CSIRO PUBLISHING Invertebrate Systematics, 2011, 25, 490–585 http://dx.doi.org/10.1071/IS10016 Cladistic analysis and revision of Piestus Gravenhorst with remarks on related genera (Coleoptera : Staphylinidae : Piestinae) Edilson Caron A,D, Cibele S. Ribeiro-Costa B and Alfred F. Newton C A Campus Palotina, Universidade Federal do Paraná, Rua Pioneiro, 2153, Jardim Dallas, 85950-000, Palotina, PR, Brazil. B Laboratório de Sistemática e Bioecologia de Coleoptera (Insecta), Departamento de Zoologia, Universidade Federal do Paraná, Centro Politécnico, Caixa Postal 19020, 81531-980, Curitiba, PR, Brazil. C Zoology Department, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, IL 60605, USA. D Corresponding author. Email: caron@ufpr.br Abstract. Rove beetles of the genus Piestus Gravenhorst, 1806 are commonly captured under the bark of or inside decaying logs from Neotropical forests. Piestus belongs to the subfamily Piestinae, historically an ill-defined dumpingground for Staphylinidae defined by plesiomorphic characters, but which has gradually been restricted in concept and currently includes only six additional extant genera worldwide. Piestinae in this restricted sense has been considered a probably monophyletic subfamily, but its status and phylogenetic position, as a possible sister-group of Osoriinae within the recently proposed Oxyteline group of staphylinid subfamilies, are uncertain and need confirmation. The main aim of the present study was to provide a morphological cladistic analysis and complete taxonomic revision of Piestus, which, as the type and most speciose genus of Piestinae, is critical for future phylogenetic studies involving the subfamily. In our study, the monophyly of Piestus is established and phylogenetic relationships among its species are proposed based on 70 adult morphological characters. Piestus is supported by 11 synapomorphies and high branch support. All species of Piestus are revised and the genus is redefined. The genus contains 43 species, including 13 species described here for the first time. The previously proposed subgenera Antropiestus Bernhauer, 1917, Eccoptopiestus Scheerpeltz, 1952, Elytropiestus Scheerpeltz, 1952, Lissopiestus Scheerpeltz, 1952, Piestus s. str., Trachypiestus Scheerpeltz, 1952 and Zirophorus Dalman, 1821 have not been confirmed, as they were found to be poly- or paraphyletic, or are here removed from Piestus, and therefore subgenera are not used. The main taxonomic changes are as follows. Lissopiestus, syn. nov. is proposed as new synonym of Eleusis Laporte, 1835 and its species, E. interrupta (Erichson, 1840), comb. rest., is transferred again to that genus. Antropiestus, syn. nov. and Eccoptopiestus, syn. nov. are proposed as new synonyms of Hypotelus Erichson, 1839 and their species, H. laevis (Solsky, 1872), comb. nov. and H. andinus (Bernhauer, 1917), comb. nov., are transferred to Hypotelus. Fourteen new synonymies are proposed (valid species listed first): P. lacordairei Laporte, 1835 = Z. furcatus Sharp, 1887, syn. nov.; P. capricornis Laporte, 1835 = P. frontalis Sharp, 1876, syn. nov.; P. pennicornis Fauvel, 1864 = P. plagiatus Fauvel, 1864, syn. nov.; P. rectus Sharp, 1876, syn. nov.; P. pygialis Fauvel, 1902, syn. nov.; P. surinamensis Bernhauer, 1928, syn. nov.; P. minutus Erichson, 1840 = P. nigrator Fauvel, 1902, syn. nov.; P. sulcatus Gravenhorst, 1806 = P. sanctaecatharinae Bernhauer, 1906, syn. nov.; P. condei Wendeler, 1955, syn. nov.; P. gounellei Fauvel, 1902 = P. wasmanni Fauvel, 1902, syn. nov.; P. mexicanus Laporte, 1835 = P. alternans Sharp, 1887, syn. nov.; P. aper Sharp, 1876 = P. schadei Scheerpeltz, 1952, syn. nov.; P. angularis Fauvel, 1864 = P. crassicornis Sharp, 1887, syn. nov.; H. andinus (Bernhauer, 1917) = P. strigipennis Bernhauer, 1921, syn. nov. One species is revalidated: P. fronticornis (Dalman, 1821), stat. rev., and one synonym is restored: P. penicillatus (Dalman, 1821) = P. erythropus Erichson, 1840, syn. rest. Neotypes are designated for P. lacordairei Laporte, 1835 and Oxytelus bicornis Olivier, 1811, and lectotypes are designated for P. puncticollis Fauvel, 1902, P. capricornis variety muticus Fauvel, 1902, P. zischkai Scheerpeltz, 1951, P. pennicornis Fauvel, 1864, P. plagiatus Fauvel, 1864, P. pygmaeus Laporte, 1835, P. niger Fauvel 1864, P. minutus Erichson, 1840, P. nigratror Fauvel, 1902, P. sulcatus Gravenhorst, 1806, P. sanctaecatharinae Bernhauer, 1906, P. sulcipennis Scheerpeltz, 1952, P. aper Sharp, 1876, P. schadei Scheerpeltz, 1952 and P. andinus Bernhauer, 1917. Received 24 May 2010, accepted 20 December 2011, published online 7 May 2012 Journal compilation
