It is included in Annex II (and IV) of the EU Habitats and Species Directive, and hence requires special measures for conservation including designation of Special Areas for Conservation. There is some habitat protection through Natura 2000, and some roosts are already protected by national legislation. There have been a number of LIFEfunded projects for this species in Spain, Italy, Romania and Germany. The species is found in many protected areas throughout its range. Care is required when fencing caves to minimise mortality. Further research is required into the causes of the recent mass mortality events. GENERAL DISTRIBUTION: Miniopterus schreibersii occurs from southwestern Europe and North and West Africa through Anatolia and the Middle East to the Caucasus. In Africa it is known from records in North Africa (Morocco, Algeria, Tunisia, Libya), and West Africa (Guinea, Sierra Leone, Liberia, Nigeria, Cameroon). It is patchily distributed over its range in some huge and vulnerable colonies. It typically occurs at altitudes of up to 1,400 m asl (commuting up to 2,600 m asl). Native: Afghanistan; Albania; Algeria; Armenia; Azerbaijan; Bosnia and Herzegovina; Bulgaria; Cameroon; Croatia; Cyprus; France [Corse]; Georgia; Gibraltar; Greece [East Aegean Isands; Kriti]; Holy See [Vatican City State]; Hungary; Israel; Italy [Sardegna, Sicilia]; Jordan; Lebanon; Liberia; Macedonia, the former Yugoslav Republic of; Malta; Monaco; Montenegro; Morocco (Benda et al., 2010a: 158; El Ibrahimi and Rguibi Idrissi, 2015: 364); Nigeria; Palestinian Territory, 636 ISSN 1990-6471 Occupied; Portugal; Romania; Russian Federation; San Marino; Serbia; Sierra Leone; Slovakia; Slovenia; Spain [Baleares]; Switzerland; Syrian Arab Republic; Tunisia (Dalhoumi et al., 2014: 53; 2016b: 867; 2019b: 26); Turkey. Possibly extinct: Austria. Regionally extinct: Germany; Ukraine. Presence uncertain: Libyan Arab Jamahiriya. DETAILED MORPHOLOGY: Bhide (1979: 1) described the anatomy, histology and histochemistry of this bat's stomach In Korea, three types of lingual papillae were on the tongue by Park and Lee (2009: 267): filiform, fungiform, and circumvallate. Cebesoy and Karakas (2013: 382) report on the muscle fiber types in primary flight muscles of specimens from Turkey, where they recognized three types: I, IIa, and IIb, which respectively make up 14 %, 70 % and 16 % of the serratus ventralis. In their study on take-off performance, Gardiner et al. (2014: 1059) determined a wing beat frequency of 11.95 ± 0.17 Hz. ECHOLOCATION: The data for 61 animals from Greece are the following: Fstart: 93.0 ± 11.84 kHz, Fend: 50.8 ± 1.16 kHz, Fpeak: 54.9 ± 5.32 kHz, bandwidth: 17.8 ± 14.30 kHz, duration 6.7 ± 1.29 msec and interpulse interval: 94.1 ± 35.73 msec (Papadatou et al., 2008b: 133). Walters et al. (2012: suppl.) report the following figures for 31 calls from France, Greece, Italy and Switzerland: duration: 6.37 ± 2.89 msec, Fmax: 73.24 ± 12.13 kHz, Fmin: 49.75 ± 1.33 kHz, bandwidth: 23.49 ± 11.29, Fpeak: 52.02 ± 1.68 kHz. Davies et al. (2013b: Table S8) report a peak frequency of 56.07 kHz and a range between 52.1 and 85.2 kHz. From Morocco, Disca et al. (2014: 226) indicate that the type of call is FM-QFC, with the following parameters: Fstart: 97.3 ± 11.3 kHz, Fend: 50.6 ± 1.4 kHz, Fpeak: 53.4 ± 1.2 kHz, FQFC-peak: 52.3 ± 1.1 kHz, bandwidth: 43.2 ± 11.4 kHz, duration: 6.2 ± 1.2 msec. For 3 calls from the Kalahari Desert, the characteristics were: Fchar: 47.6 ± 6.5 kHz, Fmax: 76.7 ± 7.7 kHz, Fmin: 53.2 ± 4.9 kHz, duration: 2.6 ± 0.4 msec(Adams and Kwiecinski, 2018: 4). Luo et al. (2019a: Supp.) reported the following data (from 2 calls): Fpeak: 54.9, 54.2 kHz, Fstart: 93, 85.2 kHz, Fend: 50.8, 52.1 kHz, Band width: 42.2, 33.1 kHz, and duration: 6.7, 5.8 msec. MOLECULAR BIOLOGY: DNA - Barragán et al. (2002), Appleton et al. (2004), Tian et al. (2004), Bilgin et al. (2006, 2008). Karyotype - Matthey and Bovey (1948) and Bovey (1949) described the karyotype from Switzerland as having a diploid number of 2n = 46, aFN = 48, X = submetacentric, Y = acrocentric. Rushton (1970: 463) reported 2n = 46, FN = 50, X = metacentric, Y = minute. Capanna and Civitelli (1965 - Italy), Baker et al. (1974 - Tunisia), Bickham and Hafner, 1978 - Yugoslavia), Zima (1978 - Czechoslovakia), Pérez-Suárez et al. (1991 - Spain), Albayrak (2006 - Turkey) found the fundamental number to be aFN = 50, and X = metacentric for Zima (1978). The autosomal chromosomes contain two large metacentrics, one medium-sized metacentric, 17 acrocentric and two dot-like pairs (one of these pairs may be bi-armed) (Arslan and Zima, 2014: 14). In Turkey, Asan Baydemir (2015: 350) found Nucleolus Organising Regions (NORs) on the secondary constriction in 2 pairs of chromosomes (pairs 18 and 20). Capanna and Civitelli (1965: 546) mention 22 pairs of autosomes, of which 3 are metacentric, 17 acrocentric, and two very small; the X chromosome is a medium sized metacentric and the Y chromosome is a very small one. Capanna and Manfredi Romanini (1971: 474) mention 2n = 45, aFN = 50, 2 pairs of metacentrics, 1 pair of submetacentrics, 19 pairs of acrocentrics. Gürün et al. (2014: 73) confirmed the pattern of local differentiation previously detected in mitochondrial DNA as they also found significant differences in the nuclear DNA between bats from North Africa, Lebanon, Cyprus, Anatolia, Russia, Thrace-Balkans, Slovakia, Italy, France, and Iberia. Gürün et al. (2019: 1866), however, could not confirm the mitochondrial differences in the microsatellite markers they examined, which would suggest a male-biased dispersal. HABITS: In southern France, Vincent et al. (2011: 57) found that during pregnancy and lactation, females might switch roosts within a 30 km radius around the maternity colony. During the night, the females travelled between 4.1 and 29.2 km to their foraging areas. They also estimated the individual homeranges for pregnant females to be 10,837 ha, and for lactating females 22,318 ha. DIET: Presetnik and Aulagnier (2013: 297) investigated the diet of M. schreibersii in Slovenia and found this to consist primarily of Lepidoptera (79 % by volume). Neuroptera (mostly Chrysopidae) accounted for 9.2 %), Diptera for 7.4 %, Trichoptera for 2.2 %, and Coleoptera for 1.4 %. African Chiroptera Report 2020 This seems to indicate that M. schreibersii is an aerial hawker. Gardiner et al. (2014: 1058) assessed its foraging strategy as "high altitude fast aerial hawking". PREDATORS: Mikula et al. (2016: Supplemental data) mention the Bat hawk (Macheiramphus alcinus Bonaparte, 1850) as diurnal avian predator. POPULATION: Structure and Density:- See Hutson et al. (2008c) [in IUCN (2009)] for information of populations outside Africa. Trend:- 2008: Decreasing (Hutson et al., 2008c; IUCN, 2009). LIFESPAN: Szekely et al. (2015: Supp.) and Lagunas-Rangel (2019: 2) reports a maximum longevity of 22 years. REPRODUCTION AND ONTOGENY: Tuttle and Stevenson (1982: 122) indicate that both sexes need about 16 months to attain sexual maturity. Kurta and Kunz (1987: 82) report that, at birth, the young weights about 20.4 % of the mother's body mass (3.4 g versus 20.4 g), and that its forearm length is 35.5 % of its mother's (16.7 versus 47.0 mm). Szekely et al. (2015: Suppl.) report that males become sexually mature after 730 days (24 months) and females after 577 days (19 months). The young is born after a gestation period of 100 days with a weight of 2.8 g (22 % of the adult). After weaning (66 days), its weight is increased to 12.8 g. In central Tunisia, females can be gravid until midJune (Dalhoumi et al., 2019a: 160). In Malawi, births occurred early in the wet season (Happold and Happold, 1990b: 568). Chari and Gopalakrishna (1984: 464) describe the development of the foetal membranes and the changes in the structure of the placenta in the Indian Miniopterus schreibersii fuliginosus, and they indicate that this bat has some unique embryological features. POSTNATAL DEVELOPMENT: In Iran, Sharifi and Vaissi (2013: 1) found that newborns had a mass of 3.74 ± 0.09 g and forearm length of 24.3 ± 0.31 mm. Both of these increased linearly over the first two weeks (at about 0.54 g/day and 1.39 mm/day), after which they remained rather stable. The epiphyseal gap of the fourth metacarpal phalangeal joint increased 637 until the thirteenth day, then decreased linearly until the 70th day and thereafter fused. The eyes opened during the first week and the ears bekame erect. The pups also began to move by that time. At 21 days, their ability to fly improved, and they started to fly freely in the cave. At that time their average body mass was 11.32 g (91.6 % of adult weight). PARASITES: Lei and Olival (2014: Suppl.) mention this species to be a host for Bartonella bacteria. Landau et al. (2012: 142) mention the Heamosporidians Polychromophilus melaniferus (Dionisi, 1899) and Polychromophilus corradetti Landau et al., 1980. In Europe, Genov et al. (1992) reported the following Nematodes: Molinostrongylus panousei Dollfus, 1954, M. skrjabini Skarbilovitch, 1934, M. alatus (Ortlepp, 1932), and M. ornatus (Mönning, 1927). Uchikawa (1985a: 22) proposed the new name Calcarmyobia dusbabeki (Acari: Myobiidae) for a mite found on Miniopterus schreibersii from amongst others Tunisia and Algeria. The Spinturnicid mites Spinturnix psi (Kolenati, 1856) and Spinturnix myoti (Kolenati, 1856) were reported by Postawa and Furman (2014: 390) from Central Anatolia and the Levant. Calcarmyobia ? parenzani Lombardini, 1956 was found on a M. schreibersii from Ras el Qued cave, Morocco. Rhipicephalus sanguineus (Latreille, 1806) (Arachnidae: Ixodida: Ixodidae) were reported from Algerian bats by Bendjeddou et al. (2013: 325). Bendjeddou et al. (2017: 15) reported the following ectoparasites from Algeria: Nycteribiidae: Nycteribia (Nycteribia) pedicularia Latreille, 1805, Penicillidia (Penicillidia) dufourii Westwood, 1835 and Phthiridium biarticulatum Hermann, 1804; Streblidae: Brachytarsina (Brachytarsina) flavipennis Macquart, 1851; and Siphonaptera: Rhinolophopsylla unipectinata arabs Jordan and Rothschild, 1921. In Italy, Voyron et al. (2011: 193) reported the following fungal entities from M. schreibersii carcasses: Aspergillus sp., Chrysosporium merdarium, Fusarium equiseti, Ophiostomatacea, Thielavia sp., Mucor hiemalis f. hiemalis. Ševcík et al. (2012: 35) report on the presence of the bat flies Nycteribia schmidlii schmidlii Schiner, 1853 and Penicillidia conspicua Speiser, 1901 on bats from Crete and Cyprus. These bat flies are 638 ISSN 1990-6471 vector hosts for the malaria parasite Polychromophilus melanipherus (see Witsenburg et al., 2013: 169). Haelewaters et al. (2017: 1) furthermore reported Nycteribia kolenati Theodor & Moscona, 1954 from Hungary. Additionally, Tortosa et al. (2013: 6) also reported the Nycteribiid bat fly Penicillidia oceanica (Bigot, 1885). The bat flies Nycteribia schmidlii Schiner, 1853, Penicillidia dufourii (Westwood, 1834), and Penicillidia conspicua Speiser, 1901 were reported by Postawa and Furman (2014: 390) from Central Anatolia and the Levant. Haelewaters et al. (2018: 794) additionally reported the flies Nycteribia parvula, Penicillidia indica, P. jenynsii, and P. oceanica from non-specified areas. In Europe, the tick Ixodes simplex Neumann, 1906 is highly specialized to M. schreibersii (see Hornok et al., 2014: 1). In Spain, Estrada-Peña and Serra-Cobo (1991: 346) found the following Acari: Ixodes (Pomerantzevella) simplex Neumann, 1906, Ixodes (Eschatocephalus) vespertilionis Koch, 1844, Macronyssus longimanus (Kolenati, 1856), Macronyssus granulosus (Kolenati, 1856), Macronyssus cyclaspis (Oudemans, 1906), Macronyssus rhinolophi (Oudemans, 1902), Macronyssus diversipilis (Vitzthum, 1920), Macronyssus ellipticus (Kolenati, 1857), Spinturnix psi (Kolenati, 1856), Eyndhovenia euryalis euryalis (Canestrini, 1884); and the bat flies Nycteribia schmidlii Schiner, 1853 and Penicillidia dufouri (Westwood, 1835). VIRUSES: Astroviridae Luis et al. (2013: suppl.) mention the presence of an unassigned bat astrovirus. Coronaviridae - Coronaviruses Luis et al. (2013: suppl.) report the occurrence of various Coronaviruses (e.g. BatCoV HKU7, Miniopterus bat coronavirus 1, Miniopterus bat coronavirus HKU8). Alphacoronavirus Kohl and Kurth (2014: 3112) report this virus from Spain and Germany. Betacoronavirus Kohl and Kurth (2014: 3112) report this virus from Bulgaria and Germany. Filoviridae - Filoviruses Cuevavirus - Reported from Spain by Kohl and Kurth (2014: 3113). Negredo et al. (2011: 1) described a new virus (Lloviu virus [LLOV]) from M. schreibersii in Spain. Flaviviridae Flavivirus de Jong et al. (2011: 10), Luis et al. (2013: suppl.) report Japanese encephalitis virus. Luis et al. (2013) also mentions Kyasanur Forest disease virus and Yokose virus. Orthomyxoviridae Despite sampling and testing of 2 individuals of this species, no evidence of influenza A-like viruses (Orthomyxoviridae) were obtained (Fereidouni et al., 2015). Papillomaviridae Dyolambda A new papillomavirus (Miniopterus schreibersii papillomavirus type 1 [MscPV1]) was described by Tse et al. (2012: 5) from Hong Kong as the first member of the novel Dyolambda-papillomavirus genus. Phenuiviridae Phlebovirus - Rift Valley fever virus: de Jong et al. (2011: 11), Luis et al. (2013: suppl.) report the occurrence of Rift Valley fever virus. Picornaviridae Zeghbib et al. (2019: 3) reported a new Mischivirus from a bat from Algeria. Rhabdoviridiae Lyssavirus - Rabies related viruses de Jong et al. (2011: 9) report the presence of West Caucasian bat virus. van Eeden et al. (2011: 67) report an unconfirmed, but possible occurrence of Duvenhage virus in a specimen from the Limpopo province, South Africa. In Algeria, Serra-Cobo et al. (2018: 2) found seven bats out of 63 testing seropositive for European bat lyssavirus 1. "Various European bat lyssaviruses" were reported by Kohl and Kurth (2014: 3114) from a variety of European countries. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Morocco, Tunisia. African Chiroptera Report 2020 639 Figure 235. Distribution of Miniopterus schreibersii Miniopterus sororculus Goodman, Ryan, Maminirina, Fahr, Christidis and Appleton, 2007 *2007. Miniopterus sororculus Goodman, Ryan, Maminirina, Fahr, Christidis and Appleton, J. Mamm., 88 (5): 1219, figs 2 - 6. Type locality: Madagascar: Province de Fianarantsoa: unnamed cave: Ambatofinandrahana, 3 km S [2034.321S 4648.530E, 1450 m] [Goto Description]. Holotype: FMNH 177259: ad ♀, skull and alcoholic. Collected by: Steven M. Goodman; collection date: 23 March 2003; original number: SMG 13560. Paratype: FMNH 177255: Collected by: Steven M. Goodman; collection date: 23 March 2003. Paratype: FMNH 177256: Collected by: Steven M. Goodman; collection date: 23 March 2003. Paratype: FMNH 177257: Collected by: Steven M. Goodman; collection date: 23 March 2003. Paratype: FMNH 177258: Collected by: Steven M. Goodman; collection date: 23 March 2003. Paratype: FMNH 177260: Collected by: Steven M. Goodman; collection date: 23 March 2003. - Etymology: The name sororculus is derived from the Latin ("sororcula") and means "small sister" (Goodman et al., 2007b: 1222). This epithet was chosen to contrast this taxon with its African counterpart fraterculus, which means "little brother", as well as this animal being slightly smaller than M. majori (Goodman et al., 2007b: 1222). (Current Combination) 2018. Miniopterus soroculus: Kemp, López-Baucells, Rocha, Wangensteen, Andriatafika, Nair and Cabeza, Agric. Ecosyst. Environ., 369: 90 (for 2019). Publication date: 1 October 2018. (Lapsus) ? Miniopterus sororculus fraterculus: (Name Combination) TAXONOMY: Specimens from Madagascar assigned to M. fraterculus by Peterson et al. (1995) and Simmons (2005) are now considered to be a distinct species (Goodman et al., 2007b). COMMON NAMES: Czech: létavec sesterský. English: Sororculus Longfingered Bat, Sororcula Long-fingered Bat. German: Madagaskar-Langflügelfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) as the species appears to have a broad distribution in the central highlands with a relatively wide elevational range. There are no obvious major threats, and based on current records seems to be at least somewhat adaptable to disturbed habitats (Jenkins et al., 2008f; IUCN, 2009; Monadjem et al., 2017t)). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017t). 2008: LC ver 3.1 (2001) (Jenkins et al., 2008f; IUCN, 2009). Regional None known. MAJOR THREATS: There are no known threats to this species and it exists in areas some distance from native forest (Goodman et al., 2007b; Jenkins et al., 2008f; IUCN, 2009; Monadjem et al., 2017t). 640 ISSN 1990-6471 CONSERVATION ACTIONS: Jenkins et al. (2008f) [in IUCN (2009)] and Monadjem et al. (2017t) report that at present, there are no known records from protected areas. This is a recently described taxon that does not appear to rely on large tracts of intact forest for its survival (Goodman et al., 2007b). Further studies are needed to understand its adaptation to high elevations and extreme climate (Goodman et al., 2007b). GENERAL DISTRIBUTION: Miniopterus sororculus is endemic to Madagascar where it has mainly been recorded from sites above 900 m elevation (Goodman et al., 2007b) although there are a few records from sites at elevations < 50 m (Goodman et al., 2008a). It is known from < 10 m confirmed localities ranging in elevation from 40 to 2,200 m above sea level. It is presumed to have a broad distribution spanning various portions of the central highlands (Goodman et al., 2007b). Schoeman et al. (2014: 28; 31) found the most suitable area for this species to be located in the central highland part of the island (higher elevations of the Central Highlands - sub-humid region). Native: Madagascar (Goodman et al., 2007b; Goodman et al., 2008a). ECHOLOCATION: Kofoky et al. (2009: 382) reported the calls as Miniopterus majori/sororculus, which produced broadband FM/QCF echolocation calls at low duty cycle, but with a lower maximum energy, at about 48.5 kHz. The fundamenetal being most intense with short duration pulses of about 4.5 ms. Ramasindrazana et al. (2011: 294) recorded 20 calls from 7 Miniopterus sororculus specimens, and found the following echolocation call values: Frequency of maximum energy: 55.3 ± 0.92 (53.9 - 56.6) kHz, Maximum frequency: 103.2 ± 9.04 (83.0 - 121.0) kHz, Minimum frequency: 51.7 ± 1.08 (50.0 - 53.0) kHz, Call duration: 3.3 ± 0.29 (2.7 - 3.9) ms, Inter pulse interval: 79.3 ± 13.06 (50.8 - 99.8) ms. HABITAT: The holotype of M. sororculus was collected in a small limestone cave surrounded by pseudosteppe at 1,450 m (Goodman et al., 2007b). This species is known to use rock crevices and deeper caves as day roosts (Goodman et al., 2007b). At Antsampandrano, this species was found occupying a day roost in the attic of a building occupied by people. Further, it has been obtained in open dry savanna in the central west of Madagascar (Goodman et al., 2007b). ROOST: Wilkinson et al. (2012: 160) mention that caves are the typical roosting sites for this species on Madagascar. POPULATION: Structure and Density:- There is no information available on the population status of this recently described endemic species (Jenkins et al., 2008f; IUCN, 2009; Monadjem et al., 2017t). Trend:- 2016: Unknown (Monadjem et al., 2017t). 2008: Unknown (Jenkins et al., 2008f; IUCN, 2009). PARASITES: Dietrich et al. (2014: Suppl.) and Gomard et al. (2016: 5) report the presence of spirochaetes of the genus Leptospira, which Dietrich et al. (2018a: 3) identified as L. borgpetersenii. Ramasindrazana et al. (2016: 6) recovered Litomosa Clade 1 and 2 (Nematoda: Onchocercidae) from this species. Tortosa et al. (2013: 4) reported the presence of Penicillidia leptothrinax, a Nycteribiid bat fly. Ramasindrazana et al. (2017: Suppl.) mentioned the bat fly Nycteribia stylidiopsis (but the specimen on which this was found - UADBA 43264 - is here considered to be a M. majori). VIRUSES: Paramyxoviridae Wilkinson et al. (2012: 160) tested 1 individuals from the Mauritius using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 1 positive results for viral nucleic acids. 2 out of 22 Madagascan specimens tested by Mélade et al. (2016b: 4) were positive for paramyxoviruses. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. African Chiroptera Report 2020 641 Figure 236. Distribution of Miniopterus sororculus Miniopterus villiersi Aellen, 1956 1956. 1965. 2006. 2006. 2017. ? Miniopterus inflatus villiersi Aellen, Bull. IFAN, (A) 18: 890. Type locality: Guinea: Dalaba: Grotte du Marché [10 42 N 12 15 W] [Goto Description]. Holotype: MHNG IFAN 54-1-1: ad ♂. Collected by: A. Villiers; collection date: 16 April 1954; original number: 2839. Former number IFAN: 54-1-1. - Comments: Considered a valid subspecies by Fahr and Ebigbo (2003: 128; 2004: 73), Fahr (2007a: 104). Aellen (1956a: 891) mentions 2 ♂♂ and 5 ♀♀ paratypes. Miniopterus schreibersii villiersi: Koopman, Am. Mus. Novit., 2219: 19. Publication date: 22 June 1965. (Name Combination) Miniopterus (schreibersii) villiersi: Fahr, Djossa and Vierhaus, Rapid assessment of bats (Chiroptera) in Déré, Diécké and Mt. Béro classified forests, southeastern Guinea; including a review of the distribution of bats in Guinée Forestière., 40: 74. (Name Combination) Miniopterus villiersi: Fahr, Djossa and Vierhaus, Rapid assessment of bats (Chiroptera) in Déré, Diécké and Mt. Béro classified forests, southeastern Guinea; including a review of the distribution of bats in Guinée Forestière: 74.. (Current Combination) Miniopterus screibersii villiersi: Reardon and Schoeman, Acta Chiropt., 19 (2): Suppl.. Publication date: December 2017. (Lapsus) Miniopterus fraterculus villiersi: TAXONOMY: For a long time, villiersi was considered to be a synonym or a subspecies of either M. inflatus, M. schreibersii or M. natalensis. Fahr et al. (2006a: 74), Weber and Fahr (2006: "4"), Jacobs et al. (2008x: 74) and Monadjem et al. (2016y: 371) indicated that villiersi represents a separate species (restricted to the Upper-Guinea forest zone), a view which is currently gaining support as detailed genetic analyses indicate the presence of multiple taxa in West Africa, and is confirmed by Monadjem et al. (2020: 237, 248). COMMON NAMES: German: Lanflügelfledermaus. GENERAL DISTRIBUTION: Probably restricted to the Upper-Guinea forest zone, including Guinea (Decher et al., 2016: 274). ECHOLOCATION: Monadjem et al. (2020: 248) found that the knee frequency of the echolocation calls from five handreleased M. villiersi specimens ranged between 51.1 and 52.4 kHz (average 51.6 kHz). HABITAT: Monadjem et al. (2016y: 371) recorded M. (schreibersii) villiersi from altitudes between 460 and 900m in the Mount Nimba area. ROOST: Monadjem et al. (2016y: 371) found Miniopterus (schreibersii) villiersi roosting in old mine adits in the Mount Nimba area. 642 ISSN 1990-6471 PARASITES: HAEMOSPORIDA Schaer et al. (2013a: 17416) report the presence of hemosporidian parasites of the genus Polychromophilus in three out of 12 investigated M. villiersi specimens from West Africa. Rosskopf et al. (2018: Suppl.) also found Polychromophilus sp. in bats from Guinea. ACARI Uchikawa (1985a: 19) indicates that Miniopterus villiersi is one of the true hosts of "typical" Calcarmyobia congoensis Uchikawa, 1982 (Acari: Myobiidae). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Cameroon, Guinea, Nigeria. Figure 237. Distribution of Miniopterus villiersi Miniopterus "incertae-sedis" 2014. 2014. 2014. 2014. 2014. 2014. 2019. Miniopterus sp. P3: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. Miniopterus sp. P4: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. Miniopterus sp. P5: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. Miniopterus sp. P6: Christides, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. Miniopterus sp. P7: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. Miniopterus X3: Christidis, Goodman, Naughton and Appleton, PLoS ONE, 9 (3) e92440: 1. Publication date: 18 March 2014. - Comments: Christidis et al. (2014: 4) found this haplotype - respresented by a single specimen - forming a sub-clade with Miniopterus sororculus and M. mahafaliensis. Miniopterus manavi 2: Demos, Webala, Lutz, Kerbis Peterhans, Goodman, CortésDelgado, Bartonjo and Patterson, Zool. Scr., 49 (1): 5. Publication date: 22 October 2019. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Madagascar. †Family PHILISIDAE Sigé, 1985 1985. Philisidae Sigé, 19: ?. TAXONOMY: Ravel et al. (2014) confirmed the family status of the Philisidae within the superfamily Vespertilionoidae based on a cladistic assesment of the dental data, and that they represent the earliest offshoot of this superfamily (p. 698). Ravel et al. (2012) suggested that Philisis and Dizzya were probably split of from the third genus of the family (Witwatia) in the Early Eocene. Ravel et al. (2012: 699), however, doubt the close relationship between the two former genera. Sigé (1985) [in Ravel et al. (2014: 696)] concluded that the Philisidae could have originated in Late or Middle Eocene from a generalized Vespertilionidae after the divergence between the Molossidae and the Vespertilionidae sensu lato. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: According to Ravel et al. (2014: 691), Philisids were among the most common African bats in the Palaeogene (66 to 23.03 mya). They were found in the Late Eocene - Early Oligocene deposits of the Fayum African Chiroptera Report 2020 Depression in Egypt, together with extant bat families, such as Rhinopomatidae, Megadermatidae, Emballonuridae and Myzopodidae Gunnell et al., 2008, 2014). 