CSIRO 2011 www.publish.csiro.au/journals/is Revision of Piestus with remarks on related genera Introduction The beetle family Staphylinidae, or rove beetles, is the most diverse family of animals known, with more than 55 400 described species (Grebennikov and Newton 2009). Piestinae Erichson, 1839 is one of the oldest of the 32 currently recognised extant staphylinid subfamilies, but historically has been poorly defined and used as a repository for a diverse assortment of rove beetles that do not fit well elsewhere; Bernhauer and Schubert (1910), for example, included nearly 30 genera in their concept of the group (as the tribe Piestini). Gradually, many of those genera have been removed to form groups of their own (Apateticinae Fauvel, 1895; Trigonurinae Reiche, 1865) or added to other well-defined subfamilies (Blackwelder 1942; Newton 1982a, 1982b), so Piestinae in the modern sense includes only seven extant genera (see below). Piestinae are now allocated within the Oxyteline group sensu Lawrence and Newton (1982) and Thayer (2005) together with another five subfamilies: Apateticinae, Osoriinae Erichson, 1839, Oxytelinae Fleming, 1821, Scaphidiinae Latreille, 1807 and Trigonurinae. Thayer (2005) noted that Piestinae in the current sense is probably a monophyletic group and sister group of Osoriinae, but no cladistic analysis has been presented for corroborating its monophyly, as well as the relationship with Osoriinae. Cladistic studies within the Oxyteline group sensu Thayer (2005) have been performed only for Scaphidiinae (Leschen and Löbl 1995, 2005), Oxytelinae (Herman 1970, 1977; Newton 1982b) and Osoriinae in part (Wu and Zhou 2007). Invertebrate Systematics Piestinae currently includes 110 species placed in eight genera: Abolescus Tikhomirova, 1968, one Jurassic fossil species found in Karatau, Kazakhstan; Eupiestus Kraatz, 1859, 21 species in the eastern Palearctic and Oriental regions; Hypotelus Erichson, 1839, eight species in the Nearctic and Neotropical regions; Parasiagonum Steel, 1950, one species from New Zealand; Piestoneus Sharp, 1889, five species in the eastern Palearctic region; Piestus Gravenhorst, 1806, 49 species in the Nearctic and Neotropical regions; Prognathoides Steel, 1950, one species from Australia; and Siagonium Kirby & Spence, 1815, 24 species in the Holarctic and Neotropical regions (Herman 2001b; Zheng 2004; Naomi 2006; Khachikov 2007). Piestus is the most speciose genus of Piestinae and has been divided into seven subgenera: Antropiestus Bernhauer, 1917, two species; Eccoptopiestus Scheerpeltz, 1952, one species; Elytropiestus Scheerpeltz, 1952, one species; Lissopiestus Scheerpeltz, 1952, one species; P. s. str. Gravenhorst, 1806, 24 species; Trachypiestus Scheerpeltz, 1952, nine species; and Zirophorus (Dalman, 1821), 11 species (Scheerpeltz 1952; Wendeler 1955; Herman 2001b). A list of all previously known valid species organised by current subgenera is given in Table 1. Most species of the genus are saprophagous and are commonly found associated with decaying logs or in leaf litter in Neotropical forests. The general purpose of this study was to initiate a modern phylogenetic study of Piestinae and its place within Staphylinidae by focusing on the monophyly and phylogenetic relationships of Piestus, the type genus and by far the most speciose genus Table 1. Subgenera and valid species of Piestus prior to the current study (Scheerpeltz 1952, Wendeler 1955, Herman 2001b) P. (Antropiestus) Bernhauer, 1917 P. (A.) andinus Bernhauer, 1917 P. (A.) strigipennis Bernhauer, 1921 P. (Eccoptopiestus) Scheerpeltz, 1952 P. (E.) laevis Solsky, 1872 P. (Elytropiestus) Scheerpeltz, 1952 P. (E.) paradoxus Bernhauer, 1917 P. (Lissopiestus) Scheerpeltz, 1952 P. (L.) interruptus (Erichson, 1840) P. (Piestus) Gravenhorst, 1806 P.(P.) alternans Sharp, 1887 P.(P.) buquetii Fauvel, 1864 P.(P.) condei Wendeler, 1955 P.(P.) erythropus Erichson, 1840 P.(P.) extimus Sharp, 1887 P.(P.) filicornis Fauvel, 1902 P.(P.) fulvipes Erichson, 1840 P.(P.) gounellei Fauvel, 1902 P.(P.) heterocephalus Fauvel 1902 P.(P.) mexicanus Laporte, 1835 P.(P.) minutus Erichson, 1840 P.(P.) niger Fauvel, 1864 P.(P.) nigrator Fauvel, 1902 P.(P.) penicillatus (Dalman, 1821) P.(P.) pennicornis Fauvel, 1864 P.(P.) plagiatus Fauvel, 1864 P.(P.) puncticollis Fauvel, 1902 P.(P.) pygialis Fauvel, 1902 491 P.(P.) pygmaeus Laporte, 1835 P.(P.) rectus Sharp, 1876 P.(P.) sanctaecatharinae Bernhauer, 1906 P.(P.) sulcatus Gravenhorst, 1806 P.(P.) Surinameensis Bernhauer, 1928 P.(P.) wasmanni Fauvel, 1902 P. (Trachypiestus) Scheerpeltz, 1952 P.(T.) angularis Fauvel, 1864 P.(T.) aper Sharp, 1876 P.(T.) chiriquensis Sharp, 1887 P.(T.) costatus Sharp, 1887 P.(T.) crassicornis Sharp, 1887 P.(T.) nevermanni Scheerpeltz, 1952 P.(T.) rugosus Sharp, 1876 P.(T.) schadei Scheerpeltz, 1952 P.(T.) sulcipennis Scheerpeltz, 1952 P. (Zirophorus) (Dalman, 1821) P.(Z.) bicornis (Olivier, 1811) P.(Z.) capricornis Laporte, 1835 P.(Z.) frontalis Sharp, 1876 P.(Z.) furcatus (Sharp, 1887) P.(Z.) lacordairei Laporte, 1835 P.(Z.) longicornis (Lacordaire, 1833) P.(Z.) longipennis Fauvel, 1864 P.(Z.) planatus (Sharp, 1887) P.(Z.) spinosus (Fabricius, 1801) P.(Z.) validus Sharp, 1876 P.(Z.) zischkai Scheerpeltz, 1951 492 Invertebrate Systematics of the subfamily. Therefore, the current work provides a morphological cladistic analysis using adults of all species of Piestus as well as selected outgroup members of all other extant genera of Piestinae, and a taxonomic revision of Piestus, including the redefinition of the genus, descriptions and illustrations of new and previously known species, and a key to all valid species. Material and methods Phylogenetic methods Terminals In total, 45 species of Piestus were included as ingroup, of which 32 are listed in Herman (2001b) and considered as valid in the present revision (see ‘Results’, ‘Taxonomic revision’), plus 13 new species. Piestus interruptus, P. fulvipes and P. longicornis were not included in analysis (for details see ‘Results’, ‘Taxonomic revision’). Given that the relationships among the genera of Piestinae are unknown, the following species (representing all other extant genera of Piestinae) were used as outgroups: Eupiestus sculpticollis Kraatz, 1859 (type species of the genus); Hypotelus pusillus Erichson, 1840 (type species of the genus); Parasiagonum hudsoni (Cameron, 1927) (monotypic genus); Piestoneus monticola Naomi, 1995; Prognathoides mjobergi (Bernhauer, 1920) (monotypic genus); and Siagonium punctatum (LeConte, 1866). Character construction In the cladistic analysis adult morphological characters from the exoskeleton and some characters from male and female genitalia were used and are listed below (see Appendix 1). All multistate characters were considered as unordered. The matrix was edited by the NEXUS program, version 0.5.0 (Page 2001). Missing data were coded as ‘?’ and the inapplicable data as ‘–’ (Table 2). Analysis Fitch’s parsimony (Fitch 1971) was adopted for optimisation criteria. The search for the most parsimonious cladograms was generated in the TNT program, version 1.1 (Goloboff et al. 2008), where the characters were treated as of equal weighting and the following settings were used in the analysis: ‘hold 30000;’, 1000 replications, 10 trees saved per replication, and tree bisection reconnection (TBR) as branch swapping algorithm. The Winclada program, version 0.9.9 (Nixon 1999) was used for representing the cladograms. The trees were rooted ‘a posteriori’ (Nixon and Carpenter 1993) in Hypotelus pusillus. Bremer support values (Bremer 1994) were used as a measure of branch support and computed in the TNT program, version 1.1 (Goloboff et al. 2008), using ‘trees suboptimal’ to 10 steps and the same settings used in the equal weighting (see above). In a subsequent analysis, the characters were treated using implied weighting (Goloboff 1993) in the TNT program, version 1.1 (Goloboff et al. 2008), using k = 3 and the same settings used in the equal weighting analysis. This method resolves conflict in favour of less homoplastic characters and was used here as an alternative estimation of phylogeny using the same data. E. Caron et al. Taxonomic revision Material examined In the current study ~5000 adult specimens of Piestinae were examined. The abbreviations cited below for each institution or private collection are used in all text citations. The name of the responsible curator or individual is given in parentheses: AMBC, Private Collection Ayr Bello, Rio de Janeiro, Brazil (A. Bello); AMNH, American Museum of Natural History, New York, USA (L. H. Herman); BMNH, The Natural History Museum, London, United Kingdom (R. G. Booth); CZUG, Centro de Estudios en Zoología, Entomología, Universidad de Guadalajara, Guadalajara, México (J. L. Navarrete-Heredia); DZUP, Coleção de Entomologia Pe. J. S. Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil (L. M. de Almeida); FMNH, Field Museum of Natural History, Chicago, USA (A. F. Newton); INPA – Coleção Sistemática de Entomologia, Instituto Nacional de Pesquisas da Amazônia, Manaus, Brazil (A. Henriques); IRSNB, Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (Y. Gérard); MNRJ, Museu Nacional, Universidade do Rio Janeiro, Rio de Janeiro, Brazil (M. A. Monné); MUSM, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru (G. Lamas); MZSP, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (S. A. Casari); NHRS, Naturhistoriska Riksmuseet, Stockholm, Sweden (B. Viklund); NMW, Naturhistorisches Museum Wien, Wien, Austria (H. Schillhammer); SEMC, Snow Entomological Museum, University of Kansas, Lawrence, USA (Z. Falin); ZMHB, Museum für Naturkunde der HumboldtUniversität, Berlin, Germany (J. Frisch); and ZMUC, Zoological Museum, University of Copenhagen, Copenhagen, Denmark (O. Martin). Type material was examined for all names listed within Piestus by Herman (2001b), except for P. bicornis (Olivier, 1811) (lost, neotype here designated), P. oxytelinus Perty, 1830 (not found, junior synonym of P. spinosus), P. longicornis (Lacordaire, 1833) (not found), P. lacordairei Laporte, 1835 (lost, neotype here designated), P. fulvipes Erichson, 1840 (syntype in BMNH), P. validus Sharp, 1876 (syntype in BMNH), Trichocoryne striata Gray, 1832 (not found, junior synonym of P. penicillatus) and Zirophorus striatus Guérin-Méneville, 1829 (not found, junior synonym of P. bicornis). For more details, see ‘Results’ in ‘Taxonomic revision’. Labels from type material are organised in sequence from top to bottom, where the data from each label are enclosed within double quotes (‘ ’), a forward slash (/) separates lines, and information enclosed by square brackets ([]) provides added details about the labels. Information from labels of additional material is organised, when complete, as follows: country: district/state/province, number of specimens, locality, extra information, date, collector (abbreviation of institution). All information from labels is listed as found, with added details enclosed by square brackets ([]) and comments in italics. Distribution of each species is listed in the text by country and followed by, when possible, district/state/province enclosed by parentheses ‘()’. Biological notes about each species are cited from labels and also from cited literature. Revision of Piestus with remarks on related genera Invertebrate Systematics 493 Table 2. Data matrix for Piestus (?) missing data, (-) inapplicable characters Taxa Hypotelus pusillus Eupiestus sculpticollis Piestus andinus Piestus laevis Siagonium punctatum Parasiagonum hudsoni Piestoneus monticola Prognathoides mjobergi Piestus heterocephalus P. lacordairei P. convexus, sp. nov. P. similis, sp. nov. P. capricornis P. puncticollis P. planatus P. spinosus P. longipennis P. zischkai P. bicornis P. fronticornis P. validus P. formicinus, sp. nov. P. termitis, sp. nov. P. surrufus, sp. nov. P. pennicornis P. rossii, sp. nov. P. pygmaeus P. buquetii P. extimus P. filicornis P. abdominalis, sp. nov. P. minutus P. niger P. penicillatus P. sulcatus P. gounellei P. mexicanus P. ecuadorensis, sp. nov. P. foveolatus, sp. nov. P. sulcipennis P. paradoxus P. nevermanni P. costatus P. chiriquensis P. imperfectus, sp. nov. P. boliviensis, sp. nov. P. angularis P. aper P. rugosus P. acuminatus, sp. nov. P. elegans, sp. nov. 0000000000 0123456789 1111111111 0123456789 2222222222 0123456789 Characters 3333333333 0123456789 4444444444 0123456789 5555555555 0123456789 6666666666 0123456789 0––0000100 0––00001–0 0––0000130 111–000100 100–000100 0––000010? 0––0000100 100–0001–0 100–000010 100–000010 110–00003? 110–00003? 110–000010 0––001002? 100–010010 100–010010 101–010020 101–010020 101–010020 101–010020 101–010020 0––0000111 0––0010100 0––0010010 0––0000021 0––000012? 0––0010121 0––1010121 0––1010021 0––1111021 0––1111021 0––1111021 0––1111021 0––1111021 0––0000111 0––0010211 0––0010211 0––0010211 0––0010211 0––0010211 0––0010201 0––0010201 0––0010201 0––0010201 0––0000201 0––001020? 