643 (Sigé, 1985; †Genus Dizzya Sigé, 1991 *1991. Dizzya Sigé, N. Jb. Geol. Paläont. Abh., 182: ?. - Comments: Type species - Dizzya exultans Sigé, 1991. (Current Combination) TAXONOMY: Currently recognized species of Dizzya: †exsultans Sigé 1991. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Early. Eocene. DENTAL FORMULA: Ravel et al. (2011: 402) indicate that Dizzya differs from the other Philisidae in showing a nyctalodont molar structure. †Dizzya exsultans Sigé, 1991 *1991. Dizzya exsultans Sigé, N. Jb. Geol. Paläont. Abh., 182: ?. Type locality: Tunisia: Chambi. Comments: Marandat (1994: 62) mentions the holotype to consist of a right M1 or M2. . (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Eocene (Ypresian - Brown et al., 2019: Suppl.). Together with Witwatia sigei, it is also the oldest. Based on data from Gunnell et al. (2009), Ravel et al. (2014: 702) estimated the body weight for this bat to be about 8.6 to 9.1 g. DETAILED MORPHOLOGY: Ravel et al. (2014: 694) indicate that Dizzya exsultans is the smallest representative of the Philisidae. †Genus Philisis Sigé, 1985 *1985. Philisis Sigé, Geol. Palaeontol., 19: ?. - Comments: Type species - Philisis sphingis Sigé, 1985. (Current Combination) TAXONOMY: May be a junior synonym of Provampyrus. Ravel et al. (2014: 699) indicate that the genus might be paraphyletic. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Ravel et al. (2014: 699) indicate that fossils of this genus were only found in deposits dating from the Early Oligocene. Currently recognized species of Philisis: †sevketi Sigé, Thomas, Sen, Gheerbrant, Roger & Al-Sulaimani 1994 - Oman; †sphingis Sigé 1985. †Philisis sphingis Sigé, 1985 *1985. Philisis sphingis Sigé, Geologica et Palaeontologica, 19: ?. Holotype: YPM MAM 034488:. Right maxilla (P4-M3): see Gunnell et al. (2008: 2). (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Oligocene (Rupelian - 32.46 - 29 mya) (see Brown et al., 2019: Suppl.). DETAILED MORPHOLOGY: Based on data from Gunnell et al. (2009), Ravel et al. (2014: 702) estimated the body weight for this bat to be between 36 to 74 g. 644 ISSN 1990-6471 †Genus Witwatia Gunnell, Simons and Seiffert, 2008 *2008. Witwatia Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 2. - Comments: Type species Witwatia schlosseri Gunnell, Simons and Seiffert, 2008. - Etymology: Wit Wat, Egyptian Arabic for large, flapping wings (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). SIMILAR SPECIES: See Gunnell et al. (2008). Ravel et al. (2014: 702) mention that the differences between W. schlosseri and W. eremicus (both found in the same locality: BQ2, Fayum, Egypt), might be attributed to the presence of a sexual dimorphism, where the former might represent the larger female and he latter the smaller male. But they also indicate that this is currently speculative. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). Currently recognized species of the genus Witwatia: †eremicus Gunnell, Simons and Seiffert, 2008; †schlosseri Gunnell, Simons and Seiffert, 2008; †sigei Ravel, Marivaux, Tabuce, Bel Haj Ali, Essid, and Vianey-Liaud, 2012. Timeframe - Ravel et al. (2014: 699) indicate that fossils were found dating back between the early and latest Eocene. GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). †Witwatia eremicus Gunnell, Simons and Seiffert, 2008 *2008. Witwatia eremicus Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 3, 4, 5. Type locality: Egypt: Fayum Depression: Quarry BQ-2: Umm Rigl Member of the Birket Qurun Formation. Etymology: Eremikos, Greek for solitude, in reference to the vast solitude of the Egyptian Western Desert (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). SIMILAR SPECIES: See Gunnell et al. (2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). Timeframe: Earliest Late Eocene (Priabonian - Brown et al., 2019: Suppl.), Umm Rigl Member of the Birket Qarum formation. GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). DETAILED MORPHOLOGY: Based on data from Gunnell et al. (2009), Ravel et al. (2014: 702) estimated the body weight for this bat to be about 40 to 98 g. †Witwatia schlosseri Gunnell, Simons and Seiffert, 2008 *2008. Witwatia schlosseri Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 2, 3, fig. 3D. Type locality: Egypt: Fayum Depression: Quarry BQ-2: Umm Rigl Member of the Birket Qarun Formation. Holotype: CGM 83668:. Left dentary (c1-m3): see Gunnell et al. (2008: 3). Etymology: Specific name for Max Schlosser who described the first fossil bat from the Fayum (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). SIMILAR SPECIES: See Gunnell et al. (2008). African Chiroptera Report 2020 PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). Timeframe: Earliest Late Eocene (Priabonian - Brown et al., 2019: Suppl.), Umm Rigl Member of the Birket Qarum formation. GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). 645 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). DETAILED MORPHOLOGY: Based on data from Gunnell et al. (2009), Ravel et al. (2014: 702) estimated the body weight for this bat to be between 57 and 116 g. †Witwatia sigei Ravel, Marivaux, Tabuce, Bel Haj Ali, Essid and Vianey-Liaud, 2012 *2012. Witwatia sigei Ravel, Marivaux, Tabuce, Bel Haj Ali, Essid and Vianey-Liaud, Palaeontology, 55 (5): 1035, 1036, fig. 2. Publication date: 18 September 2012. Type locality: Tunisia: Kasserine area: Chambi [Goto Description]. Holotype: ONM CB1-230: unknown / no information. Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Left upper molar (M2), see Ravel et al. (2012: 1036). - Etymology: In honour of Bernard Sigé, who first extensively described Paleogene bats remains, including those from Chambi (see Ravel et al., 2012: 1036). (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Eocene (Ypresian - Brown et al., 2019: Suppl.). DETAILED MORPHOLOGY: Based on data from Gunnell et al. (2009), Ravel et al. (2014: 702) estimated the body weight for this bat to be about 100 g. Family VESPERTILIONIDAE Gray, 1821 1815. Vespertilia Rafinesque, Analyse de la Nature, 54. - Comments: Type genus: Vespertilio Linnaeus, 1758. Conserved by Direction 98 (IUCN , 1958), which also corrected the name to Vespertilionidae. Originally included the subfamilies Lophinia (Rafinesque, 1815) [including the genera Rhinolophus Lacépède, 1799; Phyllostoma G. Cuvier, 1800 [= Phyllostomus Lacépède, 1799]; Vampyrum Rafinesque, 1815; and Megaderma É. Geoffroy, 1810]; and Nycteria (Rafinesque, 1815) [including the genera Pteropus Brisson, 1762; Eidolon Rafinesque, 1815; Pteronotus Rafinesque, 1815; Cephalotes É. Geoffroy, 1810 [= Nyctimene Borkhausen, 1797]; Tadaris [sic = Tadarida] Rafinesque, 1814; Vespertilio Linnaeus, 1758; Nycterus [sic = Nycteris] É. Geoffroy and G. Cuvier, 1795; Noctilio Linnaeus, 1766; Molossus É. Geoffroy, 1805; and Atalapha Rafinesque, 1814 [= Nyctalus Bowdich, 1825: 36]] (see Jackson and Groves, 2015: 264). *1821. Vespertilionidæ Gray, London Med. Repos., 15: 299. Publication date: 1 April 1821. Comments: Type genus: Vespertilio Linnaeus, 1758. Dobson (1878: 167) refers to Dobson, 1875 (p. 347), indicating that Gray's (1866) definition includes genera beloning to very distinct families. Originally included the genera Megadermes [sic = Megaderma] É. Geoffroy, 1810; Rhynolophus [sic = Rhinolophus] Lacépède, 1799; Nycterus [sic = Nycteris] É. Geoffroy and G. Cuvier, 1795; Rhynopoma [sic = Rhinopoma] É. Geoffroy, 1818; Thaphosores [sic = Taphozous] É. Geoffroy, 1818; Vespertilio Linnaeus, 1758; Pecotus [sic = Plecotus] É. Geoffroy, 1818; and Barbastella J. Gray, 1821 (see Jackson and Groves, 2015: 264). The name was conserved by Direction 98 (ICZN, 1958: 131). (Current Combination) 1825. Vespertilionina Gray, Ann. Philos., 10 (5): 339. Publication date: November 1825. Comments: Type genus: Vespertilio Linnaeus, 1758. Originally include the genera Vespertilio Linnaeus, 1758; Plecotus É. Geoffroy, 1818; Barbastellus J. Gray, 1825 [= Barbastella J. Gray, 1821]; Proboscidea Spix, 1823 [= Rhynchonycteris Peters, 1867; Thyroptera Spix, 1823; and Caelano [sic = Celaeno] Leach, 1821 [= Noctilio Linnaeus, 1766: 88] (see Jackson and Groves, 2015: 264). 1843. Gymnorhina Wagner. 646 ISSN 1990-6471 1855. 1866. 1866. 1866. 1866. 1869. 1872. 1893. 1893. 1907. 1942. 2008. 2018. ? Nycticeina Gervais, in: F. Comte de Castelnau, Exped. Partes Cen. Am. Sud., Zool, ???. Publication date: 23 July 1856. Nyctophilina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 91. Publication date: 1 February 1866. Plecotina Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 91. Publication date: 1 February 1866. Romiciana Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 90. Publication date: 1 February 1866. Vespertiliones Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 1865: 524. - Comments: Type genus: Vespertilio Linnaeus, 1758. Originally included the genera Synotus Keyserling & Blasius, 1839 [= Barbastella J. Gray, 1821]; Plecotus É. Geoffroy, 1818; Histiotus Gervais, 1855; Otonycteris Peters, 1859; Miniopterus Bonaparte, 1837; Vespertilio Linnaeus, 1758; Vesperugo Keyserling and Blasius, 1839 [= Vespertilio Linnaeus, 1758]; Vesperus Keyserling and Blasius, 1839 [= Vespertilio Linnaeus, 1758]; Murina J. Gray, 1842; Harpiocephalus J. Gray, 1842; Nycticejus [sic = Nycticeius] Rafinesque, 1819]; Atalapha Rafinesque, 1814 [= Nyctalus Bowdich, 1825]; Thyroptera Spix, 1823; and Antrozous H. Allen, 1862 (see Jackson and Groves, 2015: 265). Jackson and Groves (2015: 227) consider this (family) name a homonym of Vespertiliones Pallas, 1767, and indicated that it covers the Vespertionidae family, rather than the Vespertilioninae subfamily. Not Vespertiliones Dobson, 1878, a group name for the subfamily Vespertilioninae. Gymnorhinida Fatio, Faune des Vertébrés de la Suisse. Vespertilionidae: Gill, Smiths. Misc. Coll., 11 (1): 17. Publication date: November 1872. Comments: Type genus: Vespertilio Linnaeus, 1758. Originally included the subfamilies Vespertilioninae J. Gray, 1821 and Nycticejinae Gill, 1872 (see Jackson and Groves, 2015: 265). Nycteridae: Schulze. - Comments: Not of Van der Hoeven, 1855. Not of Dobson, 1875 (see Pavlinov et al., 1995: 93). Vespertilionini: Winge, Samling af Afhandlinger. E Museo Lundii, 2 (1): 24. - Comments: Introduced as tribe (?). Jackson and Groves (2015: 265) (erroneously?) mention Emballonura Temminck, 1838 as type genus. Originally included the genera Vespertilio Linnaeus, 1758 [including the subgenera Kerivoula J. Gray, 1842; Natalus J. Gray, 1838; and Nyctiellus Gervais, 1855]; Plecotus É. Geoffroy, 1818; Minyopterus Agassiz, 1846 [= Miniopterus Bonaparte, 1837]; Lasionycteris Peters, 1865; Vesperugo Keyserling and Blasius, 1839 [= Vespertilio Linnaeus, 1758]; Harpyiocephalus J. Gray, 1866 [= Harpiocephalus J. Gray, 1842]; Synotus Keyserling & Blasius, 1839 [= Barbastella Gray, 1821]; Chalinolobus Peters, 1865; Otonycteris Peters, 1859; Nyctophilus Leach, 1821; Atalapha Rafinesque, 1814 [= Nyctalus Bowdich, 1825]; and Antrozous H. Allen, 1862 (see Jackson and Groves, 2015: 265). Nyctophilinæ Miller, Bull. U.S. natl. Mus., 57: xi, 234. Publication date: 29 June 1907. Comments: Type genus: Nyctophilus Leach, 1821. Originally included the genera Antrozous H. Allen, 1862 and Nyctophilus Leach (see Jackson and Groves, 2015: 270). Myotini Tate, Bull. Am. Mus. Nat. Hist., 80 (7): 221, 229. Publication date: 27 November 1942. - Comments: Type genus: Myotis Kaup, 1829. Introduced as tribe, and originally included the genera Myotis Kaup, 1829; Lasionycteris Peters, 1865; Plecotus É. Geoffroy, 1818; Corynorhinus H. Allen, 1865; Idionycteris Anthony, 1923; and Euderma H. Allen, 1891 (see Jackson and Groves, 2015: 280). Vespertolionidae: Selim, Nahla and Shelfeh, Tishreen Univ. J. Res. Sci. Stud. - Biol. Sci. Ser., 30 (1): 247. (Lapsus) Vespertilionnidae: Malekani, Musaba, Gembu, Bugentho, Toengaho, Badjedjea, Ngabu, Mutombo, Laudisoit, Ewango, Van Cakenberghe, Verheyen, Asimonyo, Masudi, Bongo and Ngbolua, Nat. Conserv. Res., 3 (1): 67.. Publication date: January 2018. (Lapsus) Vespertilioninae sp.: TAXONOMY: Systematicians have long recognized that Vespertilionidae as so defined, is diagnosed by few if any apomorphies, and some data have been cited as evidence that the family is not monophyletic. For example, Tomopeas ravus (the only member of the subfamily Tomopeatinae) shares immunological affinities and several derived morphological and molecular traits with molossidis (Miller, 1907; Davis, 1970; Barkley, African Chiroptera Report 2020 1984; Pierson, 1986; Sudman et al. (1994). Miniopterines, which differ from other vespertilionids in several aspects of fetal development and adult morphology, show immunological affinity with molossids (Mein and Tupinier, 1977; Gopalakrishna and Chari, 1983; Pierson, 1986). Kerivoulines share derived morphological traits with Natalidae, Thyropteridae, Furipteridae, and Myzopodidae (Sigé, 1974; Van Valen, 1979). These patterns have lead various workers to suggest removal of one or more subfamilies from Vespertilionidae (Davis, 1970; Sigé, 1974; Mein and Tupinier, 1977; Van Valen, 1979; Gopalakrishna and Chari, 1983; Barkley, 1984; Sudman et al., 1994). Each of the eight vespertilionid subfamilies just listed appears to be monophyletic, with the exception of Vespertilioninae, which may be paraphyletic. In morphology, Vespertilioninae is "perhaps best characterized by absence of the special modifications that distinguish the other groups" (Miller, 1907: 197). Volleth and Heller, 1994) describe several derived karyotype features that diagnose Vespertilioninae (including Nyctophilinae but exclude Myotinae), but their study did not include any members of Antrozoinae or Lasiurini. Molecular phylogenetics reviewed by Hoofer and Van Den Bussche (2003). Roehrs et al. (2011: 24) suggest the following subdivision in tribes/groups: Vespertilionini (Nyctalus, Pipistrellus, Scotoecus, Vespertilio) Hypsugine group (Chalinolobus, Vespadelus, Hypsugo, Nycticeinops, Neoromicia, Laephotis, Tylonycteris) Eptesicini / Nycticeiini (Arielulus, Lasionycteris, Glauconycteris, Nycticeius, Eptesicus, Scotomanes) Perimyotine group (Parastrellus, Perimytois) Scotophilini (Scotophilus) Antrozini (Antrozous, Rhogeessa, Baeodon) Lasiurini (Lasiurus) Plecotini (Corynorhinus, Otonycteris, Barbastella, Plecotus, Idionycteris). Known genera in the subfamily Vespertilioninae: Tribe Eptesicini Volleth and Heller, 1994 - Arielulus Hill and Harrison, 1987; Eptesicus Rafinesque, 1820; Hesperoptenus Peters, 1868. Tribe Lasiurini Tate, 1942 - Lasiurus Gray, 1831. Tribe Nycticeiini Gervais, 1855 - Nycticeinops Hill and Harrison, 1987; Nycticeius Rafinesque, 1819; Rhogeessa H. Allen, 1866; Scoteanax Troughton, 1943; Scotoecus Thomas, 1901; Scotomanes Dobson, 1875; Scotophilus Leach, 1821; Scotorepens Troughton,1943. Tribe Nyctophilini Peters,1865 - Nyctophilus Leach, 1821; Pharotis Thomas, 1914. 647 Tribe Pipistrellini Tate, 1942 - Glischropus Dobson, 1875; Nyctalus Bowditch, 1825; Pipistrellus Kaup, 1829; Scotozous Dobson, 1875. Tribe Plecotini Gray, 1866 - Barbastella Gray, 1821; Corynorhinus H. Allen, 1865; Euderma H. Allen, 1892; Idionycteris Anthony, 1923; Otonycteris Peters, 1859; Plecotus E. Geoffroy Saint-Hilaire, 1818. Tribe Vespertilionini Gray, 1821 - Chalinolobus Peters, 1867; Eudiscopus Conisbee, 1953; Falsistrellus Troughton, 1943; Glauconycteris Dobson, 1875; Histiotus Gervais, 1856; Hypsugo Kolenati, 1856; Ia Thomas, 1902; Laephotis Thomas, 1901; Mimetillus Thomas, 1904; Neoromicia Roberts, 1926; Philetor Thomas, 1902; Tylonycteris Peters, 1872; Vespadelus Troughton, 1943; Vespertilio Linnaeus, 1758. Currently (Simmons and Cirranello, 2020) recognized subdivision (subfamilies, tribes and genera) of the family Vespertionidae: - Kerivoulinae Miller, 1907: Kerivoula Gray, 1842; Phoniscus Miller, 1905. - Murininae Miller, 1907: Harpiocephalus Grau, 1842; Harpiola Thomas, 1915; Murina Gray, 1842. - Myotinae Tate, 1942: Eudiscopus Conisbee, 1953; Myotis Kaup, 1829; Submyotodon Ziegler, 2003. - Vespertilioninae Gray, 1821: Antrozoini Miller, 1897: Antrozous H. Allen, 1863; Baeodon Miller, 1906; Rhogeessa H. Allen, 1866. Lasiurini Tate, 1942: Lasiurus Gray, 1831. Nycticeiini Gervais, 1866: Arielulus Hill and Harrison, 1987; Eptesicus Rafinesque, 1820; Glauconycteris Dobson, 1875; Hesperoptenus Peters, 1869; Ia Thomas, 1902; Lasionycteris Peters, 1866; Nycticeius Rafinesque, 1819; Scoteanax Troughton, 1944; Scotomanes Dobson, 1875; Scotorepens Troughton, 1944; Thainycteris Kock and Storch, 1996. Plecotini: Barbastella Gray, 1821; Corynorhinus H. Allen, 1865; Euderma H. Allen, 1892; Idionycteris Anthony, 1923; Otonycteris Peters, 1860; Plecotus E. Geoffroy Saint Hilaire, 1818. Pipistrellini (Tate, 1842): Glischropus Dobson, 1875; Nyctalus Bowdich, 1825; Pipistrellus Kaup, 1829; Scotoecus Thomas, 1901; Scotozous Dobson, 1875. Scotophilini Leach, 1821: Scotophilus Leach, 1821. Vespertilionini Gray, 1821: Cassistrellus Ruedi, Eger, Lim and Csorba, 2017; Chalinolobus Peters, 1867; Falsistrellus Troughton, 1944; Hypsugo Kolenati, 1856; Laephotis Thomas, 1901; Mimetillus Thomas, 1904; Mirostrellus Görföl, Kruskop, Tu, Estók, Son and Csorba, 2020; Nycticeinops Hill and Harrison, 1987; Nyctophilus Leach, 1821; Parahypsugo Hutterer, Decher, Monadjem and Astrin, 2019; Pharotis Thomas, 1914; Philetor Thomas, 1902; Tylonycteris Peters, 1872; Vespadelus Troughton, 1944; Vespertilio 648 ISSN 1990-6471 Linnaeus, 1758. Unassigned to tribe: Parastrellus Hoofer, Van Den Bussche and Horácek, 2006; Perimyotis Menu, 1984; Rhyneptesicus Bianchi, 1917. COMMON NAMES: Castilian (Spain): Murcielagos chicos, Murciélagos de cola envainada. Czech: netopýrovití, netopýři holonosí. Dutch: Gladneusvleermuizen. English: Evening bats, Vespertilionid bats, Vesper Bats, Plain-faced Bats, Simple-nosed Bats. Finnish: Siipat. French: Vespertilionidés. German: Glattnasen, Glattnasen-Fledermäuse. Italian: Vespertiliònidi. Norwegian: Glattneser, Glattneseflaggermus. Polish: mroczkowate. Romanian: Lilieci cu tragus. Russian: Гладконосые. Ukrainian: Лиликові [= Lylykovi]. Vietnamese: Họ dơi muỗi. ETYMOLOGY OF COMMON NAME: Rosevear (1965) uses the colloquial name vesper bats, borrowing it from the name for the family, which is derived from the Latin for a bat, vespertilio. They are sometimes called 'evening bats' because of their time of activity (Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Shi and Rabosky (2015: 1537) report the familylevel stem ages/crown ages for the Vespertilionidae (including Miniopteridae and Cistugidae) to be 52.1 and 51.1 million years ago, respectively. The oldest African fossil assigned to this family is Khonsunycteris aegypticus from the late Eocene (ca. 34 MYA) of the Fayum, Egypt (Hand et al., 2016: 7). GENERAL DISTRIBUTION: Luo et al. (2019b) investigated the geographic range of 126 vespertilionid bat species (including 20 African) in relation to their wing morphology and found that species with higher wing loading have larger distribution ranges than those with lower wing loading, and that the size of geographic ranges was associated with wing aspect ratio. MOLECULAR BIOLOGY: Sotero-Caio et al. (2017: 5) indicate that familywide the karyotype has a 2n value between 18 and 52. ROOST: López-Baucells et al. (2018: 30) examined the roost selection by synanthropic bats in rural Madagascar and found that small colonies of vespertilionid bats had a preference for small, traditional houses made of mud and wood, especially if the roof was made from leaves. PREDATORS: Mikula et al. (2016: Supplemental data) mention a Vespertiliondae bat taken as prey by a Swamp boubou (Laniarius bicolor (Verreaux, 1857)) in Botswana. PARASITES: Fain (1994: 1280) reports that three genera of Myobiidae (Acari) are occurring on members of the Vespertilionidae: Acanthophthirius, with 4 subgenera (Acanthophtirius, Myotimyobia, Chiromyobia, Mimetillobia) and 47 species, Pteracarus with 21 species, and Calcarmyobia with 18 species and 8 subspecies. The latter genus is restricted to bats of the genus Miniopterus, and since this genus is currently included in a family of its own, this implies that only two genera of Myobiidae parasite on Vespertilionidae. VIRUSES: A paramyxovirus was detected in one out of nine Vespertilionidae bats sampled in Kirindy in Madagascar (Wilkinson et al., 2014). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, Angola, Botswana, Congo (Democratic Republic of the), Kenya, Morocco, Namibia, South Africa, Sudan, Tanzania, Togo, Zambia. Subfamily Kerivoulinae Miller, 1907 *1907. Kerivoulinae Miller, Bull. U.S. natl. Mus., 57: 232. Publication date: 29 June 1907. Comments: Type genus: Kerivoula Gray, 1842. Originally included the genera Kerivoula J. Gray, 1842 and Phoniscus Miller, 1905 (see Caron et al., 2018: 265). (Current Combination) TAXONOMY: Known genera in the Kerivoulinae: †Chamtwaria Butler, 1884; Kerivoula Gray, 1842; Phoniscus Miller, 1905. COMMON NAMES: Czech: vlnouškové, kerivuly. Bats, Woolly bats. English: Forest African Chiroptera Report 2020 649 †Genus Chamtwaria Butler, 1984 *1984. Chamtwaria Butler, Palaeovert., 14 (3): 117, 187. Publication date: 15 November 1984 [Goto Description]. (Current Combination) TAXONOMY: Was assigned to the Vespertilionidae by Beard et al. (1992). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe- Early Miocene (Burdigalian). Currently recognized species of the genus Chamtwaria: †pickfordi Butler, 1984. †Chamtwaria pickfordi Butler, 1984 *1984. Chamtwaria pickfordi Butler, Palaeovert., 14 (3): 118, 187, figs 25 A, A'. Publication date: 15 November 1984. Type locality: Kenya: Tinderet (volcano) Region, Songhor, Kapurtay agglomeration: Chamtwara [Goto Description]. - Etymology: Named after Dr. Martin Pickford. (Current Combination) PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Early Miocene (Burdigalian - Brown et al., 2019: Suppl.). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola. Genus Kerivoula Gray, 1842 *1842. Kerivoula Gray, Ann. Mag. nat. Hist., [ser. 1], 10 (65): 258. Publication date: 1 December 1842. - Comments: Type species: Vespertilio pictus Pallas, 1767. Gray (1842: 258) mentions Vesp. Hardwickii Horsfield, V. picta Pallas, V. tenuis, G. Gärtneri [and Kerivoula griseus, Kerivoula Poensis]. Type species by subsequent designation (Peters, 1867c) (see Corbet and Hill, 1992: 153). Allen (1939a: 100) mentioned Vespertilio hardwickii Horsfield as genotype. - Etymology: Rosevear suggested that Kerivoula was derived from the Cingalese name for the Painted Bat, kehel vuhla. Flannery (1995a: 369) indicates that it was derived from the Ceylonese for plantain (banana) bat. (Current Combination) 1849. Kirivoula Gervais, Dictionnaire Universelle d'Histoire naturelle, 13: 213. Publication date: 1849. - Comments: Emendation (see Meester et al., 1986: 63). (Emendation) 1861. Nyctophylax Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 42: 390 (for 1860). Publication date: 1861. - Comments: "Apparently based on Vespertilio tralatitius, Temminck, 1840 =V. tralatitius Horsfield, 1824, an indeterminable name according to Laurie and Hill (1954: 66). According to Palmer (1904), Nyctophylax was 'a new name for the barbaric Kerivoula Gray, 1842'" (see Corbet and Hill, 1992: 153). Mentioned as a possible synonym (question mark) by Pavlinov et al. (1995: 93). - Etymology: From the Greek "νύξ" or "νυκτός", meaning night and "φύλαξ", meaning watcher (see Palmer, 1904: 467). 1870. Nyctophilax: Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 62 (1): 544. Publication date: November 1870. (Lapsus) 1891. Cerivoula Blanford, The fauna of British India, including Ceylon and Burma. Part II, 338. Publication date: 1891. - Comments: Emendation (see Meester et al., 1986: 63). (Emendation) 1904. Kehelvoulha Jentink, Notes Leyden Mus., 24 (4): 175 (for 1902). Publication date: 15 Jul 1904. - Comments: Nomen nudum. Jentink (1904: 175) suggests that this should be the etymologically correct name, but adds immediately that this name has no taxonomical value whatsoever. 