0––0010201 0––0000200 0––0010201 0––0010201 0––00102?? 000000000– 000000000– 000000000– 0000000011 0000000011 ?000?00011 10000??011 1000000010 0001010111 0001010111 1?01010111 00010?0111 1001000011 1??10??011 1001010111 1001010111 0001010111 0001010111 0001010111 0001010111 0001010111 0111000010 000100100– 011100100– 1211100010 1??1100010 1221100010 1221100010 1121100010 1121100010 1221100010 1221100010 1221100010 1221100010 1111000010 1111000010 1111000010 1111000010 1111000010 1001000010 1101000010 11010??010 1111000010 1111000010 1111000010 ?1010??010 1001000010 0001000010 1101000010 1101000010 ???10??010 –100–00011 –100–00011 –100–00011 0100–00011 1100–00000 0400–00000 0400–00000 –401000000 0111011000 0111011000 01???11?00 01???11?00 0111011000 01???11?00 1111011000 1111011000 0211011000 0211011000 0211011000 0211011000 0211011000 –111101100 –011101100 –011101100 –111101100 –1???01?00 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –111101100 –311101100 –1???01?00 –111101100 –311101100 –311101100 –1???01?00 –311101100 –011101100 –311101100 –311101100 –3???01?00 1001110000 1001100100 1001110000 1001110000 1111110001 1211110000 1211110001 1211110001 0101011001 0101011001 0100011001 0100011001 0111011101 0111011101 0111011101 0111011101 0101011001 0101011001 0101011001 0101011001 0101011001 0100110001 0102210101 0102010101 0101011001 0100111001 0101111001 0101111001 0101011001 0101011001 0101011001 0101011001 0101011001 0101011001 0100110111 0100210111 0100210111 0100210111 0100210111 0102200111 0102200111 0102200111 0102200111 0102200111 0102200111 0102200111 0102200111 0112200111 0102200111 0102200111 0102200111 0000–00–00 0000–01–12 0000–00–00 0000–00–00 0000–02001 0000–02000 0000–02001 0000–13011 0101012100 0101012100 0101012100 0101012100 0101012101 0101012101 0101012101 0101012101 0101112100 0101112100 0101112100 0101112100 0101112100 0101013000 0101012100 0101012100 1101012100 1101012100 0101012100 0101012100 0101012100 0101012100 0101012100 0101012100 0101012100 0101012100 1101013112 1101013112 1101013012 1101013012 1101013012 1101013112 1101013112 1101013012 1101013012 1101013012 1101013012 1111013012 1111013012 0111013012 1111013012 1111013012 ?111013012 0––0001000 1010101000 0––0001000 0––0001000 0000011000 0000010000 0000011000 0000110000 0000000110 0000000110 0000000110 0000000110 0000100110 0000100110 0000100110 0000100110 0000000111 0000000111 0000000111 0000000111 0000000111 0000000010 0100000010 0100000010 0000000110 0000000110 0000000110 0000000110 0000000110 0000000110 0000000110 0000000110 0000000110 0000000110 0000100010 0000100010 0000100010 0000100010 0000100010 0000100010 0011100010 0010100010 0010100010 0010100010 0000100010 0010100010 0010100010 0010100010 1010100010 1011100010 1010100010 1111000000 111????00? 111100000? 111100000? 0111000000 0111000000 011100000? 011100000? 0001000101 0001000101 0001100??? 0001100??? 0001000101 0001000??? 0001000101 0001000101 0001010111 0001010111 0000011110 0000011111 0000011111 0001100100 0001100100 0001100100 0001000100 0001100??? 0001100101 0001100101 0001100101 0001100101 0001100101 0001000101 0001000101 0001000101 0001100100 0001100101 0001100100 0001100100 0001100100 0001100100 0001100100 0001100100 0001100100 0001100100 0001100100 0001100?0? 0001100100 0001100100 0001100100 0001100100 000????100 The designation of lectotypes and neotypes follows ICZN (1999), which recommends that lectotype designation ‘should be done as part of a revisionary or other taxonomic work to enhance the stability of nomenclature’ (Declaration 44, Amendment of Article 74.7.3; ICZN 1999), and for neotype ‘with the express purpose of clarifying the taxonomic status or the type locality of a nominal taxon’ (Article 75.3.1; ICZN 1999). 