2005. Keivoula: Struebig, Rossiter, Bates, Kingston, Sai Sein Lin Oo, Aye Aye Nwe, Moe Moe Aung, Sein Sein Win and Khin Mya Mya, Acta Chiropt., 7 (1): 14. (Lapsus) ? Kerivoula sp.: 650 ISSN 1990-6471 TAXONOMY: Phoniscus Miller, 1906, has been included in Kerivoula as either a subgenus (Ellerman and Morrison-Scott, 1951; Ellerman et al., 1953) or a synonym (Ryan, 1965). Hill (1965) argues for the generic separation of Phoniscus from Kerivoula, and this view appears to be widely accepted (Rosevear, 1965; Hayman and Hill, 1971; Corbet and Hill, 1992), but see Koopman (1982). The species aerosa Tomes, 1858, was described from the east coast of South Africa, but Roberts (1951: 76) questions whether the description is not that of an immature argentata, although the colour is reminiscent of lanosa; Ellerman et al. (1953) point out that the type specimen resembles the Oriental species of Kerivoula (in which they include Phoniscus) rather than argentata or lanosa; Hill (1965: 553) has shown that aerosa should be transferred to the Asiatic Phoniscus. Hayman and Hill (1971) question the African origin of aerosa, and Corbet and Hill (1980) dubiously place it in Southeast Asia, while Koopman (1982) cite the provenance as 'possibly Africa'. In view of the considerable doubt about the status and origin of this species it is not treated as a member of the southern African fauna. Currently (Simmons and Cirranello, 2020) recognized species of the genus Kerivoula: africana Dobson, 1878; agnella Thomas, 1908 – Louisiade Arch., Woodlark and D’Entrecasteaux Isls (Papua New Guinea) (Simmons, 2005: 526); argentata Tomes, 1861; cuprosa Thomas, 1912; depressa Miller, 1906 – Indochina, India, Burma [=Myanmar]; dongduongana Tu, Hassanin, Furey and Csorba, 2018 – Vietnam; eriophora (Heuglin, 1877); flora Thomas, 1914 – Borneo, Lesser Sunda Isls, Bali, Sumbawa and Sumba (Indonesia), Vietnam and Thailand (Simmons, 2005: 526); furva Kuo, Soisook, Ho, Csorba, Wang and Rossiter, 2017 – Taiwan, India, Myanmar, China; hardwickii (Horsfield, 1824) – India and Sri Lanka, Burma, Laos, Cambodia, Vietnam, Thailand, China, western Malaysia, Borneo, Java, Sumatra, Nusa Penida, Mentawai Isls, Sulawesi, Bali, Lesser Sundas, Kangean Isl and Talaud Isl (Indonesia), Philippines (Simmons, 2005: 526); intermedia Hill and Francis, 1984 – Borneo, western Malaysia (Simmons, 2005: 527); kachinensis Bates, Struebig, Rossiter, Kingston, Oo and Mya, 2004 – Myanmar, Vietnam, Cambodia; krauensis Francis, Kingston and Zubaid, 2007 – Peninsular Malaysia; S Peninsular Thailand; lanosa (A. Smith, 1847); lenis Thomas, 1916 – northeastern and southern India, western Malaysia, Sabah (Simmons, 2005: 527); minuta Miller, 1898 – western Malaysia, southern Thailand, Borneo (Simmons, 2005: 527); muscina Tate, 1941 – central New Guinea (Simmons, 2005: 527); myrella Thomas, 1914 – Bismarck Arch. (Simmons, 2005: 527); papillosa (Temminck, 1840) – Thailand, Cambodia, Vietnam, western Malaysia, Sumatra, Java, Sulawesi, Borneo (Simmons, 2005: 527); pellucida (G.R. Waterhouse, 1845) – Borneo, Philippines, Java and Sumatra, western Malaysia (Simmons, 2005: 528); phalaena Thomas, 1912; picta (Pallas, 1767) – Sri Lanka; India and Nepal to Vietnam, western Malaysia and southern China, Borneo, Sumatra, Java, Bali, Lombok and Molucca Isls (Simmons, 2005: 528); smithii Thomas, 1880; titania Bates, Struebig, Hayes, Furey, Mya, Thong, Tien, Son, Harrison, Francis and Csorba, 2007 – Myanmar, Thailand, Laos, Cambodia, Vietnam; Taiwan; China (Hainan Island); whiteheadi Thomas, 1894 – Philippines, Borneo, southern Thailand, western Malaysia (Simmons, 2005: 528). COMMON NAMES: Czech: praví vlnouškové. English: Woolly Bats, Trumpet-eared Bats. French: Chauves-souris peintes. German: Wollfledermäuse. Italian: Cherìvoule. ECHOLOCATION: Keeley et al. (2018: 13) indicate that some Kerivoula species have fundamental frequencies up to 295 kHz and bandwidths up to 155 kHz, which they link to the absence of grooves in the external ear. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Côte d'Ivoire, Tanzania. Kerivoula africana Dobson, 1878 *1878. Kerivoula africana Dobson, Catalogue of the Chiroptera of the collection of the British Museum, p. xl, 331, ul 335. Publication date: June 1878. Type locality: Tanzania: Coast opposite Zanzibar Island [Goto Description]. - Comments: Type specimen: MNHN ???, ad ♂, alcoholic. (Current Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: vlnoušek východoafrický, netopýr vlnatý, kerivula africká. English: African Woolly Bat, African Chiroptera Report 2020 Tanzanian Woolly Bat. French: Kérivoule de Tanzanie, Chauve-souris peinte de Tanzanie. German: Tanzania-Wollfledermaus. CONSERVATION STATUS: Global Justification MacPhee and Flemming, 1999) considered this species to be extinct, followed by Simmons, 2005), but it was recently rediscovered, see (Burgess et al., 2000). Listed as Endangered (EN B2ab(iii) ver 3.1 (2001)) because its area of occupancy is probably less than 500 km 2, with all individuals in fewer than five locations, and the extent of its forest habitat is continuing to decline (Fahr and Jacobs, 2008; IUCN, 2009). Assessment History Global 2008: EN B2ab(iii) ver 3.1 (2001) (Fahr and Jacobs, 2008; IUCN, 2009). 2004: EN B2ab(iii) ver 3.1 (2001) (Fahr and Jacobs, 2004; IUCN, 2004). 1996: DD (Baillie and Groombridge, 1996). 1994: Ex (Groombridge, 1994). 1993: Ex (World Conservation Monitoring Centre, 1993). 1990: Ex (IUCN 1990). 1988: Ex (IUCN Conservation Monitoring Centre 1988). 651 needed into the distribution of this species to locate additional populations. GENERAL DISTRIBUTION: Endemic to eastern Tanzania, where it has only been recorded from Morogoro; the Genda Genda coastal forest; and the Tong'omba coastal forest. Native: Tanzania (Matschie, 1895; Swynnerton and Hayman, 1951; Cockle et al., 1998; Burgess et al., 2000). POPULATION: Structure and Density:- Very little is known about the population of this rarely recorded species (Fahr and Jacobs, 2008; IUCN, 2009). Trend:- 2008: Decreasing (Fahr and Jacobs, 2008; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Tanzania. Regional None known. MAJOR THREATS: It is threatened by the loss of coastal forests, largely through conversion of land to subsistence agriculture and harvesting of timber and firewood for local use (Fahr and Jacobs, 2008; IUCN, 2009). CONSERVATION ACTIONS: Fahr and Jacobs (2008) [in IUCN (2009)] report that it has been recorded from Morogoro (presumably within the Morogoro State Forest Reserve) and Genda Genda (possibly in a Forest Reserve). There is a need to prevent further loss of coastal forest habitat. Further studies are Figure 238. Distribution of Kerivoula africana Kerivoula argentata Tomes, 1861 *1861. Kerivoula argentata Tomes, Proc. zool. Soc. Lond., 1861, I: 31, 32. Publication date: May 1861. Type locality: Namibia: Ovamboland: Otjoro [=Otjihoro] [ca. 17 54 S 15 38 E] [Goto Description]. Holotype: [Unknown] ♀. Collection date: 1 December 1859. See Tomes (1861b: 33). Holotype: BMNH 1907.1.1.540: ♀. Collection date: 1 December 1859. Presented/Donated by: ?: Collector Unknown. - Comments: Ansell and Dowsett (1988: 44) indicate the type locality is probably Otjihoro, according to Shortridge (1934: 70). (Current Combination) 1865. Nycticejus nidicola Kirk, Proc. zool. Soc. Lond., 1864, III: 651. Publication date: May 1865. Type locality: Mozambique: Zambesi River: Shupanga [=Chupanga] [18 02 S 35 37 E] [Goto Description]. Syntype: BMNH 1864.1.9.37: ad. Collected by: Dr. Kirk. Syntype: BMNH 1864.1.9.38: ad. Collected by: Dr. Kirk. Syntype: BMNH 1864.1.9.39: ad. Collected by: Dr. Kirk. - Comments: (1 ad ♂ + 2 ad ♀♀). Considered a valid subspecies by Ansell and Dowsett (1988: 44. 652 ISSN 1990-6471 1924. ? ? Kerivoula nidicola zuluensis Roberts, Ann. Transv. Mus., 10: 61. Publication date: 31 January 1924. Type locality: South Africa: KwaZulu-Natal: White Umfolosi River [28 19 S 31 50 E] [Goto Description]. Holotype: TM 3025: ad ♂, skin and skull. Collected by: Austin Roberts; collection date: 17 July 1922. Note: In weaver nest with two others. Comments: Considered a valid subspecies by Taylor (1998: 57). Kerivoula argentata argentata: (Name Combination) Kerivoula argentata zuluensis: (Name Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Afrikaans: Damara-wolhaarvlermuis, Damaralandse Wolhaarvlermuis. Chinese: 银彩 蝠. Czech: vlnoušek damarský. English: Silvery Woolly Bat, Silvered Woolly Bat, Silver Woolly Bat, Damara Woolly Bat. French: Chauve-souris peinte argentée, Chauve-souris laineuse du Damara. German: Bunte Wollfledermaus. Kiluba (DRC): Kasusu. Portuguese: Morcego lanoso da Damaralandia. ETYMOLOGY OF COMMON NAME: This species was originally described from a specimen from Damaraland, Namibia (Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: K. argentata may be represented in the late Holocene at Nkupe, South Africa (Avery, 1991: 6). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008k; IUCN, 2009; Monadjem et al., 2017ch). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ch). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008k; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004h; IUCN, 2004). 1996: Lower Risk/least concern (Baillie and Groombridge, 1996). Regional South Africa:- 2016: NT C2a(i) ver 3.1 (2001) (Monadjem et al., 2016f). 2004: EN B1ab(iii)+2ab(iii) ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: There appear to be no major threats to this species as a whole (Jacobs et al., 2008k; IUCN, 2009; Monadjem et al., 2017ch). CONSERVATION ACTIONS: Jacobs et al. (2008k) [in IUCN (2009)] and Monadjem et al. (2017ch) report that there appear to be no direct conservation measures in place. It has been recorded from the Kambai Forest Reserve in Tanzania by Cunneyworth (1996b), and seems likely to be present in a number of protected areas within its range. Further studies are needed to determine if the species is truly present in Angola. GENERAL DISTRIBUTION: Kerivoula argentata is distributed in East Africa and southern Africa, with some records in southern parts of the Democratic Republic of the Congo and possibly northern Angola in Central Africa (records are uncertain from this country). In East Africa it has been recorded from Kenya and Tanzania in the north, through Zambia, Malawi and Mozambique. In southern Africa, it appears to be widespread in Zimbabwe, with additional scattered records from northeastern South Africa and central Namibia. For southern Africa, Cooper-Bohannon et al. (2016: "5") calculated a potential distribution area of 752,853 km2. Native: Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 563); Kenya (Aggundey and Schlitter, 1984); Malawi (Happold and Happold, 1997b: 822; Monadjem et al., 2010d: 563); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 563; Monadjem et al., 2010c: 383); Namibia (Monadjem et al., 2010d: 563); South Africa (Roberts, 1951; Smithers, 1983; Skinner and Smithers, 1990; Taylor, 2000; Skinner and Chimimba, 2005; Monadjem et al., 2010d: 563); Tanzania, United Republic of (Cunneyworth, 1996b); Zambia (Ansell, 1974; Monadjem et al., 2010d: 563); Zimbabwe (Smithers and Wilson, 1979; Monadjem et al., 2010d: 563). African Chiroptera Report 2020 Presence uncertain: Angola (Hill and Carter, 1941), although Taylor et al. (2018b: 63) predict it might occur in that country. ECHOLOCATION: See Taylor (1999b). Schoeman and Waddington (2011: 291) mention a peak frequency of 99 kHz and a duration of 1.7 msec for a specimen from the Mkuze Game Reserve, South Africa. 28 calls from the Okavango River Basin reported by Weier et al. (2020: Suppl.) had the following characteristics: Fmax: 87.09 ± 10.39 kHz, Fmin: 75.29 ± 9.80 kHz, Fknee: 77.89 ± 6.38 kHz, Fchar: 77.44 ± 6.73 kHz, slope: 7.86 ± 28.41 Sc, duration: 3.37 ± 1.01 msec. 653 Forest Reserve, Tanzania, at the end of November 2005. VIRUSES: Paramyxoviridae Mortlock et al. (2015: 1841) reported that the one specimen they examined from South Africa tested positive for Paramyxovirus sequences. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Congo (Democratic Republic of the), Kenya, Malawi, Mozambique, Namibia, Senegal, South Africa, Tanzania, Zambia, Zimbabwe. POPULATION: Structure and Density:- Although this species is rarely encountered, it is not thought to be especially uncommon (Jacobs et al., 2008k; IUCN, 2009; Monadjem et al., 2017ch). Trend:- 2016: Unknown (Monadjem et al., 2017ch). 2008: Unknown (Jacobs et al., 2008k; IUCN, 2009). REPRODUCTION AND ONTOGENY: Trentin and Rovero (2011: 51) reported two lactating females, captured at Uzungwa Scarp Figure 239. Distribution of Kerivoula argentata Kerivoula cuprosa Thomas, 1912 *1912. Kerivoula cuprosa Thomas, Ann. Mag. nat. Hist., ser. 8, 10 (55): 41. Publication date: 1 July 1912. Type locality: Cameroon: Ja River: Bitye [03 13 N 12 22 E, 2 000 ft] [Goto Description]. Holotype: BMNH 1912.10.25.1: ad ♂. Collected by: George Latimer Bates Esq. Collection date: 17 October 1911; original number: 564. (Current Combination) TAXONOMY: See Simmons (2005). COMMON NAMES: Czech: vlnoušek bronzový. English: Copper Woolly Bat. French: Chauve-souris laineuse cuivrée, Chauve-souris peinte cuivrée. German: Kupferfarbene Wollfledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of continuing problems with its taxonomy as well as absence of recent information on its extent of occurrence, ecological requirements and threats (Fahr, 2008b; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Fahr, 2008b; IUCN, 2009). 2004: NT ver 3.1 (2001) (Fahr, 2004m; IUCN, 2004). 1996: LR/nt (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are not well known. In parts of its range it is presumably threatened to some degree by habitat loss resulting from conversion of land to agricultural use, and harvesting of timber and firewood (Fahr, 2008b; IUCN, 2009). 654 ISSN 1990-6471 CONSERVATION ACTIONS: Fahr (2008b) [in IUCN (2009)] reports that jt is unclear if this species is present in any protected areas. Additional studies are needed into the taxonomy, distribution, natural history and threats to this little known bat. GENERAL DISTRIBUTION: Scattered records from West and Central Africa. It has apparently been reported from Guinea, Côte d'Ivoire, Liberia, possibly Ghana (see Koopman et al., 1995; Grubb et al., 1998), Cameroon and the Democratic Republic of the Congo (see Hayman and Hill, 1971). It is found up to 600 m asl. POPULATION: Structure and Density:- It appears to be a rare, or rarely recorded, species (Fahr, 2008b; IUCN, 2009). Trend:- 2008: Unknown (Fahr, 2008b; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo (Democratic Republic of the), Côte d'Ivoire. Grubb et al. (1998: 92) indicate that records west of Cameroon are erroneous and probably refer to a specimen from Oda, Ghana in the FMNH that was re-identified as K. lanosa muscilla. A record of this species from Kenya was based on a specimen subsequently reidentified as Kerivoula smithii (J. Fahr pers. Comm., in Simmons, 2005: 526; and also in Fahr (2008b) [in IUCN (2009)]). Native: Cameroon; Congo (The Democratic Republic of the); Côte d'Ivoire; Guinea (Fahr and Ebigbo, 2004: 69); Kenya (Harrison, 1957b); Liberia (Koopman, 1989b; Koopman et al., 1995); Uganda (Kityo et al., 2009b: 135). Presence uncertain: Ghana. Figure 240. Distribution of Kerivoula cuprosa Kerivoula eriophora (Heuglin, 1877) *1877. Nycticejus eriophorus Heuglin, Reise in Nordost Afrika, 2: 34. Publication date: 1877. Type locality: Ethiopia: Begemdir province: Belegaz Valley: Semian and Wogara, between [ca. 12 50 N 38 20 E] [Goto Description]. - Etymology: From the Greek "eriophora" meaning wool-bearing. ? Kerivoula eriophora: (Name Combination, Current Combination) TAXONOMY: May be conspecific with africana which it antedates; see Hayman and Hill (1971: 53) and Simmons (2005: 526). COMMON NAMES: Czech: vlnoušek habešský. English: Ethiopian Woolly Bat. French: Chauve-souris laineuse d'Ethiopie, Chauve-souris peinte d'Ethiopie. German: Heuglins Wollfledermaus. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of continuing problems with its taxonomy as well as absence of recent information on its extent of occurrence, ecological requirements and threats (Fahr, 2008c; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Fahr, 2008c; IUCN, 2009). 2004: DD ver 3.1 (2001) (Fahr, 2004p; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are not known (Fahr, 2008c; IUCN, 2009). CONSERVATION ACTIONS: Fahr (2008c) [in IUCN (2009)] reports that it is not known if the species is present in any protected areas. Further research is needed into the African Chiroptera Report 2020 655 systematic status, distribution, natural history and possible threats to this species. GENERAL DISTRIBUTION: Kerivoula eriophora has only been recorded at the type locality of 'between Semian and Wogara, Belegaz Valley, Ethiopia'. Native: Ethiopia (Heuglin, 1877: 34). POPULATION: Structure and Density:- It is known only from the holotype (Fahr, 2008c; IUCN, 2009). Trend:- 2008: Unknown (Fahr, 2008c; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: South Sudan. Figure 241. Distribution of Kerivoula eriophora Kerivoula lanosa (A. Smith, 1847) *1847. Vespertilio lanosus A. Smith, Illustrated Zoology of South African Mammals, pl. 50 and text. Publication date: December 1847. Type locality: South Africa: Cape province: Cape Town, 200 mi (332 km) E. Syntype: BMNH 1907.1.1.538: ♂. Syntype: BMNH 1907.1.1.539:. - Comments: Thorn et al. (2009: 50) indicate that Roberts (1951: 75) designated Knysna as type locality. - Etymology: From the Greek "lanosa" meaning woolly. 1878. Kerivoula brunnea Dobson, Catalogue of the Chiroptera of the collection of the British Museum, xl, 331, 334. Publication date: June 1878. Type locality: South Africa: "South Africa or Madras" [Goto Description]. Holotype: BMNH 1852.8.12.9: sad ♂, alcoholic (skull not removed). Presented/Donated by: Sir Andrew Smith. - Comments: Roberts (1951: 75) mentioned "Madras or South Africa as type locality. Sir A. Smith (Obviously the type of Smith's V. Lanosus, cf. Roberts, 1913: Knysna)". Meester et al. (1986: 64) mention "South Africa or Madras"). 1901. Kerivoula harrisoni Thomas, Proc. zool. Soc. Lond., 1900, IV: 802, footnote. Publication date: April 1901. Type locality: Ethiopia: between Lakes Zwai and Margherita [=Lake Abaya]: Walamo [c. 07 00 N 38 00 E,, 6 700 ft] [Goto Description]. Holotype: BMNH 1900.11.4.1: Collected by: J.J. Harrison; collection date: 21 February 1900. Presented/Donated by: J.J. Harrison. 1906. Kerivoula muscilla Thomas, Ann. Mag. nat. Hist., ser. 7, 18 (106): 294. Publication date: 1 October 1906. Type locality: Cameroon: Ja River [ca. 03 00 N 13 00 E] [Goto Description]. Holotype: BMNH 1908.6.23.10: ad ♂. Collected by: George Latimer Bates Esq. Collection date: 22 December 1905. Presented/Donated by: ?: Collector Unknown. - Comments: Considered a valid subspecies by Fahr and Ebigbo (2003: 128; 2004: 73). 1920. Kerivoula lucia Hinton, Ann. Mag. nat. Hist., ser. 9, 6 (32): 240. Publication date: 1 August 1920. Type locality: Zimbabwe: N'dola. [12 50 S 28 40 E] [Goto Description]. Holotype: BMNH 1920.11.3.27: ad ♂. Collected by: Captain Guy Chester Shortridge; collection date: 26 September 1919; original number: 472. 1922. Kerivoula lueia Kershaw, Ann. Mag. nat. Hist., ser. 9, 10 (55): 99. Publication date: 1 July 1922. - Comments: Erratum (see Meester et al., 1986: 64). 1959. Kerivoula harrisoni bellula Aellen, Archs Sci. Genève, 12 (2): 221. Type locality: Côte d'Ivoire: NW of Abidjan: Adiopodoumé [05 19 N 04 08 W, 0 - 30 m] [Goto Description]. Holotype: MHNG 965.038: ad ♂, alcoholic (skull not removed). Collected by: Dr. Villy Aellen; collection date: 20 July 1953; original number: 622. ? Kerivoula harrisoni lucia: (Name Combination) ? Kerivoula lanosa harrisoni: (Name Combination) ? Kerivoula lanosa lucia: (Name Combination) ? Kerivoula lanosa muscilla: (Name Combination) ? Kerivoula lanosa: (Name Combination, Current Combination) 656 ISSN 1990-6471 TAXONOMY: Includes harrisoni and muscilla; see Hill (1977b), but see Fahr and Ebigbo (2003: 128; 2004: 73). For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 636,522 km2. COMMON NAMES: Afrikaans: Klein wolhaarvlermuis. Chinese: 小彩 蝠 . Czech: vlnoušek malý. English: Lesser Woolly Bat. French: Petite Chauve-souris laineuse, Chauve-souris laineuse. German: Kleine Wollfledermaus. Portuguese: Morcego lamoso Harrison. Native: Angola (Monadjem et al., 2010d: 563; Taylor et al., 2018b: 63); Botswana (Monadjem et al., 2010d: 563); Central African Republic; Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 563); Côte d'Ivoire; Ethiopia (Kruskop and Lavrenchenko, 2009: 72 first record in over a 100 years); Gabon; Ghana; Guinea (Fahr and Ebigbo, 2003: 128); Kenya; Liberia; Malawi (Ansell and Dowsett, 1988; Monadjem et al., 2010d: 563); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 563; Monadjem et al., 2010c: 383); Nigeria (Happold, 1987); South Africa (Roberts, 1951; Smithers, 1983; Skinner and Smithers, 1990; Taylor, 1998; Skinner and Chimimba, 2005; Monadjem et al., 2010d: 563); Swaziland (Monadjem et al., 2010d: 563); Tanzania; Zambia (Ansell, 1978; Monadjem et al., 2010d: 563); Zimbabwe (Cotterill, 1996a; Monadjem et al., 2010d: 563). Presence uncertain: Namibia (Cotterill, 2004a: 262; Monadjem et al., 2010d: 563). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs et al., 2008l; IUCN, 2009; Monadjem et al., 2017ci). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017ci). 2008: LC ver 3.1 (2001) (Jacobs et al., 2008l; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs et al., 2004a; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016g). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MOLECULAR BIOLOGY: DNA - Unknown. Karyotype - Rautenbach et al. (1993) reported 2n = 28, FN = 50, BA = 24, a metacentric X chromosome, and an acrocentric Y chromosome, for specimens from South Africa. Protein / allozyme - Unknown. MAJOR THREATS: There appear to be no major threats to this species (Jacobs et al., 2008l; IUCN, 2009; Monadjem et al., 2017ci). ROOST: In the southern part of the Kruger Park, Pienaar (1964: 12) indicated that it is said that it often used disused nests of weaver birds, but he could not confirm this without doubt. CONSERVATION ACTIONS: Jacobs et al. (2008l) [in IUCN (2009)] and Monadjem et al. (2017ci) report that it is not known if the species is present in any protected areas. Further studies are needed to better understand the taxonomic status of Kerivoula lanosa. POPULATION: Structure and Density:- There appears to be little information available on the population abundance of this species (Jacobs et al., 2008l; IUCN, 2009; Monadjem et al., 2017ci). GENERAL DISTRIBUTION: Kerivoula lanosa has been recorded across subSaharan Africa, from Liberia and Guinea in the west, to Ethiopia and Kenya in the east, and ranging as far south as southern South Africa. In the latter country, its distribution is mainly affected by temperature seasonality (Babiker Salata, 2012: 49). Trend:- 2016: Unknown (Monadjem et al., 2017ci). 2008: Unknown (Jacobs et al., 2008l; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Botswana, Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Eswatini, Ethiopia, Gabon, Ghana, Guinea, Kenya, Liberia, Malawi, Mozambique, Namibia, Nigeria, South Africa, Tanzania, Zambia, Zimbabwe. African Chiroptera Report 2020 657 Figure 242. Distribution of Kerivoula lanosa Kerivoula phalaena Thomas, 1912 *1912. Kerivoula phalæna Thomas, Ann. Mag. nat. Hist., ser. 8, 10 (57): 281. Publication date: 1 September 1912. Type locality: Ghana: Western province: Inland from Denkwa: Bibianaha [=Bibiani] [06 28 N 02 20 W, 720 ft] [Goto Description]. Holotype: BMNH 1912.6.20.3: ad ♀. Collected by: Dr. H.G.F. Spurrell; collection date: 24 April 1912; original number: 224. Presented/Donated by: Dr. H.G.F. Spurrell. - Etymology: From the Greek "phalaena" meaning whale. (Current Combination) 1973. Kerivoula phalaema: Eisentraut, Bonn. zool. Monogr., 3: 46. (Lapsus) 2010. Kerivoula cf. phalaena: Monadjem, Schoeman, Reside, Pio, Stoffberg, Bayliss, Cotterill, Curran, Kopp and Taylor, Acta Chiropt., 12(2): 383. ? Kerivoula phalaena: (Current Spelling) TAXONOMY: See Simmons (2005: 528). COMMON NAMES: Czech: vlnoušek západoafrický. English: Spurrell's Woolly Bat, Spurell trumpet-eared bat. French: Chauve-souris à oreilles en trompette de Spurell, Chauve-souris peinte phalène, Chauvesouris à oreilles en trompette de Spurell. German: Spurrells Wollfledermaus. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Fahr, 2008d; Fahr, 2008d; Monadjem and Fahr, 2017a). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Fahr, 2017a). 2008: LC ver 3.1 (2001) (Fahr, 2008d; Fahr, 2008d). 2004: NT ver 3.1 (2001) (Fahr, 2004l; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The threats to this species are poorly known. Koopman et al. (1995) mention that this one of seven bat species from Liberia that have not been recorded from the lowland since 1965, and that it might have declined because of deforestation, although this may instead just indicate vagaries of collection within this country (Fahr, 2008d; IUCN, 2009; Monadjem and Fahr, 2017a). CONSERVATION ACTIONS: Fahr (2008d) [in IUCN (2009)] and Monadjem and Fahr (2017a) reports that it is not known if the species is present within any protected areas. Additional studies are needed into the distribution, abundance, natural history and threats to this little known species. GENERAL DISTRIBUTION: Recorded from Guinea, Liberia, Côte d'Ivoire and Ghana in West Africa; and from Cameroon, the Democratic Republic of the Congo to western Uganda in Central Africa. 658 ISSN 1990-6471 Native: Cameroon; Congo (The Democratic Republic of the); Côte d'Ivoire; Ghana; Guinea (Fahr and Ebigbo, 2004: 69); Liberia (Kuhn, 1965; Koopman et al., 1995; Rwanda; Uganda. Presence uncertain: Mozambique (Monadjem et al., 2010c: 383, recorded as K. cf. phalaena). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Gabon, Ghana, Liberia, Uganda. HABITAT: Monadjem et al. (2016y: 368) collected this bat in forested habitats in the Mount Nimba area, at altitudes between 450 and 700 m. POPULATION: Structure and Density:- It appears to be a rare species that is known only from 14 localities (Fahr, 2008d; IUCN, 2009; Monadjem and Fahr, 2017a). Trend:- 2016: Unknown (Monadjem and Fahr, 2017a). 2008: Unknown (Fahr, 2008d; IUCN, 2009). Figure 243. Distribution of Kerivoula phalaena Kerivoula smithii Thomas, 1880 *1880. Kerivoula Smithii Thomas, Ann. Mag. nat. Hist., ser. 5, 6 (32): 166, text-fig. Publication date: 1 August 1880. Type locality: Nigeria: Eastern region: Old Calabar [04 56 N 08 22 E]. Holotype: BMNH 1880.7.21.9: Collected by: Dr. A. Robb. Presented/Donated by: Dr. J.A. Smith. - Etymology: In honour of Dr. J.A. Smith, who presented to the type specimen to the British Museum of Natural History. (Current Combination) ? Kerivoula smithi: (Alternate Spelling) ? Kerivoula smithii: (Current Spelling) TAXONOMY: See Simmons (2005: 528). 2004a; IUCN, 2004). Groombridge, 1996). COMMON NAMES: Chinese: 史 密 斯 彩 蝠 . Czech: vlnoušek guinejský. English: Smith's Woolly Bat. French: Chauve-souris peinte de Smith. German: Smith's Wollfledermaus. Italian: Cherìvoula di Smith. Regional None known. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Fahr, 2008e; IUCN, 2009; Monadjem and Fahr, 2017b). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Fahr, 2017b). 2008: LC ver 3.1 (2001) (Fahr, 2008e; IUCN, 2009). 2004: LC ver 3.1 (2001) (Fahr, 1996: LR/nt (Baillie and MAJOR THREATS: There appear to be no major threats to this species (Fahr, 2008e; IUCN, 2009; Monadjem and Fahr, 2017b). CONSERVATION ACTIONS: Fahr (2008e) [in IUCN (2009)] and Monadjem and Fahr (2017b) reports that it is not known if the species is present within any protected areas. Further studies are needed into the distribution, abundance, and general ecology of this apparently rare species. GENERAL DISTRIBUTION: Kerivoula smithii is known from scattered records in a distribution band ranging from Nigeria and Cameroon in the west, through the Democratic Republic of the Congo, to Uganda and Kenya in African Chiroptera Report 2020 the east. It has been recorded at elevations of 900 to 2,800 m asl. Fahr (2007a: 105) indicates that the specimens from Liberia mentioned by Happold (1987: 349) and Koopman et al. (1995) are actually representatives of K. phalaena. J. Fahr pers. comm. [in Simmons (2005: 528)] also suggests that records from Côte d'Ivoire (Happold, 1987: 349) appear to be in error. 659 Trend:- 2016: Unknown (Monadjem and Fahr, 2017b). 2008: Unknown (Fahr, 2008e; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Central African Republic, Congo (Democratic Republic of the), Kenya, Nigeria, Tanzania, Uganda. Native: Cameroon; Congo (The Democratic Republic of the) (Hayman et al., 1966); Kenya (Harrison, 1963c; Aggundey and Schlitter, 1984) ; Nigeria (Happold, 1987: 77); Uganda. Presence uncertain: Côte d'Ivoire (Happold, 1987: 349). ECHOLOCATION: Eisenring et al. (2016: SI 2) reported the following values for 14 calls from the Aberdares Range in Kenya: PF: 95.9 ± 6.2 (81.5 - 104.9), HF: 126.3 ± 7.2 (115.6 - 137.2), LF: 65.3 ± 13.2 (44.9 - 91.8), DT: 0.0 ± 0.0 (0.0 - 0.1), DF: 61.0 ± 16.5 (25.1 85.5), IPI: 0.2 ± 0.1 (0.1 - 0.6). POPULATION: Structure and Density:- This species has been very rarely recorded (Happold, 1987: 77; Fahr, 2008e; IUCN, 2009; Monadjem and Fahr, 2017b). Figure 244. Distribution of Kerivoula smithii Subfamily Myotinae Tate, 1942 *1942. Myotinae Tate, Bull. Am. Mus. Nat. Hist., 80 (7): 221, 229. Publication date: 27 November 1942. - Comments: Type genus: Myotis Kaup, 1829. (Current Combination) 1942. Myotini Tate, Bull. Am. Mus. Nat. Hist., 80 (7): 221, 229. Publication date: 27 November 1942. - Comments: Type genus: Myotis Kaup, 1829. Introduced as tribe, and originally included the genera Myotis Kaup, 1829; Lasionycteris Peters, 1865; Plecotus É. Geoffroy, 1818; Corynorhinus H. Allen, 1865; Idionycteris Anthony, 1923; and Euderma H. Allen, 1891 (see Jackson and Groves, 2015: 280). 1987. Leuconoformes Menu, Palaeovert., 17 (3): 77, 6592, 133. Publication date: 31 March 1987. - Comments: Type genus: Leuconöe Boie, 1830. Introduced with unclear rank, and originally included the genera Leuconöe Boie, 1830 [= Myotis Kaup, 1829]; Pizonyx Miller, 1906 [= Myotis Kaup, 1829]; and Perimyotis Menu, 1984 (see Jackson and Groves, 2015: 280). 1998. Myotinae: Simmons, in: Kunz, T. H.; Racey, P. A., A reappraisal of interfamilial relationships of bats., 12. - Comments: Type genus: Myotis Kaup, 1829. 2005. Myotiinae: Ramírez-Pulido, Arroyo-Cabrales and Castro-Campillo, Acta Zool. Mex. (ns), 21 (1): 31. - Comments: Ramírez-Pulido et al. (2005) refer to Simmons (1998) for Myotiinae, but Simmons (1998: 5) uses the correct spelling (Myotinae). (Lapsus) TAXONOMY: Tate (1942: 229) originally named Myotini as a tribe within Vespertilioninae, in which he included the genera Myotis, Lasionycteris, Plecotus, Corynorhinus, Idionycteris and Euderma. Although Tate (1942: 230) noted that the latter four genera might alternatively be referred to their own tribe Plecotini. This suggestion was followed by subsequent authors, and Myotini has been restricted to Myotis and Lasionycteris in recent classifications (e.g. Hill and Harrison, 1987; Koopman, 1994). Volleth and Heller (1994) described karyological data which indicate that Myotini represents a lineage distinct from Vespertilioninae, and therefore they suggested that Myotini be raised to the rank of subfamily. 660 ISSN 1990-6471 Despite their suggestion, Volleth and Heller (1994) did not use the name Myotinae in their publication, so formal usage of the name at the subfamily rank dates from Simmons (1998: 3, 5), who raised it to the subfamily level, and followed in recent classifications (e.g., Bannikova et al., 2002: 722; Hoofer and Van Den Bussche, 2003: 21). May not be monophyletlic; see Hoofer and Van Den Bussche (2001), and Simmons (2005). Hoofer and Van Den Bussche (2003: 21) and Roehrs et al. (2010: 1081) only include the members of the genus Myotis, (not Lasionycteris nor Cistugo). Known genera of the subfamily Myotinae: †Khonsunycteris Gunnell, Simons and Seiffert, 2008; Myotis Kaup, 1829. COMMON NAMES: Czech: netopýrcové. Nichnytsi] Ukrainian: Нічниці [= PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Ruedi et al. (2013: 446-447) doubt the African origin of the Myotinae as proposed by Gunnell et al. (2012), and their molecular phylogeny points to eastern Asia as the cradle of Myotis evolution. Gunnell et al. (2017: 14) suggest that the basal myotines (e.g. Khonsunycteris) originated in northern Africa in the Late Eocene, followed by the appearance of Myotis in the early Oligocene of Europe. The fact that they found a plecotine bat (Quinetia misonnei) in the same Belgian location, furthermore suggests that the vespertilionid subfamilies Vespertilioninae and Myotinae had already diverged by 33.5 MYA. The discovery of the southernmost and oldest record of Myotis podlesicensis Kowalski, 1956 in Europe (Spain; Late Miocene) by Crespo et al. (2017: 324) provides support for an African origin of this taxon. †Genus Khonsunycteris Gunnell, Simons and Seiffert, 2008 *2008. Khonsunycteris Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 3, 8. - Comments: Type species - Khonsunycteris aegypticus Gunnell, Simons and Seiffert, 2008. - Etymology: Khonsu, Egyptian God of the Moon, in reference to the nocturnal habits of vesper bats (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). Gunnell et al. (2017: 9) indicate that Khonsunycteris might be the earliest known myotine bat. SIMILAR SPECIES: See Gunnell et al. (2008). GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). †Khonsunycteris aegypticus Gunnell, Simons and Seiffert, 2008 *2008. Khonsunycteris aegypticus Gunnell, Simons and Seiffert, J. Vert. Paleont., 28 (1): 3, 8, fig. 8. Type locality: Egypt: Fayum Depression: Quarry L-41: Lower Sequence, Jebel Qatrani Formation. Holotype: CGM 83673: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Left dentary c1-m2. - Comments: latest Eocene, Priabonian. - Etymology: Name for Egypt (Gunnell et al., 2008). (Current Combination) TAXONOMY: See Gunnell et al. (2008). SIMILAR SPECIES: See Gunnell et al. (2008). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: See Gunnell et al. (2008). Timeframe: Late Eocene (Priabonian - 37.97 - 33.23 - Brown et al., 2019: Suppl.). GENERAL DISTRIBUTION: Egypt (Gunnell et al., 2008). African Chiroptera Report 2020 661 GENERAL DESCRIPTION OF CRANIAL AND DENTAL MORPHOLOGY: See Gunnell et al. (2008). Genus Myotis Kaup, 1829 *1829. Myotis Kaup, Skizzierte Entwicklungs-Geschichte Natürliche Systematik der Europäische Thierwelt, 1: 106. Publication date: 1829. - Comments: Type species: Vespertilio myotis Borkhausen, 1797, by subsequent designation by Miller (1912: 167). Kaup (1829: 106) mentioned Vespertilio murinus, but for a discussion on the type species, see Jackson and Groves (2015: 280). - Etymology: From the Greek "μύξ" or "μυόξ", meaning mouse and "ούς" or "ώτός" meaning ear (see Palmer, 1904: 442). Gardner (2005:184) considers the name to be masculine (My = mouse + ot = ear(ed) + is (Latin third declension ending)). 1829. Nystactes Kaup, Skizzirte Entwicklungs-Geschichte Natürliche Systematik der Europäische Thierwelt, 1: 108. Publication date: 1829. - Comments: Type species: Vespertilio bechsteini Kuhl, 1819. Jackson and Groves (2015: 281) indicate that this name is an objective synonym of Paramyotis Bianchi. Not Nystactes Gloger, 1827 (Aves, Piciformes, Bucconidae). - Etymology: From the Greek "νυστακτής", meaning the one who nods, a sleeper (see Palmer, 1904: 467). 1830. Leuconoe Boie, Isis oder Encyclopädische Zeitung von Oken, 256. Publication date: 1830. - Comments: Type species: Vespertilio daubentonii Kuhl, 1817. Considered a valid genus based on tooth morphology: see Menu (1985, 1987) in Menu et al. (2002: 320). 1841. Capaccinius Bonaparte, Iconografia Fauna italiana, 1: Indice Distributivo: 1. - Comments: Type species: Vespertilio megapodius Temminck, 1840 [=Vespertilio capaccinii Bonaparte, 1837], by tautonomy (see Jackson and Groves, 2015: 281). - Etymology: In honour of Francesco Capaccini, under secretary of state of Rome about 1833-34, and a patron of Bonaparte's "Iconografia della Fauna Italica," published in 1832-1841 (see Palmer, 1904: 49). 1841. Selysius Bonaparte, Iconografia Fauna italiana, 1: Mamm., Introd.: 3. Publication date: 1841 [Goto Description]. - Comments: Type species: Vespertilio mystacinus Kuhl, 1817, by monotypy (see Jackson and Groves, 2015: 281). - Etymology: In honour of Baron Edmond de Selys-Longchamps, 1813 - 1900, an eminent naturalist and statesman, some time president of the Belgian Senate; author of "Etudes de Micromammalogie," 1839, and "Faune Belge," 1844 (see Palmer, 1904: 51). 1842. Trilatitus Gray, Ann. Mag. nat. Hist., [ser. 1], 10 (65): 258. Publication date: 1 December 1842. - Comments: type species: Vespertilio hasseltii Temminck, 1840, by subsequent designation (see Jackson and Groves, 2015: 281). Included Vespertilio hasselti Temminck, M. macellus Temminck, and V. blepotis Temminck (a Miniopterus) (see Hall and Kelson, 1959: 159; Hall, 1981: 183; Corbet and Hill, 1992: 119). 1843. Vespertilio Gray, Cat. specs mamm., xix, 26. Publication date: 13 May 1843. Comments: Type species: Vespertilio mystacinus Kuhl, 1817, by monotypy according to Jackson and Groves (2015: 281). Not Linnaeus, 1758, nor Vespertilio Mörch, 1852 (Mollusca, Neogastropoda, Volutidae) (see Jackson and Groves, 2015: 281). 1849. Tralatitus: Gervais, Dictionnaire Universelle d'Histoire naturelle, 13: 213. - Comments: A lapsus for Trilatitus (see Hall, 1981: 183). Modification of Trilatitus (see Ellerman and Morrison-Scott, 1951: 137; Meester et al., 1986: 47). Incorrect subsequent spelling (see Jackson and Groves, 2015: 281). (Lapsus) 1856. Brachyotus Kolenati, Allg. deutsche naturh. Zeitung, N.F., 2 (2): 131. Publication date: 1856. - Comments: Type species: Vespertilio mystacinus Kuhl, 1817, by subsequent designation by Ellerman and Morrison-Scott (1951: 137) (see Jackson and Groves, 2015: 281). Preoccupied by Brachyotus Gould, 1837 (Aves, Strigiformes, Strigidae) (see Allen, 1939a: 89; Hall, 1981: 183; Corbet and Hill, 1992: 119; Pavlinov et al., 1995: 95; Jackson and Groves, 2015: 281). Based on V. mystacinus, daubentonii and dascyneme (see Allen, 1939a: 89). 1856. Isotus Kolenati, Allg. deutsche naturh. Zeitung, N.F., 2 (2): 131. Publication date: 1856. Comments: Type species: Vespertilio nattereri Kuhl, 1817. Considered a valid subgenus by Pavlinov et al. (1995: 95). - Etymology: From the Greek "ίσος", meaning equal and "ούς" or "ώτός", meaning ear (see Palmer, 1904: 354). 662 ISSN 1990-6471 1856. Myotus: Kolenati, Allg. deutsche naturh. Zeitung, N.F., 2 (2): 131. Publication date: 1856. - Comments: Incorrect subsequent spelling (see Palmer, 1904: 442; Jackson and Groves, 2015: 282). (Lapsus) 1866. Tralatitius: Gray, Ann. Mag. nat. Hist., ser. 3, 17 (98): 90. Publication date: 1 February 1866. - Comments: Lapsus for Trilatitus Gray (seeHall, 1981: 183). Modification of Trilatitus (see Ellerman and Morrison-Scott, 1951: 137; Meester et al., 1986: 47). (Lapsus) 1867. Vespertilio (Pternopterus) Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 706. Publication date: 1867. - Comments: Type species: Vespertilio (Pternopterus) lobipes Peters, 1867 (=Vespertilio muricola Gray, 1846). 1868. Pternopterus Peters, Monatsb. k. preuss. Akad. Wiss. Berlin, 1867: 706. - Comments: Type species: Vespertilio (Pternopterus) lobipes Peters, 1867 [= Myotis muricola (J. Gray, 1846)] by monotypy (see Jackson and Groves, 2015: 282). 1870. Aëorestes Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 62 (1): 427. Publication date: October 1870. - Comments: Based upon Vespertilio villosissimus E. Geoffroy, 1806, albescens E. Geoffroy, 1806, laevis I. Geoffroy, 1824 and nigricans Schinz, 1821 (see Allen, 1939a: 90; Hall, 1981: 183 [who excludes laevis]). Hill (in litt. In Meester et al., 1986: 48) pointed out that villosissimus is a member of Lasiurus Gray, 1831, but all others are indeed Myotis representatives. If villosissimus can be regarded as the type species of Aeorestes, it should instead be included in Lasiurus (Meester et al., 1986: 48). Hoofer and Van Den Bussche (2003: 25) consider Aeorestes a valid subgenus, including all of the New World species. 1870. Comastes Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 62 (1) 9: 565. Publication date: 1870. - Comments: Type species: Vespertilio capaccinii Bonaparte, 1837, by subsequent designation (see Jackson and Groves, 2015: 282). Based upon Vespertilio capaccinii Bonaparte, megapodius, dasycneme Boie, limnophilus (see Allen, 1939a: 90). Based upon Vespertilio capaccinii and dasycneme according to Hall and Kelson (1959: 159) and Hall (1981: 183). Not Comastes Jan, 1863 (Reptilia, Squamata, Colubridae) (see Jackson and Groves, 2015: 282). - Etymology: From the Greek "κωμαστής", meaning reveler, probably referring to the animal's nocturnal habits (see Palmer, 1904: 197). 1870. Exochurus Fitzinger, Sber. k. Akad. Wiss. Wien, math. naturw. Kl., 62 (1) 6: 75. Publication date: 1870. - Comments: Type species: Vespertilio macrodactylus Temminck, 1840, by subsequent designation (see Jackson and Groves, 2015: 282). Not of Kolenati (1858). Based upon Vespertilio macrodactylus Temminck, V. horsfieldii Temminck, and V. macrotarsus Waterhouse (see Hall, 1981: 183; Corbet and Hill, 1992: 119). Not of Kolenati, 1858: see Cabrera (1914: 91). - Etymology: From the Greek "εξοχος", meaning standing out and "ούρά", meaning tail (see Palmer, 1904: 284). 1899. Euvespertilio Acloque, Faune de France, Mammifères, 38. - Comments: Includes Vespertilio emarginatus Geoffroy, murinus Schreber =myotis Borkhausen, mystacinus Kuhl, nattereri Kuhl, and bechsteinii Kuhl. Hall and Kelson (1959: 159) and Hall (1981: 183) state that Euvespertilio includes emarginatus and murinus in a composite sense. 1906. Pizonyx Miller, Proc. Biol. Soc. Wash., 19: 85. Publication date: 4 June 1906. Comments: Type species: Myotis vivesi Menegaux, 1901, by original designation (see Jackson and Groves, 2015: 282). Considered a valid genus by Hall and Kelson (1959), but only as a subgenus of Myotis by Hall (1981: 183). *1910. Chrysopteron Jentink, Notes Leyden Mus., 32: 74. Publication date: January 1910 [Goto Description]. - Comments: Type species: Chryopteron bartelsii Jentink, 1910 [=Vespertilio formosa Hodgson, 1835]. Considered a valid subgenus by Hall and Kelson (1959: 159), Hall (1981: 183), Corbet and Hill (1992: 121) and Pavlinov et al. (1995: 95). 1917. Dichromyotis Bianchi, Ann. Mus. zool. Acad. Impér. Sci. St. Petersbourg, 21: lxxviii (for 2016). Publication date: 14 May - 13 Jun 1917. - Comments: Type species: Vespertilio formosus Hodgson, 1835, by original designation according to Csorba et al. (2014: 668), by monotypy according to Jackson and Groves (2015: 283). 1917. Megapipistrellus Bianchi, Ann. Mus. zool. Acad. Impér. Sci. St. Petersbourg, 21: lxxvii (for 2016). Publication date: 14 May - 13 June 1917. - Comments: Type species: Pipistrellus annectans Dobson, 1871, by monotypy (see Jackson and Groves, 2015: 282). Considered a valid subgenus by Pavlinov et al. (1995: 95). 1917. Paramyotis Bianchi, Ann. Mus. zool. Acad. Impér. Sci. St. Petersbourg, 21: lxxix (for 2016). Publication date: 14 May - 13 Jun 1917. - Comments: Type species: Vespertilio bechsteinii African Chiroptera Report 2020 1917. 1929. 1934. 1958. 1982. 1995. 1995. 1995. 1996. ? ? 663 Kuhl, 1817. New name for Nystactes Kaup, 1829, preoccupied (see Hall and Kelson, 1959: 159; Hall, 1981: 183). Considered a valid subgenus by Pavlinov et al. (1995: 95). Rickettia Bianchi, Ann. Mus. zool. Acad. Impér. Sci. St. Petersbourg, 21: lxxvii (for 2016). Publication date: 14 May - 13 Jun 1917. - Comments: Type species: Vespertilio (Leuconoe) ricketti Thomas, 1894, by monotypy (see Jackson and Groves, 2015: 282). Valid as a subgenus (see Hall and Kelson, 1959: 159; Hall, 1981: 183). Anamygdon Troughton, Rec. Austr. Mus., 17 (2): 87. Publication date: 26 Jun 1929. Comments: Type species: Anamygdon solomonis Troughton, by original designation (see Jackson and Groves, 2015: 283) [=Vespertilio adversus Horsfield, 1824 or Leuconoe moluccarum Thomas, 1915 ?]. Brachyotis: Taylor, Monographs of the Bureau of Science, Manila, 30: 278. (Lapsus) Hesperomyotis Cabrera, Rev. Mus. Arg. Cienc. Nat., Zool., 4: 103. - Comments: Type species: Myotis simus Thomas, 1901, by original designation (see Jackson and Groves, 2015: 283). Myottis: Alberico and Orejuela, Cespedesia, 3 (suppl.): 103. (Lapsus) Capaccinus: Pavlinov, Borissenko, Kruskop and Jahonton, Arch. Zool. Mus., Moscow State Univ., 133: 95. (Lapsus) Exochirus: Pavlinov, Borissenko, Kruskop and Jahonton, Arch. Zool. Mus., Moscow State Univ., 133: 95. (Lapsus) Selisius: Pavlinov, Borissenko, Kruskop and Jahonton, Arch. Zool. Mus., Moscow State Univ., 133: 99. (Lapsus) Myotiz: Spiewak, Johansson and Wüthrich, Allergologie, 19 (11): 510 - 511. (Lapsus) Loeconoe: (Alternate Spelling) Myotis sp.: TAXONOMY: Woodman (1993) indicated that the genus name ending on -otis" is female, and therefore the species name should also be female. Benda and Tsytsulina (2000: 335) agree with Woodman (1993), but continue to understand Myotis as a masculine name, because it has been used for over 170 years, and hence was conserved in this way. Simmons (2005) follows Pritchard (1994) and retains the masculin names too. See also Gardner (2005: 184) where Myotis = My (mouse) + ot (ear[ed]) + is (Latin third declension ending) = masculine (Mammalia). Bronner et al. (2003) state that the genus Myotis has been considered to include several subgenera, but the number thereof, and placement of species, varied considerably. Meester et al. (1986) listed three subgenera from the Southern African subregion, namely Cistugo Thomas, 1912 (for seabrai and lesueuri), Chrysopteron Jentink, 1910 (for welwitschii) and Selysius Bonaparte, 1841 (for tricolor and bocagei). Menu (1987) proposed synonymising Selysius under the subgenus Leuconoe Boie, 1830 on the basis of dental characters. This was endorsed by Koopman (1993a, 1994) who transferred M. bocagei to Leuconoe but included other Selysius and Chrysopteron in the subgenus Myotis, and also by Godawa-Stormark (1998)'s phenetic analysis of dental variation in the genus. Phylogenetic analysis of mitochondrial DNA sequences, however, do not support the monophyly of the three subgenera (Selysius, Leuconoe and Myotis) analysed (Ruedi and Mayer, 2001), but Cistugo warrants generic separation (N. Simmons in litt.) owing to the distinct wing glands and unique karyotypes (see Rautenbach et al., 1993) found in seabrai and lesueuri from southern Africa. Meester et al. (1986) state that their synonymy is slightly modified from Ellerman and Morrison-Scott (1951), who include also Aeorestes Fitzinger, 1870, a South American group, in the synonymy of Myotis, as does Hall (1981). However, J.E. Hill points out (in litt) that of the species included in this genus, the first, Vespertilio villosissimus E. Geoffroy, 1806, is a Lasiurus Gray, 1831, although the others, albescens E. Geoffroy, 1806, nigricans Schinz, 1821, and levis I. Geoffroy, 1824, belong in Myotis. If, as Hill suggests, villosissimus can be regarded as the type species of Aeorestes, it should instead be included in Lasiurus. Phylogenetic analyses performed by Ruedi et al. (2013: 441) confirm that the Ethiopian clade within Myotis is highly supported with all methods of reconstruction (see also Stadelmann et al., 2004b). This clade contains all of the subSaharan species, the species inhabiting the islands in the Western Indian Ocean, as well as M. emarginatus (which occurs circum-Mediterranean) and two Asian forms (M. cf. formosus and M. formosus flavus). Patterson et al. (2019: "4') made an extensive phylogenetic analysis on the Afrotropical Myotis 664 ISSN 1990-6471 species and found that they were not monophyletic as M. welwitschii formed a sister clade with the east Asian M. rufoniger, which was separate from all other African species. All African species, together with M. rufoniger, M. formosus and M. emarginatus appeared to be grouping on a higher level. Ghazali et al. (2016: 475) state that phylogenetic relationships within Myotis reflect biogeographic affinities rather than phenotypes. They also suggest that Myotis diverged from other bats in the early Miocene, with a subsequent split between Old and New World lineages about 19 MYA, and that ecomorphs ('Leuconoe' [near-water hunters], 'Myotis' [gleaners], 'Selysius' [aerial hawkers]) emerged independently in different lineages of Myotis. Their analyses indicate the ancestor of the African (Ethiopian) clade was most likely a Myotis, which diverged from the Eurasian clade about 17.0 MYA. Depending on the calibration, Morales et al. (2019: 2271) estimated this last split to have occurred at either 27 ± 3.5 mya or at 17.8 ± 3.5 mya. Bogdanowicz et al. (2015: Fig. 2, p. 640) mention Moroccan data for Myotis nattereri, but in the supplementary information, these are mentioned as Myotis cf. nattereri. They also include the Mediterranean coastal areas of Morocco and Algeria in the distribution area of M. capaccinii. Currently (Simmons and Cirranello, 2020) recognized species of the genus Myotis: adversus (Horsfield, 1824) – islands in Indonesia, New South Wales, Taiwan, possibly Vietnam and peninsular Malaysia (Simmons, 2005: 500); albescens (E. Geoffroy Saint-Hilaire, 1806) – southern Veracruz (Mexico), Guatemala, Honduras, Nicaragua, Panama, Colombia, Venezuela, Guyana, Surinam, Equador, Peru, Brazil, Uruguay, northern Argentina, Paraguay and Bolivia (Simmons, 2005: 501); alcathoe von Helversen and Heller, 2001 – Greece, Hungary, France (Simmons, 2005: 501); altarium Thomas, 1911 – Szechwan, Kweichow (China), Thailand (Simmons, 2005: 501); ancricola Kruskop, Borisenko, Dudorova and Artyushin, 2018 – Only known from the type locality and the Ngoc Ling mountains in Vietnam; anjouanensis Dorst, 1960; annamiticus Kruskop and Tsytsulina, 2001 – Vietnam (Simmons, 2005: 501); annatessae Kruskop and Borisenko, 2013 – Vietnam; Central Laos; annectans (Dobson, 1871) – northeastern India to Burma, Thailand, Laos, Cambodia and Vietnam (Simmons, 2005: 502); atacamensis (Lataste, 1892) – southern Peru, northern Chile (Simmons, 2005: 502); ater (Peters, 1866) – Vietnam, western Sumatra, Peninsular Malaysia, Sulawesi, Togian Isl, northern Borneo, Moluccas, Papua New Guinea (Simmons, 2005: 502); attenboroughi Moratelli, Wilson, Novaes, Helgen and Gutiérrez, 2017 – Tobago Island; auriculus Baker and Stains, 1955 – Arizona and New Mexico (USA) to Jalisco and Veracruz (Mexico); Guatemala (Simmons, 2005: 502); australis Dobson, 1878) – New South Wales (Simmons, 2005: 502); austroriparius (Rhoads, 1897) – southeastern USA including Florida, north to Indiana and North Carolina, west to Texas and southeastern Oklahoma (Simmons, 2005: 502); badius Tiunov, Kruskop, and Feng, 2011 – China; bakeri Moratelli, Novaes, Bonilla and Wilson, 2019 – Known only from two localities in the departments of Lima and Lambayeque, Peru; bartelsi (Jentink, 1910) – Indonesia (Java and Bali); bechsteinii (Kuhl, 1817) – Europe to Caucasus and Iran; Bulgaria; England; southern Sweden (Simmons, 2005: 503); blythii (Tomes, 1857) – Turkey and Israel to Iraq and Iran; northwestern India and the Himalayas; northwestern Altai Mtns; Inner Mongolia and Shensi (China) (Simmons, 2005: 503); bocagii (Peters, 1870); bombinus Thomas, 1906 – Japan, Korea, southeastern Siberia, northeastern China (Simmons, 2005: 503); borneoensis Hill and Francis, 1984 – Sabah (Borneo); brandtii (Eversmann, 1845) – Britain south to Italy, Greece and Bulgaria, east to Kazakhstan and Mongolia, eastern Siberia, Kamchatka Peninsula and Kurile Isls; Ussuri region (Russia), Korea (Simmons, 2005: 503); bucharensis Kuzyakin, 1950 – Uzbekistan, Tajikistan and Afghanistan (Simmons, 2005: 504); californicus (Audubon and Bachman, 1842) – southern Alaska Panhandle (USA) to Baja California and higher elevations in the Sonoran and Chihuahuan deserts (Mexico); Guatemala (Simmons, 2005: 504); capaccinii (Bonaparte, 1837); caucensis J.A. Allen, 1914 –occurs along the Andes of Colombia, Ecuador, and Peru, including intermontane valleys and adjacent Amazon lowlands of those countries; chiloensis (G.R. Waterhouse, 1840) – central and southern Chile; Argentina (Simmons, 2005: 504); chinensis (Tomes, 1857) – Szechwan and Yunnan to Kiangsu (China); Hong Kong; northern Thailand; Burma; Vietnam (Simmons, 2005: 505); ciliolabrum (Merriam, 1886) – southern Alberta and Saskatchewan (Canada) south through eastern Colorado and western Kansas (USA) (Simmons, 2005: 505); clydejonesi Moratelli, Wilson, Gardner, Fisher and Gutiérrez, 2016 – Only known from the type specimen from Suriname; cobanensis Goodwin, 1955 – central