494 Invertebrate Systematics Morphology The morphological terminology adopted is basically from Naomi (1987–1990) and Caron et al. (2008). The numbering of abdominal segments is according to morphological origin and not indicating visible segments. Gusarov (2002) was used for describing the position of the aedeagus. Measurements were done using a micrometric ocular in a stereoscopic microscope Wild M5 or Leica MZ12.5 and follow Caron et al. (2008), in which the following abbreviations are used: BL, body length (from anterior margin of head capsule to posterior margin of tergite 8); BW, body width (across humeral region); PL, pronotum length (maximum); PW, pronotum width (maximum); EL, elytron length (maximum). Most features were observed from dried pinned specimens. Apical abdominal segments, including genitalia, were studied for all species after dissection. Mouthparts were dissected and studied for the great majority of the species, except Piestus puncticollis, P. convexus, sp. nov., P. similis, sp. nov., P. rossii, sp. nov., P. boliviensis, sp. nov. and P. elegans, sp. nov. because only holotypes or syntypes for these species have been available. The specimens were dissected following Caron et al. (2008). For type specimens, cold ammonia, NH3(aq), was used for five minutes. The dissections were carried out under a Zeiss Stemi SV6 or Leica MZ12.5 stereoscopic microscope. The dissected parts were put inside micro vials with glycerin or on plastic board covered with Canada balsam, both pinned with the specimen. Drawings were made under a stereoscopic microscope, Zeiss Stemi SV6 or Leica MZ12.5, and also with a compound microscope, Zeiss Standard 20 or Wild M20, all with a drawing tube attached. The final drawings were produced as in Caron et al. (2008). Photographs of most of the species were taken as in Caron et al. (2008), except for Piestus puncticollis, P. interruptus, P. laevis and P. andinus, which were taken using a Microptics ML Macro XLT digital imaging system. The final photographs were edited as in Caron et al. (2008). Phylogenetic analysis The cladistic analysis using equal weighting generated eight equally parsimonious cladograms (Length, L = 170; Consistency Index, CI = 0.49; Retention Index, RI = 0.88) (Fig. 1). The strict consensus tree is shown in Figs 2 and 3 (L = 176, CI = 0.47, RI = 0.87). Species of Piestus form a distinct monophyletic group, excluding P. andinus and P. laevis, both here transferred to Hypotelus (see ‘Taxonomic revision’ below) (Fig. 2). Within Piestus there are four major clades (A, B, C and D), and two different hypotheses concerning the relationships among Piestus formicinus and the clades C and D. In hypothesis 1, P. formicinus is sister group of clade C+D (Fig. 1; H1); in hypothesis 2, the clade C is sister group to P. formicinus + clade D (Fig. 1; H2). The other topologies concern the relationships among three species of the outgroup (Piestoneus monticola, Prognathoides mjobergi and Parasiagonum hudsoni) (Fig. 1; H1, H3) and the relationships within clade A (Fig. 1; H1, H2) and within clade C (Fig. 1; H1, H2). The analysis utilising implied weighting confirmed the monophyly of Piestus, excluding P. andinus and P. laevis, and E. Caron et al. showed a similar topology, with a few differences among the relationships of internal terminals (Fig. 4). This analysis supports the hypothesis of clade C as a sister group of P. formicinus + clade D. According to Scheerpeltz’s classification, Piestus contained seven subgenera (Table 1) (Scheerpeltz 1952). In the current analysis, the species of the subgenus Antropiestus, P. andinus and the species of the subgenus Eccoptopiestus, P. laevis, are excluded from Piestus. The species of the subgenera Zirophorus, Piestus s. str., Elytropiestus and Trachypiestus are discussed below. The discussions of characters are based on the results of the strict consensus cladogram from equal weighting (Figs 2, 3). When pertinent, the characters from strict consensus with implied weighting are also discussed. Piestus is a strongly monophyletic group supported by 13 unambiguous changes (Fig. 3) (Bremer support, >10; Fig. 2), of which 11 are synapomorphies: two moderately sized punctures with setae on the basal half of the dorsal margin of the eyes (Fig. 86); antennomere 4 densely covered with microsetae (Fig. 106); openings of glands at the external margin of mandibles (Fig. 114); maxillary palpomere 3 as wide as long (Fig. 131); absence of pair of conspicuous setae on apical margin of the ligula, near the median sclerotised plate (Fig. 133); carinate mesoventrite process (Fig. 9); groove on ventral margin of the anterior femur (Fig. 139); very small emargination on each posterolateral area on sternite 6, external openings of abdominal defensive gland complex (Fig. 11) (Caron et al. 2008); absence of fringe on apical margin of tergite 10 (Fig. 149); one or two pairs of setae at the apex of tergite 10 (Fig. 149); and one or two setae on the apex of the ovipositor of females (Fig. 153). The analysis suggested Piestus is closely related to the clade Siagonium punctatum + (Parasiagonum hudsoni+Piestoneus monticola + Prognathoides mjobergi) as supported by six synapomorphies (Fig. 3) (Bremer support, 6; Fig. 2): not emarginate anterior angles of the mentum; absence of pair of conspicuous setae at the apex of the median sclerotised plate of the ligula; narrowly separated gular sutures (Fig. 8); anterior margin of the prosternum anteriorly projected at the middle (Fig. 8); five complete longitudinal striae on each elytron (Fig. 5); and absence of short setae on apical margin of sternite 8 of the female. However, a specific phylogenetic study is necessary to confirm this hypothesis. Clade A Within Piestus, clade A comprises 13 species that share seven unambiguous changes, two of which are synapomorphies (Fig. 3) (Bremer support, 4; Fig. 2): long epipharynx visible dorsally, ~2.5 times or so the median length of labrum (Fig. 110); and maxillary palpomere 4 shorter than 2 and 3 combined (Fig. 130). This clade is similar to the species group proposed as the subgenus Zirophorus by Scheerpeltz (1951, 1952), except for P. heterocephalus and P. puncticollis, which were both previously allocated in the subgenus Piestus s. str. (Table 1). Scheerpeltz (1952) grouped the species within P. (Zirophorus) based mainly on the presence of the frontal process on the head and mandibles anteriorly projected. The Revision of Piestus with remarks on related genera Invertebrate Systematics Fig. 1. Three of eight equally parsimonious cladograms showing relationships among species of Piestus using equal weighting of characters. H1, H2 and H3: hypothesis 1, 2 and 3, respectively. 495 496 Invertebrate Systematics E. Caron et al. Clade A Clade B Clade C Clade D Fig. 2. Strict consensus cladogram of eight equally parsimonious trees using equal weighting of characters. Bremer support values shown below branches. frontal process (Fig. 22) occurs in other groups of Piestinae, such as Hypotelus (H. laevis, see ‘Taxonomic revision’) (Fig. 56), Siagonium (S. punctatum) and Prognathoides (P. mjobergi), and thus seems to be a plesiomorphic character, and it is not found in P. puncticollis (Fig. 16). The other character indicated by Scheerpeltz (1952) was observed in the Revision of Piestus with remarks on related genera Invertebrate Systematics Clade A Clade B Clade C Clade D Fig. 3. Strict consensus cladogram of eight equally parsimonious trees using equal weighting of characters. Only unambiguous changes are shown. Black square, synapomorphies; white square, homoplasies. 497 498 Invertebrate Systematics E. Caron et al. Clade A Clade B Clade C Clade D Fig. 4. Strict consensus cladogram of three using implied weighting of characters (L = 174, CI = 0.48, RI = 0.87). Only unambiguous changes are shown. Black square, synapomorphies; white square, homoplasies. current analysis as a homoplasy for clade A, not found in P. capricornis and P. puncticollis (Figs 63, 64). The implied weighting scheme showed a different topology among the terminals within clade A (compare Fig. 1, H1, H2, and Fig. 4). Furthermore, the current characters used with equal weighting do not produce a good resolution at the base of clade A, in which there is a polytomy (Fig. 3). However, both weighting schemes confirmed two different clades: clade P. convexus + P. similis and clade P. longipennis + P. zischkai + (P. bicornis + P. fronticornis + P. validus). The clade P. convexus + P. similis was supported only by homoplasies with equal weighting (Fig. 3) (Bremer support, 8, Fig. 2). The homoplasies were: short frontal process on head (Fig. 13); line with more than five moderately sized punctures with setae; curved sides of pronotum (Fig. 59); and curved apex of the median lobe of the aedeagus (Fig. 158). The clade Revision of Piestus with remarks on related genera P. longipennis + P. zischkai + (P. bicornis + P. fronticornis+ P. validus) was supported by three homoplasies and five synapomorphies (Fig. 3) (Bremer support, 7; Fig. 2). The following synapomorphies were observed: two acute teeth on the internal border of the right mandible, not close (Fig. 121); long stout setae at the groove on ventral margin of the anterior femur; longitudinal set of long setae near each lateral margin on sternite 7; emarginate apex of the median lobe of the aedeagus of the male (Fig. 181); and emarginate