Guatemala (Simmons, 2005: 505); crypticus Ruedi, Ibáñez, Salicini, Juste and Puechmaille, 2019 – Mountain areas of N + C Spain, S France, Italy, SW Austria?, W Switzerland, SE France; csorbai Topál, 1997 – Nepal (Simmons, 2005: 505); dasycneme (Boie, 1825) – France and African Chiroptera Report 2020 Sweden east to Yenisei River (Russia), south to Ukraine, northwestern Kazakhstan, China (Simmons, 2005: 505); daubentonii (Kuhl, 1817) – Europe (including Britain and Ireland; Scandinavia) east to Kamtschatka, Vladivostok, Sakhalin and Kurile Isls (Russia), Japan, Korea, Manchuria, northern and eastern China (including Tibet), Vietnam (Simmons, 2005: 505); davidii (Peters, 1869) – northern China (Simmons, 2005: 506); dieteri M. Happold, 2005; diminutus Moratelli and Wilson, 2010 – Ecuador, Colombia; dinellii Thomas, 1902 – Bolivia, Uruguay, Argentina (incl. Entre dos Rios; Patagonia); Brazil; dominicensis Miller, 1902 – Dominica, Guadeloupe (Simmons, 2005: 506); elegans Hall, 1962 – Mexico to Costa Rica (Simmons, 2005: 506); emarginatus (E. Geoffroy Saint-Hilaire, 1806); escalerai Cabrera, 1904 – France, Spain, Portugal; evotis (H. Allen, 1864) – southern British Columbia, southern Alberta, southern Saskatchewan (Canada) to New Mexico (USA) and Baja California (Mexico) (Simmons, 2005: 506); federatus Thomas, 1916 – Peninsular Malaysia; fimbriatus (Peters, 1871) – southeastern China (Simmons, 2005: 507); findleyi Bogan, 1978 – Trés Marías Isls (Mexico) (Simmons, 2005: 507); formosus (Hodgson, 1835) – Afghanistan to northern India, Nepal, Tibet, Kweichow, Kwangsi, Kiangsu and Fukien (China), Taiwan, Korea, Tsushima Isl (Japan), Malaysia, Philippines, Sumatra, Java, Sulawesi and Bali (Simmons, 2005: 507); fortidens Miller and Allen, 1928 – Sonora and Veracruz (Mexico) to Guatemala (Simmons, 2005: 507); frater G. M. Allen, 1923 – eastern Siberia, Ussuri Region, Krasnoyarsk Region (Russia) to Korea, Heilungkiang (China), southeastern China; Japan (Simmons, 2005: 507); gomantongensis Francis and Hill, 1998 – Sabah (Borneo, Malaysia) (Simmons, 2005: 508); goudoti (A.Smith, 1834); gracilis Ognev, 1927 – E Palearctic, ranging from Lake Baikal region to Ussuri region, Sakhalin Isl., Kamchatka Peninsula, Kurille Isls., Amur region, Korea, and Hokkaido (Japan); grisescens A. H. Howell, 1909 – Florida Panhandle to Kentucky, Indiana, Illinois, eastern Kansas and northeastern Oklahoma (USA) (Simmons, 2005: 508); handleyi Moratelli, Gardner, de Oliveira and Wilson, 2013 – Known from two cordilleras in northern Venezuela; hasseltii (Temminck, 1840) – eastern India, Sri Lanka, Burma, Thailand, Cambodia, Vietnam, western Malaysia, Sumatra, Mentawai Isls, Riau Arch., Java, Borneo (Simmons, 2005: 508); hermani Thomas, 1923 – Sumatra (Indonesia) (Simmons, 2005: 508); horsfieldii (Temminck, 1840) – India, Andaman Isls, southeastern China, Thailand, Burma, Laos, Vietnam, western Malaysia, Java, Bali, Sulawesi, Borneo, Philippines (Simmons, 2005: 508); hoveli Harrison, 1964 – Israel, Lebanon, Syria, Turkey; hyrcanicus 665 Benda, Reiter and Vallo, 2012 – Only known from the type locality in Iran; ikonnikovi Ognev, 1912 – Ussuri region and North Korea to Lake Baikal (Russia), the Altai Mtns and Mongolia, northeastern China; Sakhalin Isl (Russia) and Honshû and Hokkaido Isls (Japan) (Simmons, 2005: 509); indochinensis Son, Görföl, Francis, Motokawa, Estók, Endo, Thong, Dang, Oshida and Csorba, 2013 – Vietnam; insularum (Dobson, 1878) – Samoa (Simmons, 2005: 509); izecksohni Moratelli, Peracchi, Dias and Oliveira, 2011 – Brazil; keaysi J. A. Allen, 1914 – Tamaulipas (Mexico) to Bolivia, northern Argentina, Peru, Ecuador, Venezuela and Trinidad (Simmons, 2005: 509); laniger (Peters, 1871) – southern China including Tibet, Vietnam, eastern India (Simmons, 2005: 509); lavali Moratelli, Peracchi, Dias and Oliveira, 2011 – Brazil, Paraguay, Argentina; leibii (Audubon and Bachman, 1842) – eastern North America from southern Ontario, southern Quebec (Canada) and southern Maine (USA) to southwards to Georgia and western to eastern Oklahoma (USA) (Simmons, 2005: 509); levis (I. Geoffroy Saint-Hilaire, 1824) – Bolivia, Argentina, southeastern Brazil, Uruguay (Simmons, 2005: 510); longipes (Dobson, 1873) – Afghanistan, northeastern India, Nepal (Simmons, 2005: 510); lucifugus (Le Conte, 1831) – Alaska (USA) to Labrador and Newfoundland (Canada), south to southern California, northern Arizona, northern New Mexico (USA) (Simmons, 2005: 510); macrodactylus (Temminck, 1840) – Japan, Kunashir Isl and Kurile Isls (Russia), southeastern Siberia, Korea (Simmons, 2005: 510); macropus (Gould, 1854) – southern Australia (Simmons, 2005: 511); macrotarsus (G.R. Waterhouse, 1845) – Philippines, northern Borneo (Simmons, 2005: 511); martiniquensis LaVal, 1973 – Martinique, Barbados (Lesser Antilles) (Simmons, 2005: 511); melanorhinus (Merriam, 1890) – British Columbia (Canada) south to central Mexico and east to western Oklahoma (USA) (Simmons, 2005: 511); midastactus Moratelli and Wilson, 2014 – Bolivia, Paraguay; moluccarum (Thomas, 1915) – Ambon and Kai Isls (Moluccas), northern and western Australia, Seram, Waigeo Isl (West Palua, Indonesia), Papua New Guinea, Bismarck Arch., Solomon Isls (Simmons, 2005: 511); montivagus (Dobson, 1874) – Yunnan to Fukien and Chihil (China), northeastern India, Burma, Vietnam, Laos, northeastern Thailand, western Malaysia, Borneo (Simmons, 2005: 511); morrisi Hill, 1971; muricola (Gray, 1846) – Afghanistan through northern India and Nepal to Taiwan, Vietnam, Malaysia, Indonesia and New Guinea (Simmons, 2005: 512); myotis (Borkhausen, 1797) – central and southern Europe, east to Ukraine; southern England, most Mediterranean islands; Asia Minor; Lebanon, Syria and Israel (Simmons, 2005: 512); mystacinus (Kuhl, 1817); nattereri (Kuhl, 1817) – 666 ISSN 1990-6471 Ireland, Great Britain, Europe (except northern Scandinavia), Morocco, northern Algeria, Turkey, Israel, Jorden, Lebanon, Iraq, Iran, Bulgaria, Crimea and Caucasus to Turkmenistan (Simmons, 2005: 513); nesopolus Miller, 1900 – northeastern Venezuela; Curaçao and Bonaire (Netherlands Antilles) (Simmons, 2005: 513); nigricans (Schinz, 1821) – Nayarit and Tamaulipas (Mexico) to Peru, Bolivia, northern Argentina, Paraguay and southern Brazil; Trinidad and Tobago; St. Martin, Montserrat, Grenada (Lesser Antilles) (Simmons, 2005: 513); nyctor LaVal and Schwartz, 1975 – Barbados; Grenada (?); occultus Hollister, 1909 – southern California to Arizona, New Mexico and Colorado (USA), south to Distrito Federal (Mexico) (Simmons, 2005: 514); oxyotus (Peters, 1867) – Venezuela to Bolivia; Colombia, Ecuador, Panama; Costa Rica; Peru; possibly Guyana; pequinius Thomas, 1908 – Hong Kong, Hopeh, Shantung, Honan and Kiangsu (China) (Simmons, 2005: 514); petax Hollister, 1912 – Kamtschatka, Vladivostok, Sakhalin and Kurile Isls (Russia), Japan, Korea, Manchuria, E and S China Mongolia, Tibet, Vietnam; peytoni Wroughton and Ryley, 1913 – S + EC India; phanluongi Borisenko, Kruskop and Ivanova, 2009 – Vietnam; pilosatibialis LaVal, 1973 – Colombia; pilosus (Peters, 1869) – China, Hong Kong, Vietnam, Laos, NE India; planiceps Baker, 1955 – Coahuila, Nuevo León and Zacatecas (Mexico) (Simmons, 2005: 515); pruinosus Yoshiyuki, 1971 – Honshu and Shikoku (Japan) (Simmons, 2005: 515); punicus Felten, Spitzenberger and Storch, 1977; ridleyi Thomas, 1898 – western Malaysia, Sumatra, Borneo (Simmons, 2005: 515); riparius Handley, 1960 – Honduras south to Uruguay, eastern Brazil, Argentina, Paraguay and Bolivia; Trinidad (Simmons, 2005: 515); rosseti (Oey, 1951) – Cambodia, Thailand (Simmons, 2005: 516); ruber (E. Geoffroy Saint-Hilaire, 1806) – southeastern Brazil, southeastern Paraguay, northeastern Argentina (Simmons, 2005: 516); rufoniger (Tomes, 1858) – Korea, Japan, China, Taiwan, Laos, Vietnam; rufopictus (G.R. Waterhouse, 1845) – Philippines; schaubi Komos, 1934 – Arminia and western Iran (Simmons, 2005: 516); scotti Thomas, 1927; secundus Ruedi, Csorba, Lin and Chou, 2015 – Taiwan; septentrionalis (Trouessart, 1897) – eastern USA and Canada west to British Columbia, eastern Montana, eastern Wyoming; south to Alabama, Georgia, and Florida Panhandle (Simmons, 2005: 516); sicarius Thomas, 1915 – northeastern India and Nepal (Simmons, 2005: 516); siligorensis (Hodgson, 1855) – northern India to southern China, Burma, Vietnam and Laos; south to western Malaysia; Borneo (Simmons, 2005: 516); simus Thomas, 1901 – Colombia, Ecuador, Peru, northern Brazil, Bolivia, northeastern Argentina and Paraguay (Simmons, 2005: 517); sodalis Miller and G.M. Allen, 1928 – New Hampshire to Florida Panhandle, west to Wisconsin and Oklahoma (USA) (Simmons, 2005: 517); soror Ruedi, Csorba, Lin and Chou, 2015 – Taiwan (only known from the type specimen); stalkeri Thomas, 1910 – Kai and Gebe Isls (Molucca Isls), Waigeo Isl. (Prov. of Papua, Indonesia) (Simmons, 2005: 517); thysanodes Miller, 1897 – Chiapas (Mexico) to southwestern South Dekota (USA) and southcentral British Columbia (Canada) (Simmons, 2005: 517); tricolor (Temminck, 1832); velifer (J. A. Allen, 1890) – Honduras to Kansas and southeastern California (USA) (Simmons, 2005: 517); vivesi Menegaux, 1901 – coast of Sonora and Baja California (Mexico), mainly on small islands (Simmons, 2005: 518); volans (H. Allen, 1866) – Jalisco to Veracruz (Mexico); Alaska Panhandle (USA) to Baja California (Mexico), east to northern Nuevo León (Mexico), South Dekota (USA) and central Alberta (Canada) (Simmons, 2005: 518); weberi (Jentink, 1890) – Indonesia (Sulawesi); welwitschii (Gray, 1866); yanbarensis Maeda and Matsumura, 1998 – Northern Okinawa Isl (Japan) (Simmons, 2005: 518); yumanensis (H. Allen, 1864) – Hidalgo, Morelos and Baja California (Mexico) north to British Columbia (Canada), east to Montana and western Texas (USA) (Simmons, 2005: 518); zenatius Ibáñez, Juste, Salicini, Puechmaille and Ruedi, 2019. COMMON NAMES: Czech: netopýři myší, ouška myší, netopýrcové. English: Mouse-eared Bats, Little Brown Bats, Hairy Bats, Large-footed Bats. French: Murins. German: Mausohr, Mausohren. Polish: nocki. PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: In Africa Myotis sp. have been found deposits of the Plio-Pleistocene, at Bolt's Farm, South Africa (Broom, 1948). Early Pleistocene, at Olduvai (Butler, 1978). Geraads et al. (2010: 279) reported the presence of Myotis in Late Cenozoic (ca. 2.5 MYA) deposits in Ahl al Oughlam, Morocco. Molecular dating performed by Ruedi et al. (2013: 437-441) suggest an origin of all recent Myotis in the early Miocene (about 21 MYA with 95 % highest posterior density interval 23 - 20 MYA). Shi and Rabosky (2015: 1532) report the approximate fossil date for crown Myotinae to be 27 MYA. However, a recent study by Gunnell et al. (2017) reported on a fossil Myotis found in a Belgian Oligocene locality, dating 33 MYA. Morales et al. (2019: 2264) find this to be consistent with other fossiles of a primitive Myotis-like form found in Africa suggest and dating from the late Eocene African Chiroptera Report 2020 (37.8 - 33.9 mya), which suggest that the early diversification of the subfamily might have occurred on that continent. However, these authors also indicate that this interpretation depends on the correct identification of the "Myotis-like" material. Alternatively, the radiation of Myotis could only date from about 18 mya. The Ethiopian clade (see Taxonomy) would have been separated about 12.33 (15 - 10) MYA. MOLECULAR BIOLOGY: Baker et al. (1974) recorded the karyotype of Myotis as 2n = 44, AA = 50, but Dippenaar et al. (1983) give it as 2n = 50, AA = 48 in both seabrai and lesueuri. PREDATORS: In Algeria, Djilali et al. (2016: 159) found the Shorteared owl (Asio flammeus) to feed primarily on 667 bats of the genus Myotis. They made up 37.8 % of all prey items (195 specimens on 516 prey items). The second one was Gerbillus nanus with only 5.4 % of all preys. PARASITES: From non-specified Myotis specimens, the following nycteribiids were reported by Vermeil (1960) from Tunisia: Penicillidia conspicua Speiser, 1900, Nycteribia vexata vexata Westwood, 1835, Nycteribia biarticulata Hermann, 1804, Nycteribia latreillii Leach, 1817, and Nycteribia schmidlii Schiner, 1853. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo (Democratic Republic of the), Egypt, Ethiopia, Gabon, Kenya, Malawi, Tanzania, Tunisia, Uganda. †Myotis darelbeidensis Gunnell, Eiting and Geraads, 2011 *2011. Myotis darelbeidensis Gunnell, Eiting and Geraads, N. Jb. Geol. Paläont. Abh., 260 (1): 59, figs 3C, 5A-C, 8A-C. Publication date: January 2011. Type locality: Morocco: Ahl al Oughlam [ca. 33 35 N 07 30 W] [Goto Description]. Holotype: INSAP AaO 4571: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Right dentary fragment with p2-m3. Paratype: INSAP AaO 4589: Collected by: ?: Collector Unknown. Presented/Donated by: ?: Collector Unknown. Left maxilla P4-M1. - Etymology: Name taken from Dar el Beida (“white house”), the Arabic name for Casablanca (see Gunnell et al., 2011: 60). (Current Combination) TAXONOMY: Gunnell et al. (2011: 60) describe it as a large species of Myotis, similar in size to M. blythii and M. myotis, much larger than most other extant Mediterranean Myotis species (including M. capaccinii, M. bechsteinii, M. emarginatus). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Timeframe: Pliocene (Brown et al., 2019: Suppl.). Myotis anjouanensis Dorst, 1960 *1960. Myotis Goudoti anjouanensis Dorst, Bull. Mus. natn. Hist. nat., Paris, sér. 2, 31: 476 (for 1959). Type locality: Comoros: Anjouan Island [Goto Description]. Holotype: MNHN ZM-MO-1886-1536: ad ♀, skull and alcoholic. Collected by: Léon Humblot. Presented/Donated by: ?: Collector Unknown. - Etymology: The specific name refers to the island where the type specimen was collected. ? Myotis anjouanensis (Name Combination, Current Combination) TAXONOMY: Considered a synonym of goudoti by Koopman (1993a: 211), and a subspecies of goudoti by Peterson et al. (1995: 89), but a valid species by Simmons (2005). COMMON NAMES: Czech: netopýr anžuánský. English: Anjouan Myotis, Anjouan Mouse-eared Bat. French: Murin d'Anjouan, Vespertilion de l'Ile d'Anjouan. German: Anjouan-Mausohr. Csorba et al. (2014: 667) include anjouanensis in the subgenus Chrysopteron. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since it has only recently been recognised as a distinct species, and there is still very little information on 668 ISSN 1990-6471 its extent of occurrence, status, threats and ecological requirements (Jacobs, 2008c; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Jacobs, 2008c; IUCN, 2009). 2004: DD ver 3.1 (2001) (Jacobs, 2004k; IUCN, 2004). POPULATION: Structure and Density:- There is currently little information on the abundance of this species (Jacobs, 2008c; IUCN, 2009). Specimens were collected by Goodman, 2007) from Anjouan in 2007. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Comoros. Regional None known. MAJOR THREATS: The threats to this species are not known (Jacobs, 2008c; IUCN, 2009). CONSERVATION ACTIONS: Jacobs (2008c) [in IUCN (2009)] reports that it is not known if the species is present in any protected areas. Further studies are needed into the natural history and possible threats to this littleknown species. GENERAL DISTRIBUTION: Myotis anjouanensis is endemic to the island of Anjouan in the Comoros Islands. Native: Comoros. Figure 245. Distribution of Myotis anjouanensis Myotis bocagii (Peters, 1870) *1870. Vespertilio Bocagii Peters, J. Sci. mat. phys. nat., ser. 1, 3 (10): 125. Publication date: December 1870. Type locality: Angola: Duque de Bragança [08 55 S 16 10 E]. Lectotype: ZMB 3973: ad ♂, skull and alcoholic. Presented/Donated by: José Vicente Barboza du Bocage. Lectotype designated by Turni and Kock (2008: 57). - Comments: Turni and Kock (2008: 57) also indicate that Peters did not mention the number of specimens he examined, therefore it might be possible that additional syntypes might have existed in the Lisbon museum, but were destroyed by the fire in 1978. Thorn et al. (2009: 51) situate the type locality at 09 23 S 15 53 E. - Etymology: In honour of J.V. Barbosa du Bocage, an eminent Portuguese zoologist, who worked during the nineteenth century, and who named material from the Congo and Angola collected by his Portuguese colleague Snr. M. Jose d'Anchieta (Smithers, 1983: 93). 1976. Myotis boccagei: Brosset, Z. Tierpsychol., 42 (1): 50. (Lapsus) 1996. Myotis bocagi: Kityo and Kerbis, J. East Afr. Nat. Hist., 85: 63. (Lapsus) 2019. Myotis bocageii: Hranac, Marshall, Monadjem and Hayman, Epidemics, Suppl.. Publication date: 16 November 2019. (Lapsus) ? Myotis (Selysius) bocagei: (Name Combination, Alternate Spelling) ? Myotis bocagei: (Alternate Spelling) ? Myotis bocagii: (Name Combination, Current Combination) TAXONOMY: Mentioned as bocagei by Corbet (1978: 49), Smithers (1983: 93), Meester et al. (1986: 49), Koopman (1993a: 209), Horácek et al. (2000: 121) Wilson and Cole (2000: 58), Hutson et al. (2001: 28), Fenton and Bogdanowicz (2002: 1008) but as bocagii by Ansell and Dowsett (1988: 37), Harrison and Bates (1991: 76), Grubb et al. (1998: 82), Stadelmann et al. (2004b: 178); and as bocagi by Kityo and Kerbis (1996: 63). Csorba et al. (2014: 667) include bocagii in the subgenus Chrysopteron, but see Ruedi and Mayer (2001: 436), Ghazali et al. (2016: 476) and African Chiroptera Report 2020 Patterson et al. (2019: "2") who reject the subgeneric division. The phylogenetic analyses by Patterson et al. (2019: "5") revealed the presence of two groups within M. bocagii: one including a single Senegalese specimen and two all the remainder, including material from Ghana, DRC, Kenya, and Tanzania. These findings do not support the currently accepted division into two subspecies: bocagii and cupreolus, although the authors stress that no material from Angola (where the type of 669 Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Jacobs, 2017a). 2008: LC ver 3.1 (2001) (Jacobs, 2008d; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs, 2004f; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa:- 2016: LC ver 3.1 (2001) (Monadjem et al., 2016j). 2004: DD ver 3.1 (2001) (Friedmann and Daly, 2004). 1986: Indeterminate (Smithers, 1986). MAJOR THREATS: There appear to be no major threats to this species as a whole (Jacobs, 2008d; IUCN, 2009; Monadjem and Jacobs, 2017a). Figure 246. Myotis bocagii (TM 46637) caught near Haziview, Mpumalanga, South Africa. bocagii was collected) was included in the analyses (p. "9"). As such, it might be possible that two subspecies might still be present, with bocagii restricted to Angola. Until this has been resolved, we do retain the two traditionally recognized subspecies. Patterson et al. (2019) also indicated that M. bocagii forms a sister group with M. goudoti + M. anjouanensis, which in turn form a sister group to M. tricolor and M. emarginatus (based on nuclear genes). COMMON NAMES: Afrikaans: Rooi langhaarvlermuis. Castilian (Spain): Murciélago de Orejas de Ratón. Chinese: 棕 红 鼠 耳 蝠 . Czech: netopýr Bocageův. English: Rufous Mouse-eared bat, Bocage's Mouse-eared Bat, Rufous Mouse-eared Myotis, Rufous Hairy Bat, Bocage Banana Bat, Bocage's Hairy Bat. French: Murin roux, Murin de Du Bocage. German: Kupferfarbenes Mausohr, Bananen-Mausohr. Portuguese: Morcego lanudo de Bocage. CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs, 2008d; IUCN, 2009; Monadjem and Jacobs, 2017a). CONSERVATION ACTIONS: Jacobs (2008d) [in IUCN (2009)] and Monadjem and Jacobs (2017a) reports that it has been recorded from Kruger National Park, South Africa (Skinner and Chimimba, 2005), Dzanga-Sangha Special Dense Forest Reserve in the Central African Republic (Lunde et al., 2001), and from the Manga Forest Reserve in Tanzania (Doggart et al., 1999b), and in view of the species wide range it is presumed to be present in many more protected areas. There are no direct conservation measures currently needed for this species as a whole. GENERAL DISTRIBUTION: Myotis bocagii is widespread throughout much of sub-Saharan Africa. It occurs from Sierra Leone and Senegal in West Africa, eastwards through Cameroon and Central Africa, to Ethiopia and East Africa, being recorded as far south as northeastern South Africa. Outside of Africa it has been recorded from southern Yemen. For southern Africa, Cooper-Bohannon et al. (2016: Table S2) calculated a potential distribution area of 594,469 km2. Native: Angola (Crawford-Cabral, 1989; Monadjem et al., 2010d: 564); Benin; Burkina Faso (Kangoyé et al., 2015a: 618); Burundi; Cameroon; Central African Republic; Congo (The Democratic Republic of the) (Allen, 1917; Hayman et al., 1966; Van Cakenberghe et al., 1999; Monadjem et al., 2010d: 564); Côte d'Ivoire; Equatorial Guinea; Ethiopia; Gabon; Ghana; Guinea (Fahr et al., 2006a: 73); Kenya; Liberia (Monadjem and Fahr, 2007: 50); Malawi (Happold et al., 1988; Monadjem et al., 2010d: 564); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 564; Monadjem et al., 2010c: 383); Nigeria; Rwanda; Senegal; Sierra Leone (Weber et al., 670 ISSN 1990-6471 2019: 25); South Africa (Monadjem et al., 2010d: 564); Swaziland (Monadjem et al., 2010d: 564; Shapiro and Monadjem, 2015: 353); Sudan; Tanzania (Stanley and Goodman, 2011: 43); Togo; Uganda; Yemen; Zambia (Monadjem et al., 2010d: 564); Zimbabwe (Monadjem et al., 2010d: 564). Bates et al. (2013: 339) reject the occurrence of M. bocagii in the Republic of Congo as Happold (2013dn: 693) only provided a map without any supporting data. ECHOLOCATION: Schoeman and Waddington (2011: 291) mention a peak frequency of 43.2 ± 0.2 kHz and a duration of 2.0 ± 0.1 msec for specimens from Durban, South Africa. Monadjem et al. (2017c: 179) reported for one specimen the following parameters from Swaziland: Fmin: 30.1 kHz, Fknee: 41.2 kHz, Fc: 36.6 kHz, and duration: 2.8 msec. They were also to capture the call up to a distance of 5 m. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Rautenbach et al. (1993) reported 2n = 44, FN = 50, BA = 8, a submetacentric X chromosome, and an acrocentric Y chromosome, for specimens from South Africa. These results are the same as those obtained for M. tricolor specimens from South Africa and Zimbabwe (Rautenbach et al., 1993). Protein / allozyme - Unknown. HABITAT: In southern Africa, this species primarily (82 %) occurs in Savanna habitats (Cooper-Bohannon et al., 2016: Table S2). In the Mount Nimba area, Monadjem et al. (2016y: 369) collected one specimen in old growth forest at 420 m. Weier et al. (2019b: 6) investigated droppings from this species in the Levubu region (Limpopo, RSA) and found DNA sequences of Bathycoelia distincta Distant, 1878 (Hemiptera). POPULATION: Structure and Density:- Although this species is difficult to catch it is possibly not very rare (Jacobs, 2008d; IUCN, 2009; Monadjem and Jacobs, 2017a). Trend:- 2016: Unknown (Monadjem and Jacobs, 2017a). 2008: Unknown (Jacobs, 2008d; IUCN, 2009). REPRODUCTION AND ONTOGENY: Happold and Happold (1990b: 567) reported that young were born in the middle of the wet season in Malawi. Stanley and Goodman (2011: 43) report on a male with scrotal testes collected in the East Usambara Mountains (Tanzania) on 30 July 1992. One female captured on 4 June in Sierra Leone by Weber et al. (2019: 25) was pregnant. VIRUSES: Coronaviridae - Coronaviruses SARS-CoV - Müller et al. (2007b) tested between 1986 and 1999, for antibodies to SARS-CoV in serum of one individual from Limpopo Province, South Africa, which tested negative (0/1). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Angola, Benin, Burundi, Cameroon, Central African Republic, Congo, Congo (Democratic Republic of the), Côte d'Ivoire, Equatorial Guinea, Eswatini, Ethiopia, Gabon, Ghana, Guinea, Kenya, Malawi, Mozambique, Niger, Rwanda, Senegal, South Africa, Tanzania, Togo, Uganda, Zambia, Zimbabwe. HABITS: Brosset (1976) found that harem groups of female M. bocagii's remained very stable over a period of several years. The males in these groups were replaced more often. DIET: From the Umbilo River in KwaZulu Natal (RSA), Naidoo et al. (2011: 26) reported the following prey volume percentages: during summer: Lepidoptera (1.3 ± 1.9), Coleoptera (0.8 ± 1.0), Hemiptera (8.5 ± 13.1), Diptera (37.8 ± 25.8), and Trichoptera (51.8 ± 29.9); during winter: Lepidoptera (7.5 ± 10.6), Coleoptera (32.5 ± 46.1), Hemiptera (20.0 ± 28.3), Diptera (33.4 ± 33.0), and Trichoptera (6.7 ± 4.7). Figure 247. Distribution of Myotis bocagii African Chiroptera Report 2020 671 Myotis bocagii bocagii (Peters, 1870) *1870. Vespertilio Bocagii Peters, J. Sci. mat. phys. nat., ser. 1, 3 (10): 125. Publication date: December 1870. Type locality: Angola: Duque de Bragança [08 55 S 16 10 E] [Goto Description]. 1904. Myotis Hildegardeæ Thomas, Ann. Mag. nat. Hist., ser. 7, 13 (75): 209. Publication date: 1 March 1904. Type locality: Kenya: Fort Hall [=Muranga] [00 43 S 37 09 E, 4 000 ft] [Goto Description]. Holotype: BMNH 1903.3.2.2: ♂. Collected by: Mrs. Hildegarde Hinde; collection date: 17 October 1902; original number: 115. ? Myotis bocagei bocagei: (Alternate Spelling) ? Myotis bocagei hildegardeae: (Name Combination, Alternate Spelling) ? Myotis bocagii bocagii: (Name Combination, Current Combination) GENERAL DISTRIBUTION: Uganda to Ethiopia, south to Angola, Zambia, Malawi, and Transvaal (South Africa). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Angola, Cameroon, Congo (Democratic Republic of the), Kenya, Malawi, South Africa, Tanzania, Uganda. Myotis bocagii cupreolus Thomas, 1904 *1904. Myotis Bocagei cupreolus Thomas, Ann. Mag. nat. Hist., ser. 7, 13 (78): 407. Publication date: 1 June 1904. Type locality: Cameroon: Efulen [02 46 N 10 42 E] [Goto Description]. Holotype: BMNH 1903.2.4.6: ♂. Collected by: George Latimer Bates Esq. Collection date: 14 August 1901. - Comments: Evenhuis (2003: 36) states that pages 329 to 424 were included in part 77, which appeared on 1 May 1904. However, the title page of the publication indicates that the part is 78, which was published on 1 June 1904. (Current Combination) ? Myotis bocagii cupreolus: (Current Spelling) GENERAL DISTRIBUTION: West Africa. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Cameroon, Central African Republic, Congo (Democratic Republic of the), Côte d'Ivoire, Ghana, Liberia, Nigeria, Sierra Leone. Myotis capaccinii (Bonaparte, 1837) *1837. Vespertilio Capaccinii Bonaparte, Iconografia Fauna italiana, 1, fasc. 20 and pl., fig. Publication date: 1837. Type locality: Italy: Sicily [Goto Description]. Holotype: BMNH 1907.1.1.734:. - Etymology: In honour of Francesco Capaccini, under secretary of state of Rome about 1833-34, and a patron of Bonaparte's "Iconografia della Fauna Italica," published in 1832-1841 (Palmer, 1904: 49). (see Palmer, 1904: 49; Kozhurina, 2002: 15). ? Myotis capaccinii capaccinii: (Name Combination) ? Myotis capaccinii: (Name Combination, Current Combination) TAXONOMY: Most recent authors consider M. capaccinii as a monotypic species (Spitzenberger and von Helversen, 2001; Simmons, 2005). COMMON NAMES: Albanian: Lakuriq nate gishtgjatë. Arabian: Khaffash. Armenian: Միջերկրածովային գիշերաչղջիկ. Azerbaijani: Uzunbarmaq gecə şəbpərəsi. Basque: Saguzar hatz-luze, Saguzar hankahandia. Belarusian: Начніца даўгапалая. Bosnian: Dugoprsti šišmiš. Breton: Gousperell Capaccini. Bulgarian: Дългопръст нощник. Castilian (Spain): Murciélago patudo, Murciélago ratonero patudo. Catalan (Spain): Ratpenat de peus grans, Rat penat de peus grans. Croatian: Dugonogi šišmiš, Dugoprsti šišmiš. Czech: Netopýr dlouhonohý. Danish: Capaccinis flagermus. Dutch: Capaccini's vleermuis. English: Capaccinii's Mouse-eared Bat, Long- 672 ISSN 1990-6471 fingered Bat, Long-fingered Myotis. Estonian: Pikkjalg-lendlane. Finnish: Pitkäsormisiippa. French: Murin de Capaccini, Vespertilion de Capaccini, Vespertilion des montagnes. Frisian: Grutfuottige flearmûs. Galician (Spain): Morcego de pés grandes. Georgian: კაპაჩინის მღამიობი. German: Langfußfledermaus, Freischeinige Stelzfussfledermaus. Greek: Ποδαρομυωτίδα. Hebrew: ‫גדות נשפון‬, Nishpon Gadot. Hungarian: Hosszúlábú denevér, Nycteris i macropus. Irish Gaelic: Ialtóg fhadmhéarach. Italian: Vespertilio di Capaccini. Latvian: Garpirkstu naktssikspārnis. Lithuanian: Ilgapirštis pelėausis. Luxembourgish: Laangfoussfliedermaus. Macedonian: Долгопрст ноћник [= Dolgoprst Nokjnik]. Maltese: Farfett il-Lejl tas-Swaba Twal. Montenegrin: Dugoprsti večernjak. Norwegian: Langfotflaggermus. Polish: Nocek długopalcy. Portuguese: Morcego de Capaccini. RhaetoRomance: Vespertil da Capaccini. Romanian: Liliacul cu picioare lungi, Liliac-cu-picioare-lungi. Russian: Ночница средиземная (итальянская). Serbian: Дугопрсти вечерњак [= Dugoprsti večernjak]. Scottish Gaelic: Ialtag fhadmheurach. Slovak: Netopier dlhonohý. Slovenian: Dolgonogi netopir. Swedish: Storfotsfladdermus. Turkish: Uzunayaklı Yarasa. Ukrainian: Нічниця довгопала. Welsh: Ystlum hirfys. CONSERVATION STATUS: Global Justification The species occupies specialised habitat (caves and associated water systems). In the eastern part of the range it congregates in winter in a few sites which are threatened by human disturbance. It has declined between 30 and 50 % in Spain in the last 10 years, and there are indications of declines in other parts of the range. It only hunts in watercourses and is therefore threatened by water pollution and the development of tourist infrastructure, which is expected to continue in the future. It is suspected that population declines are underway that will exceed 30 % over 18 years (3 generations), and for that reason the species is considered Vulnerable (VU A4bce ver 3.1 (2001)) (Hutson et al., 2008k; IUCN, 2009; Paunovic, 2016). Assessment History Global 2016: VU A4bce ver 3.1 (2001) (Paunovic, 2016). 2008: VU A4bce ver 3.1 (2001) (Hutson et al., 2008k; IUCN, 2009). 1996: VU (Baillie and Groombridge, 1996). 1994: VU (IUCN, 1994). 1990: VU (IUCN, 1990). 1988: VU (IUCN, 1988). Regional None known. MAJOR THREATS: Threats include changes in water quality through pollution and dam building, and loss of water bodies and watercourses. Damage or disturbance to caves (tourism, fires and vandalism) used as roosts may also be a problem, as the species is very dependent on caves. The species is collected for medicinal purposes in North Africa (Hutson et al., 2008k; IUCN, 2009; Paunovic, 2016). The species's small distribution range, cave or tree roosting, and aerial hawking feeding style are considered to be the most important risk factors related to climatic change (Sherwin et al., 2012: 174). Bilgin et al. (2012: 434) predict that the range retraction for M. capaccinii as a result of climatic change may pose severe threats to its survival as this species has the bulk of its range in the Mediterranean region. CONSERVATION ACTIONS: Paunovic (2016) supports Hutson et al. (2008k) [in IUCN (2009)] who report that it is protected by national legislation in most range states. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention in the range states where these apply. It is included in Annex II (and IV) of EU Habitats and Species Directive, and hence requires special measures for conservation including designation of Special Areas for Conservation. Some habitat protection through Natura 2000. In Spain, fences are in place to protect several known colonies. Measures needed include protection of colonies (these measures should avoid the blocking of any cave entrances with gates and control of tourist access) and improvement of water quality. GENERAL DISTRIBUTION: Myotis capaccinii is sparsely distributed from eastern Iberia, Spain through the northern Mediterranean to coastal Asia Minor and Israel, Lebanon and Jordan, and also in Mesopotamia from Turkey to Iran and in north-west Africa (limited to the Mediterranean fringe of western Maghreb: north Morocco and northwest Algeria). It occurs from sea level to 900 m. Bilgin et al. (2012) examined climate change scenarios within the Asia Minor and Levant region, they predict that M. capaccinii will have a range contraction, which may pose severe threats to their survival of this species. Native: Albania; Algeria (Kowalski and RzebikKowalska, 1991: 91); Andorra; Austria; Bosnia and Herzegovina; Bulgaria; Croatia; Cyprus (Benda et al., 2007: 71); France [Corse]; Greece [Kriti]; Holy See [Vatican City State]; Iran, Islamic Republic of; African Chiroptera Report 2020 Iraq; Israel; Italy [Sardegna, Sicilia]; Jordan; Lebanon; Macedonia, the former Yugoslav Republic of; Montenegro; Morocco (Aulagnier and Thevenot, 1986: 44); Romania; Russian Federation; Serbia; Slovenia; Spain [Baleares]; Syrian Arab Republic; Tunisia (Dalhoumi et al., 2019b: 26); Turkey; Uzbekistan. Regionally extinct: Switzerland. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Almenar et al. (2006: 158) indicate that the large hind feet and pointed wings of this species are well suited for capturing prey while trawling through the water surface. DETAILED MORPHOLOGY: In their study on take-off performance, Gardiner et al. (2014: 1059) determined a wing beat frequency of 12.96 ± 0.18 Hz. FUNCTIONAL MORPHOLOGY: Gardiner and Nudds (2011: 2187), in their study of flight-initiating jumps, found that M. capaccinii did not show a reduced jumping performance, which could have been expected due to the large feet, which leads to extra weight at the extremities. ECHOLOCATION: Highly variable depending on activity and habitat; these data refer to cruising calls in the main habitat. Search-phase call-shape (while hunting above water surface): hyperbolic FM. Startfrequency: 43 kHz; end-frequency: 22 kHz; frequencies with maximum energy: 28 - 35 kHz; call-duration: 6 - 9 msec, inter-call interval: 145 165 msec (Barataud, 2002, 2005). The data for 56 animals from Greece were the following: Fstart: 89.7 ± 7.35 kHz, Fend: 35.1 ± 2.18 kHz, Fpeak: 48.9 ± 4.76 kHz, bandwidth: 14.2 ± 9.93 kHz, duration 4.6 ± 0.77 msec and interpulse interval: 75.4 ± 22.97 msec (Papadatou et al., 2008b: 132). Walters et al. (2012: suppl.) report the next figures for 30 calls of bats from France and Italy: duration: 3.62 ± 1.40 msec, Fmax: 73.51 ± 9.28 kHz, Fmin: 36.6 ± 3.18 kHz, bandwidth: 36.91 ± 10.73 kHz, Fpeak: 52.2 ± 6.93 kHz. The following data for 20 calls from Iran were reported by Benda et al. (2012a: 182): Fstart: 66.0 ± 12.3 (53.1 - 87.1) kHz, Fend: 48.9 ± 2.0 (46.6 - 52.6) kHz, Fpeak: 52.9 ± 1.9 (49.9 - 55.3) kHz, duration: 5.3 ± 0.7 (3.9 - 6.3) msec, and interpulse interval: 49.1 ± 20.4 (19.4 - 84.4) msec. Mehdizadeh et al. (2018: 1) reported that isolation calls of 1 to 4 day old Iranian M. capaccinii were FM shallow calls with longer duration (7.54 ± 1.83 ms) and lower Fpeak (20.07 ± 0.89 kHz) as compared to adult female calls (2.35 ± 0.75 ms and 54.02 ± 4.34 673 kHz). Between days 12 and 16, the calls began to sounc more like these of the adults. Luo et al. (2019a: Supp.) reported the following data (from two calls): Fpeak: 50.4, 48.9 kHz, Fstart: 83.6, 89.7 kHz, Fend: 39.7, 35.1 kHz, Band width: 43.9, 54.6 kHz, and duration: 3.8, 4.6 msec. Aizpurua et al. (2013: 4) indicate that M. capaccinii is able to regulate the temporal component of the echolocation call so that they can modify their capturing technique depending on whether they are hunting for aquatic arthropods or fish. MOLECULAR BIOLOGY: DNA - See Hoofer and Van Den Bussche (2003). Karyotype - Capanna et al. (1968) and Rushton (1970: 463) reported 2n = 44, aFN = 50, X = metacentric, Y = acrocentric for specimens from Italy, while Pérez-Suárez et al. (1991) reported 2n = 44, aFN = 52, X = submetacentric, Y = acrocentric for specimens from Spain. From Italy, Capanna and Manfredi Romanini (1971: 474) mention 2n = 44, aFN =60, 3 metacentric pairs of chromosomes, 1 submetacentric and 17 acrocentric; with the X chromosome being metacentric and the Y chromosome acrocentric. Karaman et al. (2017: 119) indicate that the ddRAD and mtDNA structure suggests that three lineages in the Mediterranean area could be considered different species, but this needs to be investigated further. Protein / allozyme - Unknown. HABITAT: In the eastern Iberian peninsula, Almenar et al. (2006: 157) report that M. capaccinii only used aquatic habitats as foraging sites, and that most of their foraging activity was on rivers. M. capaccinii preferentially used river stretches characterised by an open course and smooth water surfaces (Almenar et al., 2012). They shifted foraging stretches seasonally, likely following the spatiotemporal dynamics of prey production over the watershed (Almenar et al., 2012). HABITS: Aihartza et al. (2008: 287) describe the fishing behaviour of M. capaccinii as observed in a flight tent containing an artificial pool. They distinguished two patterns: "A) long series of circular flights, skimming along the water and dipping in softly twice or three times in each roundabout; B) long figure-eight loops with bats flying faster and higher, swooping down on the centre of the pond, where they snapped their hind feet hard into the water." They also suggest (p. 674 ISSN 1990-6471 297) that the bats detect the presence of fish by touching them when they dip their feet into the water. ROOST: Within the Iberian Peninsula, within the lower basin of the Xúquer River, Almenar et al. (2012) reported a main roost within a limestone cave with a variable colony size through the season: 81 M. capaccinii during pre-breeding, 219 during lactation and 113 during weaning of 2004. More than 100 individuals inhabit the second roost during lactation, located 8.4 km from the first. Both roosts are shared with other bat species (Rhinolophus euryale, Rhinolophus mehelyi, Myotis blythii, Myotis myotis, Myotis emarginatus, Myotis nattereri and Miniopterus schreibersii) (Almenar et al., 2012). DIET: Aihartza et al. (2008: 288) indicate that in the Mediterranean arean aquatic arthropods (e.g. Diptera: Culicidae and Chironomidae) form the main prey of this species, supplemented by Trichoptera, Neuroptera, Lepidoptera, Heteroptera and arachnids. But its diet also contains fish. Although some M. capaccinii populations are known to eat fish in the wild Aihartza et al., 2003; Levin et al., 2006; Biscardi et al., 2007), the diet of the surveyed individuals appeared to be exclusively arthropods (Almenar, 2008). On the Iberian peninsula, Aizpurua et al. (2011: 48) examined 2600 guano samples and retrieved 97 otoliths, all from the mosquitofish (Gambusia holbrooki) which would have been between 1.5 and 3.6 cm in length (maximum size of fishes: 5 cm in males and 8 cm in females, see Aihartza et al., 2003: 196). They also found that one of the bats was able to eat between 8 and 15 fishes in one or two nights. POPULATION: Structure and Density:- See Hutson et al., 2008k) [in IUCN (2009)] and Paunovic (2016). Trend:- 2016: Decreasing (Paunovic, 2016). 2008: Decreasing (Hutson et al., 2008k; IUCN, 2009). Myotis capaccinii populations within the Asia Minor and Levant region are predicted to have a declining population trend, under climate change scenarios (Bilgin et al., 2012: 433). REPRODUCTION AND ONTOGENY: In Greece, most females were pregnant by the second half of April and had given birth before the end of May. The first flying young were found by the end of June or the beginning of July (Papadatou et al., 2008a 509). In Iran, Mehdizadeh et al. (2018: 1) found the forearm length of neonates to be 19.59 ± 1.23 mm, with a linear growth of 0.74 mm/day until day 28. The weight at day one was 3.59 ± 0.23 g, and this increased linearly with 0.15 g/day until day 28. PARASITES: In Italy, Voyron et al. (2011: 193) reported the following fungal entities from carcasses: Alternaria sp., Candida palmioleophila, Chrysosporium/Gymnoascus, Cladosporium cladosporioides, Gimnoascacea, Verticillium lecanii, Trichosporon chiropterorum, Mucor hiemalis f. hiemalis, Mucor racemosus. Ševcík et al. (2012: 35) report on the presence of the bat flies Nycteribia pedicularia Latreille, 1805, Nycteribia schmidlii schmidlii Schiner, 1853, and Penicillidia dufourii Westwood, 1835 on bats from Crete and Cyprus. Nycteribia kolenati Theodor & Moscona, 1954 and Phthiridium biarticulatum Hermann, 1804 were reported for the first time by Haelewaters et al. (2017: 1) from bats from Hungary. From a non-specified area, Haelewaters et al. (2018: 794) added Penicillidia conspicua Speiser, 1901. In Algeria, Bendjeddou et al. (2013: 325) reported two ectoparasites: Cimex lectularius (Linnaeus, 1758) (Hemiptera: Cimicidae) and Rhipicephalus sanguineus (Latreille, 1806) (Arachnidae: Ixodida: Ixodidae). Bendjeddou et al. (2017: 15) added the following species: Nycteribia (Nycteribia) latreillii (Leach, 1817) and Phthiridium biarticulatum Hermann, 1804 (Nycteribiidae); Brachytarsina (Brachytarsina) flavipennis Macquart, 1851 (Streblidae) and Spinturnix myoti (Kolenati, 1856) and Ixodes vespertilionis Koch, 1844 (Arachnida). VIRUSES: Paramyxoviridae Morbillivirus This virus was reported from Bulgaria, Germany and Romania by Kohl and Kurth (2014: 3113). UTILISATION: The species is collected for medicinal purposes in North Africa (Hutson et al., 2008k; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Morocco, Tunisia. African Chiroptera Report 2020 675 Figure 248. Distribution of Myotis capaccinii Myotis dieteri M. Happold, 2005 *2005. Myotis dieteri M. Happold, Acta Chiropt., 7 (1): 11, figs 1 - 2. Type locality: Congo: Loudima: Grotte du Viaduc [04 15 S 13 00 E] [Goto Description]. Holotype: MNHN ZMMO-1985-1925: ad ♀, alcoholic and cranium only (no mandibula). Collected by: JeanPaul Adam; collection date: between 1961 and 1968. - Etymology: In honour of Dr. Dieter Kock of the Senkenberg Museum, Frankfurt, Germany, in recognition of the great contribution he made to the knowledge of African mammals, in recognition of the kindness and generosity with which he has helped so many others with their studies, and in gratitude for our long firendship (see Happold, 2005: 11). (Current Combination) TAXONOMY: Known only from the type specimen (Happold, 2005: 11). MAJOR THREATS: The threats to this species remain unknown (Happold, 2008; IUCN, 2009). Csorba et al. (2014: 668) indicate that dieteri is probably the only sub-Saharan Myotis that doesn't belong to the subgenus Chrysopteron, but molecular data are missing to confirm this. CONSERVATION ACTIONS: Happold (2008) [in IUCN (2009)] reports that it is not known if the species is present within any protected areas. Additional studies are needed into the distribution, abundance, breeding biology and general ecology of this species. There is a need to determine if the species is still present at Grotte du Viaduc à Loudima, and to locate additional populations through directed surveys. COMMON NAMES: Czech: netopýr Kockův. French: Murin de Kock. German: Kocks Mausohr. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) since it has only recently been described, and there is still very little information on its extent of occurrence, status, threats and ecological requirements (Happold, 2008; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Happold, 2008; IUCN, 2009). GENERAL DISTRIBUTION: Myotis dieteri has only been recorded from type locality of 'Grotte du Viaduc à Loudima, Republic of Congo (04º15'S, 13º00'E) (Happold, 2005: 11). Despite extensive surveys of other seemingly suitable localities, it has not been recorded from additional caves in the Loudima-Kimongo region, or from caves in the Mayombe and lower Kouilou regions of Congo, the Haut-Ivindo region of Gabon, or Kikwit in the Democratic Republic of Congo (Happold, 2005). Regional None known. Native: Congo (Happold, 2005: 11; Bates et al., 2013: 337). 676 ISSN 1990-6471 POPULATION: Structure and Density:- It is known only from the holotype collected by Jean-Paul Adam sometime between 1961 and 1968, from a small number of this species present in the cave (Happold, 2005). Trend:- 2008: Unknown (Happold, 2008; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Congo. Figure 249. Distribution of Myotis dieteri Myotis emarginatus (E. Geoffroy St.-Hilaire, 1806) *1806. Vespertilio emarginatus E. Geoffroy Saint-Hilaire, Ann. Mus. Hist. nat. Paris, 8: 198. Type locality: France: Ardennes: Givet: Charlemont. Holotype: MNHN 819-202: ad, mounted skin (skull not removed). Additional skull A6953: see Rode (1941: 244), who mentions Abbeville as locality, and M. Baillon as collector. ? Myotis emarginata: - Comments: corrected gender for specifc name. (Alternate Spelling) ? Myotis emarginatus emarginatus: (Name Combination) ? Myotis emarginatus: (Name Combination, Current Combination) TAXONOMY: Assigned to the subgenus Isotus by Pavlinov et al. (1995: 97), but Csorba et al. (2014: 667) include it in the subgenus Chrysopteron. Woodman (1993) notes that many mammalian generic names ending in -otis use the wrong gender for specific names therefore this name should be M. emarginata. However, see also Benda and Tsytsulina (2000: 335), and Simmons (2005). COMMON NAMES: Albanian: Lakuriq nate i Geoffroy-it. Arabian: Khaffash. Armenian: Եռագույն գիշերաչղջիկ. Azerbaijani: Üçrəng gecə şəbpərəsi. Basque: Geoffroy saguzar. Belarusian: Начніца трохкаляровая. Bosnian: Trobojni šišmiš. Breton: Gousperell Geoffroy. Bulgarian: Трицветен нощник. Castilian (Spain): Murciélago orejirroto, Murciélago de Geoffroy, Murciélago ratonero pardo. Catalan (Spain): Ratpenat d'orelles dentades, Ratapinyada d'orella escapçada. Croatian: Riđi šišmiš, Trobojni šišmiš. Czech: Netopýr brvitý, Netopýra brvitého, Nedopír wraubkowaný, Stejnoušan zejkatý, Netopýrec brvitý. Danish: Geoffroys flagermus. Dutch: Ingekorven vleermuis. English: Geoffroy's Bat, Notch-eared Bat, Geoffroy's Myotis. Estonian: Ripslendlane. Finnish: Ruskosiippa. French: Murin à oreilles échancrées, Vespertilion à oreilles échancrées, Vespertilion échancré. Frisian: Ynsniene flearmûs. Galician (Spain): Morcego de orellas fendidas. Georgian: სამფერი მღამიობი. German: Wimperfledermaus, Kerbohrige Ohrenfledermaus, Fledermaus mit gerändelten Ohren. Greek: Πυρρομυωτίδα. Hebrew: Nishpon Pgoom Ozen. Hungarian: Csonkafülű denevér, Nycteris i blephardoti. Irish Gaelic: Ialtóg Geoffroy. Italian: Vespertilio smarginato, Vespertilio smarginato. Latvian: Skropstainais naktssikspārnis. Lithuanian: Trispalvis pelėausis. Luxembourgish: Wimperefliedermaus. Macedonian: Тробоен ноќник [= Troboen Nokjnik]. Maltese: Farfett ilLejl ta' Widnejħ Imnaqqxa. Montenegrin: Riđi večernjak. Norwegian: Vinkeløreflaggermus, Geoffroyflaggermus. Polish: Nocek orzęsiony. Portuguese: Morcego-lanudo. Rhaeto-Romance: Vespertil cun tschegls, Vespertil cun tscheglias. Romanian: Liliacul cărămiziu, Liliac-cu-gene-lungi. Russian: Ночница трёхцветная, Трехцветная нu1086\чница [= Tryokhtsvetnaya nochnitsa]. Serbian: Риђи вечерњак [= Riđi večernjak]. Scottish Gaelic: Ialtag Geoffroy. Slovak: Netopier brvitý. Slovenian: Vejicati netopir. Swedish: Flikörad fladdermus. Turkish: Kirpikli Yarasa. African Chiroptera Report 2020 Ukrainian: Нічниця триколірна. Geoffroy. Welsh: Ystlum CONSERVATION STATUS: Global Justification The species experienced a significant decline in at least parts of its range from the 1960s to the 1990s, but now it is expanding in central Europe and is stable or not significantly declining elsewhere. The range is still large and the species is not specialized to restricted habitat, although it has a very specialized diet. Assessed as Least Concern (LC ver 3.1 (2001)) (Hutson et al., 2008m; IUCN, 2009; Piraccini, 2016a). Assessment History Global 2016: LC ver 3.1 (2001) (Piraccini, 2016a). 2008: LC ver 3.1 (2001) (Hutson et al., 2008m; IUCN, 2009). 1996: VU A2c ver 2.3 (1994) (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: In Europe the species is mainly associated with agricultural landscapes, therefore all agricultural activities can affect populations of this species. Loss of and disturbance to roost sites in buildings (including remedial timber treatment in attics) and underground sites are also threats. In the African part of the range, cave habitats, where the species roosts, are being destroyed by fires and vandalism. The species is also collected for traditional medicine practices in North Africa (Hutson et al., 2008m; IUCN, 2009; Piraccini, 2016a). CONSERVATION ACTIONS: Piraccini (2016a) supports Hutson et al. (2008m) [in IUCN (2009)] who report that it is protected by national legislation in most range states. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention in the range states where these apply. It is included in Annex IV of EU Habitats and Species Directive, and there is some habitat protection through Natura 2000. Protection of roosts and promotion of awareness about the lack of medicinal value of the species is required. GENERAL DISTRIBUTION: Myotis emarginatus occurs in southern Europe from Portugal in the west to the Balkans in the east and southern part of western and central Europe and non-arid parts of southwestern Asia from Asia Minor, Caucasus region and Palestine to Oman, Uzbekistan and Tajikistan; also in north-west 677 Africa (recorded from northern Maghreb (Algeria, Morocco and Tunisia)). In the eastern part of the Mediterranean it occurs in various parts of Turkey, Syria, Lebanon and Israel. It occurs from sea level to 1,800 m, highest records in the Alps are 812 m (maternity colony) and 1,505 m (hibernaculum) (Spitzenberger, 2002). Native: Afghanistan; Albania; Algeria; Andorra; Armenia; Austria; Azerbaijan; Belgium; Bosnia and Herzegovina; Bulgaria; Croatia; Cyprus; Czech Republic; France [Corse]; Georgia; Germany; Gibraltar; Greece [Kriti] (Benda et al., 2007: 71); Hungary; Iran, Islamic Republic of; Israel; Italy [Sardegna]; Jordan; Kazakhstan; Kyrgyzstan; Lebanon; Luxembourg; Macedonia, the former Yugoslav Republic of; Monaco; Montenegro; Morocco (El Ibrahimi and Rguibi Idrissi, 2015: 360); Netherlands; Oman; Poland; Portugal; Romania; Russian Federation; San Marino; Saudi Arabia; Serbia; Slovakia; Slovenia; Spain [Baleares]; Switzerland; Syrian Arab Republic; Tajikistan; Tunisia (Dalhoumi et al., 2014: 53); Turkey; Turkmenistan; Ukraine; Uzbekistan. ECHOLOCATION: Highly variable depending on activity and habitat; these data refer to cruising calls in the main habitat. Search-phase call-shape (close to foliage): steep linear FM. Start-frequency: 110 kHz; end-frequency: 39 kHz; frequency with maximum energy: 65 kHz; call-duration: 2.6 msec; inter-call interval: 26-32 msec (Barataud, 2005). The data for 42 animals from Greece were the following: Fstart: 110.5 ± 10.90 kHz, Fend: 42.6 ± 5.57 kHz, Fpeak: 72.8 ± 7.28 kHz, bandwidth: 43.4 ± 8.29 kHz, duration 3.3 ± 0.73 msec and interpulse interval: 79.4 ± 23.83 msec (Papadatou et al., 2008b: 132). Walters et al. (2012: suppl.) report the following figures for 26 calls (24 sequences) from France and Germany: duration: 2.29 ± 0.70 msec, Fmax: 93.23 ± 20.18 kHz, Fmin: 42.00 ± 3.47 kHz, bandwidth: 51.23 ± 17.78 kHz, Fpeak: 60.23 ± 15.14 kHz. MOLECULAR BIOLOGY: DNA - See García-Mudarra et al. (2009). Karyotype - Bovey (1949) and Pérez-Suárez et al., 1991) reported 2n = 44, aFN = 50, X = submetacentric, Y = acrocentric, while Radjhabli et al. (1970) reported aFN = 56 and Zima (1978) aFN = 52 with X = metacentric. The autosomal component consists of three large metacentric, one small metacentric, 15 acrocentric and two dotlike pairs (with one being bi-armed) (Arslan and Zima, 2014: 10). The latter authors also refer to 678 ISSN 1990-6471 the X chromosome as being a medium-sized metacentric. HABITAT: In Spain, Goiti et al. (2011: 19) found that M. emarginatus flew on average 5 km to its foraging areas, with a maximum distance of almost 10 km. The foraging area covered between 120 and 371 ha, and consisted mainly of scrubland (35 % of mean individual foraging time) and pine plantations (34 % of mean individual foraging locations), followed by isolated trees (oak, pine, or eucalyptus - on average 11 %), riparian woods (7 %), oak dehesas (6 %), and eucalyptus plantations (5 %). DIET: Goiti et al. (2011: 20) found that, in Spain, spiders made up the major part of the diet by both volume (mean 79 %) and frequency (in feces of all examined bats, and in 95 % of the feces). The next most important group was formed by moths, with a mean consumption of 7 % of the diet by volume; dipterans (flies and midges - 6.3 %), and lacewings (especially the family Hemerobiidae 4.4 %). Earwings, harvestmen, wasps, and beetles all made up less than 2 % of the diet. Benda et al. (2012a: 325) summarizes the European food groups as: Araneae, Lepidoptera, brachyceran Diptera, and Lepidoptera larvae. In the Middle East Diptera, Lepidoptera, Coleoptera, and Homoptera are prevailing in Azerbaijan, Araneae and Brachycera in Syria, and Hemiptera, Araneae, and Brachycera in Turkey. In Iran, there was a high proportion of Tipulidae, and in Jordan of scarabaeid beetles. This lead Benda et al. (2012a: 325) to conclude that a certain level of flexibility of the trophic niche in M. emarginatus. POPULATION: Structure and Density:- Locally it can be rare or common. The species experienced a significant decline from the 1960s to the 1990s, but in recent time the numbers in several regions have increased and the species has spread into new areas. It lives in large colonies (up to 1,200 individuals in Austria in one maternity colony) (Hutson et al., 2008m; IUCN, 2009; Piraccini, 2016a). Arachnida: Eyndhovenia euryalis (G. Canestrini, 1885), Macronyssus rhinolophi (Oudemans, 1902), Steatonyssus periblepharus Kolenati, 1858, Spinturnix emarginata (Kolenati, 1856). Insecta: Basilia nana Theodor & Moscona, 1954, Basilia italica Theodor, 1954, Cimex pipistrelli Jenyns, 1839, Ischnopsyllus emarginatus, Ischnopsyllus simplex Rothschild, 1906, Penicillidia dufourii (Westwood, 1835), Phthiridium biarcticulatum, Rhinolophopsylla unipectinata (Taschenberg, 1880). Trematoda - Digenea: Lecithodendrium linstowi Dollfus, 1931, Prosthodendrium aelleni Dubois, 1956, Prosthodendrium chilostomum (Mehlis, 1831). Cestoda: Myotolepis grisea (Beneden, 1873). Bendjeddou et al. (2017: 15) mention the following parasites from bats from Algeria: Nycteribiidae: Nycteribia (Nycteribia) latreillii (Leach, 1817) and Penicillidia (Penicillidia) dufourii Westwood, 1835; Arachnida: Spinturnix myoti (Kolenati, 1856) and Ixodes vespertilionis Koch, 1844. VIRUSES: Orthomyxoviridae Despite sampling and testing of 3 individuals of this species, no evidence of influenza A-like viruses (Orthomyxoviridae) were obtained (Fereidouni et al., 2015). UTILISATION: The species is also collected for traditional medicine practices in North Africa (Hutson et al., 2008m; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Algeria, France, Macedonia, Morocco, Tunisia. Trend:- 2016: Stable (Piraccini, 2016a). 2008: Stable (Hutson et al., 2008m; IUCN, 2009). M. emarginatus populations within the Asia Minor and Levant region are predicted to have a stable population trend, under climate change scenarios (Bilgin et al., 2012: 433). PARASITES: Frank et al. (2015: 8) mention the following parasites for European M. emarginatus: Figure 250. Distribution of Myotis emarginatus African Chiroptera Report 2020 679 Myotis goudoti (A. Smith, 1834) *1834. Vespertilio goudoti A. Smith, S. Afr. Quart. J., ser. 2, 2: 244. Publication date: June 1834. Type locality: Madagascar: "Madagascar" [Goto Description]. 1858. Vespertilio Madagascariensis Tomes, Proc. zool. Soc. Lond., 1858, XXVI (ccclii): 89. Publication date: 27 April 1858. Type locality: Madagascar: "Madagascar". Holotype: BMNH 1907.1.1.503:. 1870. Vespertilio sylvicola A. Grandidier, Rev. Mag. Zool., ser. 2, 22: 49. Publication date: February 1870. Type locality: Madagascar: Ambaravantao forest [Goto Description]. Comments: The type locality was mentioned as "Forests of Ambaravaranvatou" by Allen (1939a: 91). Dobson (1878: 301, footnote) indicates that the type has been lost and that the description is insufficient. He, therefore, included it with a question mark. ? Myotis goudoti goudoti: (Name Combination) ? Myotis goudoti: (Name Combination, Current Combination) TAXONOMY: Simmons (2005) states that it does not appear to include anjouanensis; see Peterson et al. (1995). Csorba et al. (2014: 667) include goudoti in the subgenus Chrysopteron, but Ruedi and Mayer (2001: 436), Ghazali et al. (2016: 476) and Patterson et al. (2019: "2") reject the division in subgenera. Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem et al., 2017bv). 2008: LC ver 3.1 (2001) (Andriafidison et al., 2008h; IUCN, 2009). 1996: LR/nt ver 2.3 (1994) (Baillie and Groombridge, 1996). Regional None known. COMMON NAMES: Czech: netopýr madagaskarský. English: Malagasy Mouse-eared Bat. French: Murin de Madagascar, Chauve-souris malgache à oreille de souris, Vespertilion de Madagascar. German: Madagaskar-Mausohr. MAJOR THREATS: There are no known threats to this species although additional research is needed to determine the impact of forest degradation on its persistence in cave roosts (IUCN, 2009; IUCN, 2009; Monadjem et al., 2017bv). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: Samonds (2007: 60) found samples in deposit (SS2), which was collected from the surface near the main cave entrance, dating attemps revealed this sample was contaminated, therefore no date is known for this locality in Anjohibe cave, Madagascar. CONSERVATION ACTIONS: Andriafidison et al. (2008h) [in IUCN (2009)] and Monadjem et al. (2017bv) report that this is a widespread species that is found in many of Madagascar’s protected areas, including those in the east and west of the island (Goodman, 1999; Russ et al., 2003; Goodman et al., 2005a). Weyeneth (2010) indicated that, during the Pleistocene, populations of this speciesi diverged into a southern and a northern component, with subsequent expansion and adaptation to all biomes. GENERAL DISTRIBUTION: Myotis goudoti is endemic to the island of Madagascar (Simmons, 2005) where it is widely distributed across a range of different vegetation types (Peterson et al., 1995; Goodman, 1999; Eger and Mitchell, 2003; Russ et al., 2003; Goodman et al., 2005a), including Nosy Be and Nosy Komba (Rakotonandrasana and Goodman, 2007: 6). CONSERVATION STATUS: Global Justification This species is listed as Least Concern (LC ver 3.1 (2001)) in view of its widespread distribution across Madagascar, and the absence of any major threats. Close monitoring of roosting colonies in areas where the forest is subject to disturbance is needed (Andriafidison et al., 2008h; IUCN, 2009; Monadjem et al., 2017bv). Native: Madagascar (Peterson et al., 1995; Goodman, 1999; Eger and Mitchell, 2003; Russ et al., 2003; Goodman et al., 2005a; Rakotonandrasana and Goodman, 2007: 6). 680 ISSN 1990-6471 DETAILED MORPHOLOGY: Baculum: Rakotondramanana and Goodman (2017: 64) reported the structure as being triangular, and in some cases, with a pronounced proximal indentation; length: 0.87 ± 0.111 (0.68 - 1.02) mm, width: 0.53 ± 0.081 (0.43 - 0.67) mm. ECHOLOCATION: Russ et al. (2003) describe the echolocation as a single frequency-modulated sweep from 75 kHz to 55 kHz. Kofoky et al. (2009: 382) reported the calls of 11 individuals, which were characterized by broadband FM sweeps produced at low duty cycle. The most energy always found in the fundamental at about 64.4 kHz, and the pulses were very short about 3 ms. MOLECULAR BIOLOGY: DNA.- Unknown. Karyotype.- Richards et al. (2010) examined one female and one male having a 2n=44 and a FNa= 50, a submetacentric X and small acrocentric Y chromosome. In addition, Volleth and Eick (2012: 167) mention a segment number of 22. HABITAT: Dammhahn and Goodman (2013: 108) mention the lower portion of forest structure and partially open areas as the foraging habitat for this species. ROOST: Wilkinson et al. (2012: 160) indicate that caves are the typical roosting sites for these bats on Madagascar. Ravelomanantsoa et al. (2019: 110) added crevices and possibly tree holes. DIET: Razakarivony et al. (2005) found Araneae in 27 % of stomachs examined. Rakotoarivelo et al. (2007) in Madagascar found M.goudoti to mainly consumed Hymenoptera, Neuroptera, Araneae and fragments of spiders were present in 55 % of the fecal samples. Rakotondramanana et al. (2015: 78) report the following diet items (volume percent, frequency percent): Coleoptera (34.0, 100.00); Hymenoptera (4.0, 60.0); Lepidoptera (62.0, 100.0). Rasoanoro et al. (2015: 64) found the following volume percentages for two specimens: Aranea (40.7), Coleoptera (43.7), Hymenoptera (8.4), Isoptera (4.6), Lepidoptera (2.5). Kemp et al. (2018: Suppl.) used DNA metacarcoding to detect insect pest species in the diet of these bats and found the following prey orders (in descending order): Coleoptera, Hymenoptera, Lepidoptera (Homona sp., Thaumatotibia batrachopa (Meyrick, 1908)), Diptera (Simulium lineatum (Meigen, 1804), Culex sp., Culex annulioris Theobald, 1901, Coquillettidia sp., Anopheles sp.), Sarcoptiformes, Ephemeroptera, Hemiptera, Araneae, Blattodea, Trichoptera, Symphypleona, Mesostigmata, Trombidiformes. PREDATORS: Goodman et al. (2015c: 78) found the remains of one individual in pellets of Bat Hawk Macheiramphus alcinus Bonaparte, 1850 in western central Madagascar. POPULATION: Structure and Density:- Although widespread, there is a paucity of information on the population status of M. goudoti. It is frequently trapped in mist nets during bat surveys and appears to be at least locally abundant in west (Kofoky et al., 2007; Rakotoarivelo and Randrianandrianina, 2007), but is more difficult to net, or is less abundant in the east (Russ et al., 2003; Randrianandriananina et al., 2006). There are few data available on roosting colonies but aggregations of up to 1,000 individuals have been recorded (Monadjem et al., 2017bv). Trend:- 2016: Unknown (Monadjem et al., 2017bv). 2008: Unknown (Andriafidison et al., 2008h; IUCN, 2009). ACTIVITY AND BEHAVIOUR: Rakotoarivelo et al. (2007) suggest that M. goudoti is better adapted to forage in cluttered areas and may have a closer association with forest vegetation. They also suggest that M. goudoti may be a gleaner. PARASITES: BACTERIA Lagadec et al. (2012: 1696), Dietrich et al. (2014: Suppl.) and Gomard et al. (2016: 5) report the presence of spirochaetes bacteria of the genus Leptospira, which are assigned to L. borghpetersenii by Dietrich et al. (2018a: 3). Lei and Olival (2014: Suppl.) mention M. goudoti as host for Bartonella bacteria. HAEMOSPORIDA Megali et al. (2010: 1042) and Duval et al. (2012: 1563) reported Genbank data for Polychromophilus sp. from M. goudoti. Ramasindrazana et al. (2018: Suppl.) identified this parasite as Polychromophilus murinus Dionisi, 1899. Ramasindrazana et al. (2016: 6) reported the presence of an unnamed filariod in this bat species. African Chiroptera Report 2020 DIPTERA Nycteribiidae: Ramasindrazana et al. (2017: Suppl.) found the bat flies Nycteribia stylidiopsis and Penicillidia sp. on specimens from this bat species. ACARI Trombiculidae: Stekolnikov (2018a: 116) reported Trisetica aethiopica (Hirst, 1926) from this bat species. 681 Rhabdoviridae Of the 11 bats tested by Mélade et al. (2016a: 6), four showed a seroreaction. UTILISATION: See Goodman et al. (2008d). DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Madagascar. VIRUSES: Astroviridae Lebarbenchon et al. (2017 Suppl.) tested 14 animals and three of them was positive for Astroviruses. Paramyxoviridae Wilkinson et al. (2012: 160) tested 2 individuals from the Madagascar using RT-PCR specific for Respirovirus/Morbillivirus/Henipahvirus (RMH) and Paramyxovirinae (PMV) and found 0 positive results for viral nucleic acids. Wilkinson et al. (2014) tested 8 individuals from Anjohikinakia and Bekoaky in Madagascar with an RT-PCR specific for the Respiro-, Morbilli- and Henipavirus genera, two of the 8 individuals tested positive for paramyxovirus RNA. 48 Madagascan M. goudoti specimens were tested by Mélade et al. (2016b: 4) and five of them were positive for paramyxoviruses. Figure 251. Distribution of Myotis goudoti Myotis morrisi Hill, 1971 *1971. Myotis morrisi Hill, in: Hill and Morris, Bull. Br. Mus. (nat. Hist.) Zool., 21 (2): 43, pl. 1, 2 (a, b). Publication date: 28 April 1971. Type locality: Ethiopia: Walaga, Blue Nile Gorge: Didessa River mouth: "Forward Base Three" [10 05 N 35 38 E, ca. 1 000 m] [Goto Description]. Holotype: BMNH 1970.488: ad ♀, skin and skull. Collected by: The Great Abbai Expedition; collection date: 28 August 1968; original number: A107. (Largen, Yalden & King). Also see Hill and Morris (1971: 43). - Etymology: In honour of Dr. Pat Morris of Royal Holloway College, University of London, in recognition of his many services to the study of the Ethiopian fauna while with the Great Abbai Expedition (Hill in Hill and Morris, 1971: 44). (Current Combination) TAXONOMY: Known only from two specimens (see Stadelmann et al., 2004b: 179). COMMON NAMES: Czech: netopýr Morrisův. English: Morris's Mouse-eared Bat, Morris's Bat, Morris's Myotis. French: Murin d'Ethiopie, Vespertilion de Morris. German: Morris' Mausohr. CONSERVATION STATUS: Global Justification Listed as Data Deficient (DD ver 3.1 (2001)) in view of the absence of sufficient information on its extent of occurrence, threats and conservation status (Jacobs et al., 2008o; IUCN, 2009). Assessment History Global 2008: DD ver 3.1 (2001) (Jacobs et al., 2008o; IUCN, 2009). 2004: VU D2 ver 3.1 (2001) (Jacobs et al., 2004b; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. 682 ISSN 1990-6471 MAJOR THREATS: The threats to this species are unclear. It may be threatened by cave disturbance and subsistence harvesting for food (Jacobs et al., 2008o; IUCN, 2009). Trend:- 2008: Unknown (Jacobs et al., 2008o; IUCN, 2009). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia, Nigeria. CONSERVATION ACTIONS: Jacobs et al. (2008o) [in IUCN (2009)] report that it is not known if the species is present in any protected areas. Further studies are needed into the distribution, abundance, natural history, and threats to this poorly known species. GENERAL DISTRIBUTION: Myotis morrisi has only been recorded from two very disjunct sites, the type locality of 'Didessa River mouth, Walaga' in Ethiopia (Hill and Morris, 1971) and from Numan, Adamawa Province, in northeastern Nigeria (Hill et al., 1988). Native: Ethiopia (Hill and Morris, 1971); Nigeria (Hill et al., 1988). POPULATION: Structure and Density:- Little information is available on the population abundance or size of this species (Jacobs et al., 2008o; IUCN, 2009). Figure 252. Distribution of Myotis morrisi Myotis mystacinus (Kuhl, 1817) *1817. Vespertilio mystacinus Kuhl, Die Deutsche Fledermäuse, 7, 58. Type locality: Germany. Syntype: ZMB 461: ♂, skin and skull. Unknown locality, presumably coll. Kuhl, purchased from Kuhl, 09.1818.: see Turni and Kock (2008: 60). ? Myotis mystacinus: (Name Combination, Current Combination) TAXONOMY: Despite the fact that numerous species have been recently recognized within this complex, the whiskered bats from Morocco are still included in the nominal form (von Helversen et al., 2001; Mayer et al., 2007; García-Mudarra et al., 2009). At present, this species includes three forms differing in morphological traits: M. mystacinus mystacinus (most of European range), M. mystacinus occidentalis (Iberia and Morocco) and M. mystacinus caucasicus (Caucasus region). COMMON NAMES: Albanian: Lakuriq nate me mustaqe. Armenian: Բեղավոր գիշերաչղջիկ. Azerbaijani: Bığlı gecə şəbpərəsi. Basque: Saguzar biboteduna. Belarusian: Начніца вусатая. Bosnian: Mali brkati šišmiš. Breton: Gousperell varvek. Bulgarian: Мустакат нощник. Castilian (Spain): Murciélago bigotudo, Murciélago ratonero bigotudo. Catalan (Spain): Ratpenat de bigotis, Rat penat de bigotis. Croatian: Brkati šišmiš, Mali brkati šišmiš. Czech: Netopýr vousatý, nedopír walausatý, netopýrec vousatý. Danish: Skægflagermus. Dutch: Baardvleermuis, Gewone pisang blad vleermuis. English: Whiskered Mouse-eared Bat, Whiskered Bat, Whiskered Myotis, Mouse-eared bat. Estonian: Habelendlane. Finnish: Viiksisiippa. French: Murin à moustaches, Vespertilion à moustaches. Frisian: Burdflearmûs. Galician (Spain): Morcego bigotudo, Morcego de bigotes. Georgian: ულვაშა მღამიობი. German: Bartfledermaus, Kleine Bartfledermaus, Schnauzbärtige Fledermaus. Greek: Μουστακονυχτερίδα. Hungarian: Bajuszos denevér, Nycteris i mystakophoros. Indonesian: Kelelewar myotis pucuk. Irish Gaelic: Ialtóg ghiobach. Italian: Vespertilio mustacchino. Latvian: Bārdainais naktssikspārnis. Lithuanian: Ūsuotasis pelėausis. Luxembourgish: Kleng Baartfliedermaus. Macedonian: Мустаќест ноќник [= Mustakjest Nokjnik]. Maltese: Farfett ilLejl Daqni. Montenegrin: Brkati večernjak. Norwegian: Skjeggflaggermus. Polish: Nocek wasatek. Portuguese: Morcego-de-bigodes. Rhaeto-Romance: Vespertil dal barbis, Vespertil African Chiroptera Report 2020 pitschen dal barbis. Romanian: Liliacul mustăcio, Liliac-cu-mustãþi. Russian: Ночница усатая, Усатая ночница [= Usataya nochnitsa]. Serbian: Тамнолики бркати вечерњак [= Tamnoliki brkati večernjak]. Scottish Gaelic: Ialtag ghibeach. Slovak: Netopier fúzatý. Slovenian: Brkati netopir, Skupina brkatih netopirjev. Swedish: Mustaschfladdermus. Turkish: Bıyıklı Siyah Yarasa. Ukrainian: Нічниця вусата. Welsh: Ystlum barfog. CONSERVATION STATUS: Global Justification Myotis mystacinus has a large population size and a wide distribution. No declines in population size have been detected, and there are no known widespread major threats. Assessed as Least Concern (LC ver 3.1 (2001)) (Hutson et al., 2008r; IUCN, 2009; Coroiu, 2016). Assessment History Global 2016: LC ver 3.1 (2001) (Coroiu, 2016). 2008: LC ver 3.1 (2001) (Hutson et al., 2008r; IUCN, 2009). Regional None known. MAJOR THREATS: There are no major threats to this species overall, but the species is affected by loss of woodland and other aspects of land management and development. It is also affected by loss of and damage to roost sites in trees, buildings and underground habitats. In the African part of the species' range, cave habitats where the species roosts are being destroyed by fires and vandalism. This species is also collected for medicine, but not at a level that constitutes a threat to the species (Hutson et al., 2008r; IUCN, 2009; Coroiu, 2016). CONSERVATION ACTIONS: Coroiu (2016) supports Hutson et al. (2008r) [in IUCN (2009)] who report that it is protected by national legislation in most range states. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention. It is included in Annex IV of EU Habitats and Species Directive, and there is some habitat protection through Natura 2000. It occurs in protected areas throughout its range. Protection of cave roost sites is required. GENERAL DISTRIBUTION: Myotis mystacinus is a western Palaearctic species, occurring in western and central Europe, southern parts of Scandinavia, the British Isles, Morocco, northern parts of eastern Europe, western parts of the Caucasus and the Urals. 683 The occurrence in southeast Europe is questionable but likely. It is marginal in Africa, restricted to Moroccan mountains from the Rif in the north to the southern slope of High Atlas in the south. It has been recorded from sea level up to 1,920 m asl (Gerell, 1999). Native: Andorra; Armenia; Austria; Azerbaijan; Belgium; Bosnia and Herzegovina; Bulgaria; China; Croatia; Czech Republic; Denmark; Estonia; Finland; France [Corse]; Georgia; Germany; Greece [Kriti]; Hungary; Ireland; Italy; Latvia; Liechtenstein; Lithuania; Luxembourg; Macedonia, the former Yugoslav Republic of; Montenegro; Morocco (Benda et al., 2004d; El Ibrahimi and Rguibi Idrissi, 2015: 360); Netherlands; Norway; Poland; Romania; Russian Federation; Serbia; Slovakia; Slovenia; Spain; Sweden; Switzerland; Turkey; Ukraine; United Kingdom. Regionally extinct: Portugal. Presence uncertain: Albania; Lebanon. ECHOLOCATION: Papadatou et al. (2008b: 132) report the following data for 1 Greek specimen belonging to M. mustacinus bulgaricus: Fstart: 104.0 kHz, Fend: 33.6 kHz, Fpeak: 47.7 kHz, bandwidth: 9.1 kHz, duration 3.8 msec and interpulse interval: 72.7 msec. Data for 51 calls from France, Greece, Switzerland, and the UK, presented by Walters et al. (2012: suppl.) include: duration: 2.36 ± 0.59 msec, Fmax: 84.76 ± 14.10 kHz, Fmin: 37.04 ± 3.24 kHz, bandwidth: 47.72 ± 14.14 kHz, Fpeak: 51.55 ± 7.70 kHz. MOLECULAR BIOLOGY: DNA - See von Helversen et al. (2001), Mayer et al. (2007), García-Mudarra et al. (2009). Karyotype - Bovey (1949) reported a 2n = 44, aFN = 50, X = metacentric, Y = acrocentric, while Zima (1978) found aFN = 52. Arslan and Zima (2014: 9) added an FN of 54 and that the autosomal component consists of three large metacentric, one small metacentric, 15 acrocentric and two dotlike pairs. POPULATION: Structure and Density:- In Europe, it is one of the more common species within the regular distribution area. It is very rare in North Africa, with only 25 specimens in 4 locations (Hutson et al., 2008r; IUCN, 2009; Coroiu, 2016). Trend:- 2016: Unknown (Coroiu, 2016). 2008: Unknown (Hutson et al., 2008r; IUCN, 2009). 684 ISSN 1990-6471 PARASITES: In Europe, Genov et al. (1992) reported the presence of the following Nematodes: Molinostrongylus alatus (Ortlepp, 1932), M. skrjabini Skarbilovitch, 1934, M. spasskii Andrejko, Pinchuk and Skvorzov, 1968, and M. vespertilionis Morosov and Spassky, 1961. Frank et al. (2015: 8) provide the following overview for parasites in European M. mystacinus: Arachnida: Argas vespertilionis (Latreille, 1796, Ixodes sp., Ixodes vespertilionis Koch, 1844, Macronyssus ellipticus (Kolenati, 1857), Macronyssus flavus (Kolenati, 1856), Spinturnix kolenatii Oudemans, 1910, Spinturnix myoti (Kolenati, 1856), Spinturnix mystacinus (Kolenati, 1857), Steatonyssus periblepharus Kolenati, 1858, Steatonyssus spinosus Wilmann, 1936. Insecta: Basilia nana Theodor & Moscona, 1954, Basilia nattereri Kolenati, 1857, Basilia italica Theodor, 1954, Cimex pipistrelli (Jenyns, 1839), Ischnopsyllus hexactenus (Kolenati, 1856), Ischnopsyllus mysticus Jordan, 1942, Ischnopsyllus octactenus (Kolenati, 1856), Ischnopsyllus simplex Rotschild, 1906, Ischnopsyllus variabilis (Wagner, 1898), Myodopsylla trisellis Jorda, 1929, Nycteribia kolenatii Theodor and Moscona, 1954, Nycteribia pedicularia Latreille, 1805, Nycteridopsylla longiceps Rotschild, 1908, Nycteridopsylla pentactena (Kolenati, 1856). Pintschuk and Skvorzov, 1968, Molinostrongylus vespertilionis Morosov and Spassky, 1961. Cestoda: Vampirolepis balsaci (Joyeux and Baer, 1934). VIRUSES: Orthomyxoviridae Despite sampling and testing of 44 individuals of this species, no evidence of influenza A-like viruses (Orthomyxoviridae) were obtained (Fereidouni et al., 2015). UTILISATION: Species are also collected for medicine, but not at a level that constitutes a threat to the species (Hutson et al., 2008r; IUCN, 2009; Coroiu, 2016). DISTRIBUTION BASED ON VOUCHER SPECIMENS: Morocco. Trematoda - Digenea: Lecithodendrium linstowi Dollfus, 1931, Plagiorchis vespertilionis (Muller, 1784), Prosthodendrium chilostomum (Mehlis, 1831). Nematoda: Molinostrongylus alatus (Ortlepp, 1932), Molinostrongylus spasskii Andrejko, Figure 253. Distribution of Myotis mystacinus Myotis punicus Felten, 1977 1875. Vespertilio murinus africanus Dobson, Ann. Mag. nat. Hist., ser. 4, 16 (94): 260. Publication date: 1 October 1875. Type locality: Gabon: "Gaboon". Holotype: BMNH 1873.4.16.5:. . - Comments: The type locality was originally mentioned as 'Gaboon', but the specimen is more probably from Kashmir, see Blanford (1888-91) and Hayman and Hill (1971: 35) (in Corbet and Hill, 1992: 120). However, see also Grubb (2004: 92); who indicates that Gabon might not be rejected. *1977. Myotis blythii punicus Felten, in: Felten, Spitzenberger and Storch, Senckenb. biol., 58 (1/2): 39. Publication date: 22 August 1977. Type locality: Tunisia: Cap Bon: El Haouaria cave [37 03 N 11 01 E] [Goto Description]. Holotype: SMF 44104: ad ♂, skull and alcoholic. Collected by: Nagel, Schubert and Indulis Evalds Vesmanis; collection date: 25 March 1971. See Felten et al. (1977: 39). Paratype: SMF 18946: ♂, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 15 May 1960. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 18947: ♂, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 15 May 1960. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 18948: ♂, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 15 May 1960. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 18949: ♀, skull and African Chiroptera Report 2020 alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: alcoholic. Collected by: Presented/Donated by: ?: 685 ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18950: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18951: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18952: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18955: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18956: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18957: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18958: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18959: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18960: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18961: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18962: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18963: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 18964: ♀, skull and ?: Collector Unknown; collection date: 15 May 1960. Collector Unknown. Paratype: SMF 22214: ♂, skull and ?: Collector Unknown; collection date: 12 March 1963. Collector Unknown. Paratype: SMF 22217: ♀, skull and ?: Collector Unknown; collection date: 12 March 1963. Collector Unknown. Paratype: SMF 22218: ♀, skull and ?: Collector Unknown; collection date: 12 March 1963. Collector Unknown. Paratype: SMF 22219: ♀, skull and ?: Collector Unknown; collection date: 12 March 1963. Collector Unknown. Paratype: SMF 22220: ♀, skull and ?: Collector Unknown; collection date: 12 March 1963. Collector Unknown. Paratype: SMF 43386: ♀, skull and ?: Collector Unknown; collection date: 25 March 1971. Collector Unknown. Paratype: SMF 43387: ♀, skull and ?: Collector Unknown; collection date: 25 March 1971. Collector Unknown. Paratype: SMF 43388: ♀, skull and ?: Collector Unknown; collection date: 25 March 1971. Collector Unknown. Paratype: SMF 43389: ♀, skull and ?: Collector Unknown; collection date: 25 March 1971. Collector Unknown. Paratype: SMF 43390: ♀, skull and ?: Collector Unknown; collection date: 25 March 1971. Collector Unknown. Paratype: SMF 43392: ♀, skull and ?: Collector Unknown; collection date: 25 March 1971. Collector Unknown. Paratype: SMF 43961: ♀, skull and ?: Collector Unknown; collection date: 28 August 1972. Collector Unknown. Paratype: SMF 43962: ♀, skull and ?: Collector Unknown; collection date: 28 August 1972. Collector Unknown. Paratype: SMF 43963: ♀, skull and ?: Collector Unknown; collection date: 28 August 1972. Collector Unknown. Paratype: SMF 43964: ♀, skull and ?: Collector Unknown; collection date: 28 August 1972. Collector Unknown. Paratype: SMF 43965: ♀, skull and ?: Collector Unknown; collection date: 28 August 1972. Collector Unknown. Paratype: SMF 43966: ♀, skull and 686 ISSN 1990-6471 2000. ? ? ? ? ? ? alcoholic. Collected by: ?: Collector Unknown; collection date: 28 August 1972. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 43967: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 28 August 1972. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 43968: ♂, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 28 August 1972. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44097: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44098: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44099: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44100: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44101: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44102: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44103: ♂, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Paratype: SMF 44618: ♀, skull and alcoholic. Collected by: ?: Collector Unknown; collection date: 25 March 1971. Presented/Donated by: ?: Collector Unknown. Myotis cf. punicus Castella, Ruedi, Excoffier, Ibanez, Arlettaz and Hausser, Mol. Ecol., 9: 1761. Myotis blythii oxygnathus: Myotis blythii: Myotis myotis myotis: Myotis myotis: Myotis oxygnathus: Myotis punicus: (Name Combination, Current Combination) TAXONOMY: Originally described as a subspecies of M. blythii, but recently shown to lie outside a clade including blythii, myotis and oxygnathus (Ruedi and Mayer, 2001). Considered a valid species by Castella et al. (2000), Dietz and von Helversen (2004: 71), Benda et al. (2004d) and Evin et al. (2008). Biollaz et al. (2010) suggest that the Corsican and Sardinian populations are currently isolated from the continental gene pool and therefore should be considered as different evolutionary significant units. Baron (2012: 5) found that the population on Malta is a single panmitic unit with an tendency towards becoming isolated mating systems. COMMON NAMES: Albanian: Lakuriq nate maghrebian. Armenian: Փյունիկյան գիշերաչղջիկ. Azerbaijani: Siçanqulaq gecə şəbpərəsi. Basque: Saguzar arratoi-belarri mairutar. Belarusian: Начніца фінікійская, Начніца магрыбская. Bosnian: Punski šišmiš. Breton: Gousperell ar Magreb. Bulgarian: Магребски нощник. Castilian (Spain): Murciélago ratonero moruno. Catalan (Spain): Ratpenat rater africà. Croatian: Sjevernoafrički oštrouhi šišmiš. Czech: Netopýr punský. Danish: Mahgreb museøre. Dutch: Fenicische vale vleermuis. English: Maghrebian Mouse- eared Bat, Felten's Myotis. Estonian: Vahemere lendlane. Finnish: Tunisiansiippa. French: Murin du Maghreb. Frisian: Maghreb mûsearflearmûs. Galician (Spain): Morcego rateiro africano. Georgian: მაღრიბული მღამიობი. German: Punisches Mausohr, Maghreb-Mausohr. Greek: Τρανομυωτίδα της Καρχηδόνας. Hungarian: Magrebi denevér. Irish Gaelic: Ialtóg luch-chluasach Mhaigribeach. Italian: Vespertilio maghrebino. Latvian: Pūnijas naktssikspārnis. Lithuanian: Finikinis pelėausis. Luxembourgish: Maghreb-Mausouer. Macedonian: Магребски ноќник. Maltese: Farfett il-Lejl Widnet il-ġurdien ta' Maghreb, Farfett il-Lejl Widnet il-Gurdien. Montenegrin: Punski več ernjak. Norwegian: Mahgrebmusø re. Polish: Nocek punicki. Portuguese: Morcego-rato do Magrebe. Rhaeto-Romance: Vespertil ureglia-mieur punic. Romanian: Liliacul din Maghreb. Russian: Ночница финикийская. Serbian: Магребски велики вечерњак [= Magrebski veliki večernjak]. Scottish Gaelic: Ialtag luch-chluasach Mhaigribeach. Slovak: Netopier magrebský. Slovenian: Punski netopir. Swedish: Medelhavsmusöra. Turkish: Mahrep Farekulaklı Yarasası. Ukrainian: Нічниця магрибська. Welsh: Ystlum clustlydan Maghreb. African Chiroptera Report 2020 CONSERVATION STATUS: Global Justification Juste and Paunovic (2016a) reports that insufficient information about population dimension and demographic trends to make an assessment. It is probably declining but needs further research. Assessment History Global 2016: DD ver 3.1 (2001) (Juste and Paunovic, 2016a). 2008: NT ver 3.1 (2001) (Aulagnier et al., 2008c; IUCN, 2009). 2004: DD ver 3.1 (2001) (Aulagnier and Juste, 2004; IUCN, 2004). Regional In the Maltese Island Myotis punicus is listed as Vulnerable (VU) (Baron and Vella, 2010). Legal Status Aulagnier et al. (2008c) [in IUCN (2009)] report that it is protected by national legislation in its European range states. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention, in parts of its range where these apply. It is included in Annex IV of the EU Habitats and Species Directive, and some habitat protection may be provided through Natura 2000. MAJOR THREATS: Human disturbance is a major threat, as colonies are located in tourist areas, in caves that are well known and popular. Changes in land management and agricultural pollution are also threats (Aulagnier et al., 2008c; IUCN, 2009). Juste and Paunovic (2016a) report that in the African part of the range, the cave habitat where the species roosts is being destroyed by fires and vandalism. In North Africa the species is collected for traditional medicinal use (Aulagnier et al., 2008c; IUCN, 2009; Juste and Paunovic, 2016a). The species is a cave or tree rooster, its water stress and long-distance dispersal were found to be the major risk factors related to climatic change (Sherwin et al., 2012: 174). CONSERVATION ACTIONS: Juste and Paunovic (2016a) reports that no conservation actions are in place and further research into population trends, establishment and management of protected areas, education, and implementation of nationalscale legislation are needed. While Aulagnier et al. (2008c) [in IUCN (2009)] report that there are ongoing project for the conservation of this species. Appropriate 687 conservation measures include fencing cave entrances (but not gating) and obtaining legal protection for the species. In North Africa further research into population trends, establishment and management of protected areas, education, and implementation of national-scale legislation are needed (Aulagnier et al. (2008c) [in IUCN (2009)). Biollaz et al. (2010) suggest that special care is needed for the Sardinian population as an important source of genetic diversity. GENERAL DISTRIBUTION: Myotis punicus occurs in North-west Africa (Morocco, Algeria, Tunisia, and to western Libya) and on the Mediterranean islands of Corsica (to France), Sardinia (to Italy), and Malta (including Gozo [Baron and Borg, 2013: 407]). Native: Algeria; Libyan Arab Jamahiriya; Malta; Morocco (Benda et al., 2004d; 2010a: 157; El Ibrahimi and Rguibi Idrissi, 2015: 361); Tunisia (Dalhoumi et al., 2014: 53; 2016b: 867, 2019b: 26). BIOGEOGRAPHY: Biollaz et al. (2010: 8)'s study suggests that the islands (Corsica and Sardinia) were colonized by M. punicus in a stepping-stone manner and predates human colonization. Mitochondrial analysis revealed that the insular populations were well differentiated from the mainland, with mean haplotypes and nucleotide diversities decreading from Tunisia to Sardinia, and from Sardinia to Corsica. While a Baysian and ML trees showed that the Corsican haplotypes are nested within the Sardinan ones, suggesting that the Corsica might have been colonized from Sardinia (Biollaz et al., 2010: 8). Compared with the North African populations, the two islands harbour significant lower allelic richness as well as observed and expected heterozygosities, which could reflect recent population crashes or a bottleneck during the colonization of the islands (Biollaz et al., 2010). The Sardinian and Corsican populations were estimated to date back to the early and midPleistocene, respectively (Biollaz et al., 2010). During the glacial episodes that punctuated this period, low sea levels exposed land that was previously underwater. Geographical distances between the islands and the mainland were thus reduced with the emergence of land bridges between some of the islands, favouring island colonization, with the first the colonization of Sardinia from North Africa and then the colonization of Corsica from Sardinia. As the Ice Age ended, sea levels rose and islolated the two islands, explaining the strong reduction of gene flow currently observed (Biollaz et al., 2010). 688 ISSN 1990-6471 The large divergence between the Moroccan and Tunisian populations suggest that both populations had been isolated since the Pleistocene (Biollaz et al., 2010), despite current nuclear gene flow, no female exchange seems to have occured since then. The confinement of Moroccan populations in the High Atlas Range and the philopatric behaviour of females could have enhanced the isolation of these populations, explaining the low haplotype diversity (Biollaz et al., 2010). In North Africa, poor nuclear differentiation between colonies associated with a strong mitochondrial differentiation suggested current male-biased dispersal on the continental area. GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Dietz and von Helversen (2004) and Baron (2012: 1) mention the following external characters: large size (comparable to M. myotis), the wing membrane starting at the base of the toes, a lancet shaped tragus and distinct dorsal (light brown) and ventral (white) fur coloration. ECHOLOCATION: Walters et al. (2012: suppl.) report the following figures for 26 calls (6 sequences) from French bats: duration: 3.62 ± 0.96 msec, Fmax: 60.42 ± 15.65 kHz, Fmin: 27.07 ± 2.26 kHz, bandwidth: 33.35 ± 14.25 kHz, Fpeak: 35.82 ± 4.21 kHz For this species, Disca et al. (2014: 226) report two call types: A - FM APMF, with the following parameters: Fstart: 79.2 ± 16.4 kHz, Fend: 25.0 ± 2.1 kHz, Fpeak: 37 ± 7.9 kHz, bandwidth: 54.2 ± 15.5 kHz, duration: 5.5 ± 0.6 msec; B - FM APBF, with the following parameters: Fstart: 77.3 ± 6.6 kHz, Fend: 20.2 ± 1.2 kHz, Fpeak: 32.0 ± 5.2 kHz, bandwidth: 57.1 ± 7.1 kHz, duration: 8.6 ± 0.6 msec. MOLECULAR BIOLOGY: DNA - See Castella et al. (2000) and Biollaz et al. (2010). Karyotype - Unknown. ROOST: Benda et al. (2014c: 38-39) reported that, in Libya, these bats are only found in ruins, which might serve as temporary roosts or perhaps as maternity roost. DIET: Ahmim and Moali (2011: 45, 48) found that the diet of M. punicus in Algeria consisted of 96.06 % of insects, 2.82 % of chilopods, and 1.12 % of spiders. Among the insects, Diptera (46.32 %), Lepidoptera (20.33 %), and Hemiptera (10.16 %) were found most frequently in the bat's guano. On Malta, Baron and Borg (2013: 415) report that the major food sources are Orthoptera (65 %), Lepidoptera (20 %), and Coleoptera (15 %), whereas on Corsica, the Orthoptera only account for 36 %. In Libya, Benda et al. (2014c: 44) analyses two groups of pellets. One set (from one individual) contained only large spiders (Aranea), while the other consisted primarily of Coleoptera (Scarabaeidae: Melolonthinae), which were supplemented by solifuges (Solpugida), cockroaches (Blattodea) and small scorpions (Scorpionida). POPULATION: Structure and Density:- Over 10,000 individuals, found in large colonies (300 - 500 individuals) (Juste and Paunovic, 2016a). On Corsica, there are around 4,000 individuals in four colonies. The total population size on Corsica, Sardinia and Malta is estimated at between 7,000 and 9,000 individuals. This is a cave dwelling species, therefore there are few colonies. On Malta, over 50 % of the population was lost between the latter half of the 1980s and the early 1990s. Numbers appear to have stabilised at 400 - 450 individuals (Borg, 2002, J.J. Borg unpubl. data [in Aulagnier et al. (2008c) in IUCN (2009)]). Trend:- 2016: Unknown (Juste and Paunovic, 2016a). 2008: Decreasing (Aulagnier et al., 2008c; IUCN, 2009). REPRODUCTION AND ONTOGENY: At Sabratha (Libya), Benda et al. (2014c: 40) collected a pregnant female on 2 April and a juvenile female on 14 April. Dalhoumi et al. (2019a: 158) reported about 50 neonates in a cave in Tunisia on 19 May. PARASITES: Bruyndonckx et al. (2010) collected Spinturnix myoti (Kolenati, 1856) in 11 colonies in North Africa, Corsica and Sardinia. From Libya, Benda et al. (2014c: 44) reported the following ectoparasites: Nycteribiidae: Nycteribia vexata Westwood, 1835, Nycteribia latreillii (Leach, 1817), Spinturnicidae: Spinturnix myoti (Kolenati, 1856), Macronyssidae: Steatonyssus occidentalis (Ewing, 1933), Ischnopsyllidae: Rhinolophopsylla unipectinata (Taschenberg, 1880). From Algeria, Bendjeddou et al. (2013: 325) reported the following ectoparasites: Nycteribiidae: Penicillidia dufouri (Westwood, 1825), Phthiridium biarticulatum Hermann, 1804, Spinturnicidae: Spinturnix myoti (Kolenati, 1856), African Chiroptera Report 2020 Ixodidae: Rhipicephalus sanguineus (Latreille, 1806). Bendjeddou et al. (2016a: 109) added Ixodes vespertilionis Koch, 1844. Bendjeddou et al. (2017: 15) mentioned the following species from Algeria: Nycteribiidae: Nycteribia (Nycteribia) latreillii (Leach, 1817) and Phthiridium biarticulatum Hermann, 1804; Streblidae: Brachytarsina (Brachytarsina) flavipennis Macquart, 1851 and Arachnida: Spinturnix myoti (Kolenati, 1856) and Ixodes vespertilionis Koch, 1844. 689 Serra-Cobo et al. (2018: 2) tested 63 Algerian bats for European bat lyssavirus type 1 and found one being seropositive. This was also the case for 15 out of 262 Moroccan bats. DISTRIBUTION BASED ON VOUCHER SPECIMENS: "Africa", Algeria, Libya, Morocco, Tunisia. Blackwell (1980: 145) and Haelewaters et al. (2017: Suppl.) mention the presence of the Nycteribiid bat-fly Nycteribia vexata Westwood, 1835 on "M. oxygnathus" in Tunisia, which in turn was infected by the fungus Arthrorhynchus nycteribiae (Peyritsch) Thaxter. VIRUSES: Coronaviridae - Coronaviruses Alphacoronavirus Ar Gouilh et al. (2018: 92) reported the EPI 9 virus from Tunisia (3 infected out of 8 bats). Figure 254. Distribution of Myotis punicus Rhabdoviridae Lyssavirus Myotis scotti Thomas, 1927 *1927. Myotis scotti Thomas, Ann. Mag. nat. Hist., ser. 9, 19 (113): 554. Publication date: 1 May 1927. Type locality: Ethiopia: Shoa province: ca. 40 mi (64 km) W Addis Ababa: Djemdjem Forest [09 00 N 38 12 E, 8 000 ft]. Holotype: BMNH 1927.3.4.1:. Paratype: BMNH 1927.3.4.2:. Paratype: BMNH 1927.3.4.3:. Paratype: BMNH 1927.3.4.4:. Paratype: BMNH 1927.3.4.5:. (Current Combination) TAXONOMY: See Simmons (2005). Csorba et al. (2014: 667) include scotti in the subgenus Chrysopteron. COMMON NAMES: Czech: netopýr habešský. English: Scott's Mouse-eared Bat, Scott's Mouse-eared Myotis. French: Murin de Scott, Vespertilion de Scott. German: Scotts Mausohr. CONSERVATION STATUS: Global Justification Listed as Vulnerable (VU B2ab(iii) ver 3.1 (2001)) because its area of occupancy (AOO) is possibly less than 2,000 km 2, its distribution is likely to be severely fragmented, and there is continuing decline in the extent and quality of its forest habitat in Ethiopia (Benda and Lavrenchenko, 2008; IUCN, 2009; Benda and Lavrenchenko, 2017). Assessment History Global 2016: VU B2ab(iii) ver 3.1 (2001) (Benda and Lavrenchenko, 2017). 2008: VU B2ab(iii) ver 3.1 (2001) (Benda and Lavrenchenko, 2008; IUCN, 2009). 2004: VU B2ab(iii) ver 3.1 (2001) (Benda and Lavrenchenko, 2004; IUCN, 2004). 1996: VU (Baillie and Groombridge, 1996). Regional None known. MAJOR THREATS: The forest habitat of this species is being impacted through conversion of land to agricultural use and logging (Benda and Lavrenchenko, 2008; IUCN, 2009; Benda and Lavrenchenko, 2017). CONSERVATION ACTIONS: Benda and Lavrenchenko (2008) [in IUCN (2009)] and Benda and Lavrenchenko (2017) report that there appear to be no conservation measures in 690 ISSN 1990-6471 place, and it is not known if the species is present in any protected areas. There is a need to conserve areas of suitable montane forest habitat for this species. Further studies are needed into the abundance, biology and ecology. DISTRIBUTION BASED ON VOUCHER SPECIMENS: Ethiopia. GENERAL DISTRIBUTION: Myotis scotti is endemic to the highlands of Ethiopia, where it has been recorded from ten localities between 1,300 and 2,500 m asl. Native: Ethiopia. POPULATION: Structure and Density:- It appears to be a rare species that is known from only about 20 specimens (Benda and Lavrenchenko, 2008; IUCN, 2009; Benda and Lavrenchenko, 2017). Trend:2016: Unknown (Benda and Lavrenchenko, 2017). 2008: Decreasing (Benda and Lavrenchenko, 2008; IUCN, 2009). Figure 255. Distribution of Myotis scotti Myotis tricolor (Temminck, 1832) *1832. V[espertilio] tricolor Temminck, in: Smuts, Enumer. Mamm. Capensium, 106. Publication date: 1832. Type locality: South Africa: Cape province: Cape Town [33 56 S 18 28 E] [Goto Description]. - Comments: Type specimen: RMNH ???. 1924. Eptesicus loveni Granvik, Acta Univ. Lund, ser. 2, 21 (3): 12. Publication date: 1924. Type locality: Kenya: E slopes of Mount Elgon [01 07 N 34 35 E, 8 000 ft]. 2013. Myotis tricolor 1: Ruedi, Stadelmann, Gager, Douzery, Francis, Lin, Guillén-Servent and Cibois, Mol. Phylog. Evol., 69 (3): Suppl. Publication date: 27 August 2013. 2013. Myotis tricolor 2: Ruedi, Stadelmann, Gager, Douzery, Francis, Lin, Guillén-Servent and Cibois, Mol. Phylog. Evol., 69 (3): Suppl. Publication date: 27 August 2013. ? Myotis (Selysius) tricolor: (Name Combination) ? Myotis tricolor A: ? Myotis tricolor: (Name Combination, Current Combination) TAXONOMY: (2001: 436), Ghazali et al. (2016: 476) and Patterson et al. (2019: "2"), who reject the division into subgenera. The phylogenetic analyses on mitochondrial DNA performed by Patterson et al. (2019: "5", "9"') identified three separate groups, two corresponding to South African samples and a third from Kenya (close to the type locality of loveni, which might possibly represent a separate (sub)species. The Cytb sequences of these groups differed by 3.7 - 3.8 %. Figure 256. Myotis tricolor (TM 48188) caught at the Farm Schaapplaats, Free State, South Africa. Includes Eptesicus loveni, see Aggundey and Schlitter (1986). Csorba et al. (2014: 667) include tricolor in the subgenus Chrysopteron, but see Ruedi and Mayer COMMON NAMES: Afrikaans: Temminck se langhaarvlermuis, Temminck-langhaarvlermuis, Kaapse Langhaarvlermuis. Chinese: 南非鼠耳蝠. Czech: netopýr trojbarvý. English: Temminck's Mouse-eared Bat, Cape Myotis, Cape Hairy Bat, Tricoloured Mouse-eared Bat, Cape Hairy Myotis, Temminck's Hairy Bat, Three-coloured bat. French: Murin tricolore, Petit murin brun, Murin velu du Cap, African Chiroptera Report 2020 Vespertilion d'Angola, Murin d'Angola. German: Dreifarb-Mausohr. ETYMOLOGY OF COMMON NAME: The colloquial name is after C.J. Temminck, who was the author of Monographies de Mammalogie (1827) (Taylor, 2005). PALAEONTOLOGICAL OR ARCHAEOLOGICAL RECORDS: This species was represented in the very late Holocene at Border Cave, South Africa (Avery, 1991: 6). CONSERVATION STATUS: Global Justification Listed as Least Concern (LC ver 3.1 (2001)) in view of its wide distribution, presumed large population, and because it is unlikely to be declining fast enough to qualify for listing in a more threatened category (Jacobs, 2008e; IUCN, 2009; Monadjem and Jacobs, 2017b). Assessment History Global 2016: LC ver 3.1 (2001) (Monadjem and Jacobs, 2017b). 2008: LC ver 3.1 (2001) (Jacobs, 2008e; IUCN, 2009). 2004: LC ver 3.1 (2001) (Jacobs, 2004e; IUCN, 2004). 1996: LR/lc (Baillie and Groombridge, 1996). Regional South Africa: 2016: LC ver 3.1 (2001) (Monadjem et al., 2016k). 2004: NT ver 3.1 (2001) (Friedmann and Daly, 2004). MAJOR THREATS: There appear to be no major threats to this species as a whole (Jacobs, 2008e; IUCN, 2009; Monadjem and Jacobs, 2017b). CONSERVATION ACTIONS: Jacobs (2008e) [in IUCN (2009)] and Monadjem and Jacobs (2017b) report that it has been recorded from the Virunga National Park in the Democratic Republic of the Congo (Baeten et al., 1984) and in view of its East African range, it seems likely that it is present in additional protected areas. Further studies are needed into the range of this species in West and Central Africa. GENERAL DISTRIBUTION: Myotis tricolor has been patchily recorded in subSaharan Africa. In West Africa the species has currently only been reported from the northwestern uplands of Liberia (Koopman et al., 1995), while in Central Africa it is known only from a few records in the Democratic Republic of the Congo and Rwanda (Hayman et al., 1966; Baeten et al., 1984). Myotis tricolor is much more widely recorded in East Africa, ranging from Ethiopia in 691 the north, through Uganda, Kenya, Tanzania, Malawi, Zambia, Mozambique and Zimbabwe through to southern South Africa. In the RSA, its distribution is best predicted by geology (Babiker Salata, 2012: 50). Native: Congo (The Democratic Republic of the) (Hayman et al., 1966; Monadjem et al., 2010d: 564); Ethiopia; Kenya; Lesotho (Lynch, 1994: 193; Monadjem et al., 2010d: 564); Liberia (Fahr, 2007a: 104); Malawi (Happold et al., 1988; Monadjem et al., 2010d: 564); Mozambique (Smithers and Lobão Tello, 1976; Monadjem et al., 2010d: 564; Monadjem et al., 2010c: 384); Rwanda; Saõ Tomé (Rainho et al., 2010a: 31); South Africa (Monadjem et al., 2010d: 564); Swaziland (Monadjem et al., 2010d: 565); Tanzania; Zambia (Ansell, 1978; Monadjem et al., 2010d: 565); Zimbabwe (Monadjem et al., 2010d: 565). GENERAL DESCRIPTION OF EXTERNAL MORPHOLOGY: Schoeman and Jacobs (2002: 157) mention the following parameters for 1 specimen from the Algeria Forestry Station (RSA): Fa: 46.9 mm, Wing area: 156.6 cm 2, Wing span: 30.6 cm, Wing loading: 6.9 N/m2, and aspect ratio: 6.0. DETAILED MORPHOLOGY: Baculum - Kearney et al. (2002). FUNCTIONAL MORPHOLOGY: Jordaan and Jacobs (2009: 35) indicate that the wing morphology of M. tricolor is intermediate between that of a typical aerial forager, gleaner (takes insects off a substrate) and trawler (takes prey from the water surface). ECHOLOCATION: See Taylor (1999b). Rainho et al. (2010a: 21) reports the calls of 5 individuals from Saõ Tomé. From Algeria Forest Station (RSA), Schoeman and Jacobs (2002: 157) reported the type of calls to be low duty echolocation dominated by frequency modulated calls, with a Fpeak: 50.0 kHz. Jordaan and Jacobs (2009: 35) found that its echolocation call structure is intermediate between that of typical aerial foragers, gleaners and trawlers. Schoeman and Waddington (2011: 291) mention a peak frequency of 40.8 kHz and a duration of 4.4 msec for a specimen from Durban, South Africa. In Waterberg, South Africa and Swaziland, Taylor et al. (2013b: 17) recorded 6 calls with the following parameters: Fmax: 86.1 ± 5.92 (76.6 91.9) kHz, Fmin: 42.7 ± 4.43 (37.1 - 50.0) kHz, Fknee: 692 ISSN 1990-6471 57.6 ± 7.25 (48.5 - 67.5) kHz, Fchar: 52.4 ± 6.13 (45.3 - 62.2) kHz, duration: 2.0 ± 0.40 (1.6 - 2.6) msec. At Farm Welgevonden, South Africa, Taylor et al. (2013a: 556) report a Fknee value of 58 (48 - 68) kHz. Four calls recorded at Mapungubwe National Park (RSA) by Parker and Bernard (2018: 57) had the following values: Fchar: 42.68 ± 0.70 kHz, Fmax: 70.40 ± 3.85 kHz, Fmin: 40.88 ± 1.98 kHz, Fknee: 52.68 ± 3.43 kHz, duration: 3.03 ± 0.70 msec and 11.20 ± 7.85 calls/sec. Release calls recorded at Telperion Nature Reserve by Kearney et al. (2019: 37) had a Fchar: 37.04 kHz, slope at the start of the call: 372.62, time from start of call to Fchar: 2.62 msec and duration 3.13 msec. From Swaziland, Monadjem et al. (2017c: 179) reported Fmin: 38.0 ± 3.78 (34.2 - 41.7) kHz, Fknee: 55.4 ± 10.8 (44.6 - 66.2) kHz, Fc: 49.4 ± 11.4 (38.0 - 60.8) kHz, duration: 2.4 ± 0.33 (2.07 - 2.72) msec. The maximum distance over which they were able to detect the calls was 10 m (mean: 4.6 ± 0.73 m). Adams and Kwiecinski (2018: 4) reported Fchar: 41.8 ± 4.3 kHz, Fmax: 81.5 ± 6.2 kHz, Fmin: 38.