external margin of the ovipositor of the female (Fig. 154). The clade P. capricornis + P. puncticollis+(P. planatus + P. spinosus) was also supported in equal weighting only by homoplasies (Fig. 3) (Bremer support, 2; Fig. 2): long antennae of male; projected anterior angles of pronotum (Fig. 65); a slight emargination in each side of pronotum, anterior to basal constriction (Fig. 65); fine punctures on striae of elytra; and sternite 3 with conspicuous transverse carina. This clade was not confirmed in implied weighting (Fig. 4). Clade B+(P. formicinus + clade C+D) This clade is supported by one homoplasy and two synapomorphies (Fig. 3) (Bremer support, 2; Fig. 2), the synapomorphies being: strongly acute small basal tooth on external margin of the mandibles (Fig. 113); and subquadrate mentum (Fig. 135). The topology of the clade P. formicinus + clades C+D is supported in equal and implied weightings (Figs 3, 4) only by character 9 as a unique change: the antennal length in the male conspicuously longer than in the female. This character is not found in P. aper (Table 2). The clade P. formicinus + clades C+D has low branch support (Bremer support, 1; Fig. 2). In the implied weighting, clade C is sister group of P. formicinus + clade D. The clade P. formicinus + clade D is supported by two homoplasies: curved sides of the pronotum (Fig. 87), and six complete longitudinal striae on each elytron. This topology appears in one of the most parsimonious trees estimated using equal weighting (Fig. 1; H2). Clade B Clade B has only two species and is strongly supported by four homoplasies and two synapomorphies (Fig. 3), the latter being: two lobes at each apex of the epipharynx (Fig. 111); and undulate microstriae on integument of the interstriae of the elytra. Clade B has a branch support value of 4 (Fig. 2). Invertebrate Systematics 499 occurs in most of the species of clade D, except in P. aper (Table 2). This clade also occurs in equal weighting (Fig. 1; H2). The clade P. pygmaeus + (P. buquetii + (P.extimus + (P. filicornis + P. abdominalis + (P. minutus + P.niger + P. penicillatus)))) is supported by two homoplasies and one synapomorphy (Fig. 3) (Bremer support, 2; Fig. 2): enlarged area on basal half of exposed dorsal face of the antennal scape of males and females. The clade P. filicornis + P. abdominalis + (P. minutus + P.niger + P. penicillatus) is supported by two synapomorphies (Fig. 3) (Bremer support, 3; Fig. 2): a slightly concave region at the middle of the front; and a transverse carina at the base of the vertex (Fig. 82). Clade C contains most of the species grouped by Scheerpeltz (1952) in the subgenus Piestus s. str. (Table 1). The rest of the species grouped in Piestus s. str. occur in clade A (P. heterocephalus and P. puncticollis) and in clade D (P. sulcatus, P. gounellei and P. mexicanus) (Fig. 2). Clade D Clade D is the largest clade within Piestus and contains 17 species, which share four homoplasies and one unique synapomorphy (Fig. 3): a small lateral tooth on the side of the pronotum, anterior to the basal constriction (Fig. 7). However, clade D has good branch support (Bremer support, 6; Fig. 2). The clade P. sulcipennis + (P. paradoxus + (P. nevermanni + (P. costatus + (P. chiriquensis + P. imperfectus)) + (P. sp. n. L.+ (P. angularis + P. aper) + (P. rugosus + P. acuminatus+ P. elegans))))) is distinct and shares three homoplasies (Fig. 3) (Bremer support, 3; Fig. 2): enlarged area on basal half of exposed dorsal face of the antennal scape only in males; sinuous sides of pronotum (Fig. 100); and inconspicuous median longitudinal sulcus on pronotum (Fig. 92). All species of this clade were treated as the subgenera Trachypiestus and Elytropiestus by Scheerpeltz (1952), based on large punctures on the pronotum (Table 1). However, similar punctures occur in Eupiestus sculpticollis, Piestus gounellei, P. mexicanus, P. ecuadorensis and P. foveolatus. Within clade D, the topologies remain almost the same between the different weighting schemes (compare Figs 3 and 4). Thus, clade D is a mixture of species from the subgenera Piestus s. str., Trachypiestus and Elytropiestus of Scheerpeltz (1952). Conclusions Clade C Clade C contains 10 species and is supported by three homoplasies and two synapomorphies (Fig. 3) (Bremer support, 3; Fig. 2): long setae on basal half of exposed dorsal face of the antennal scape of the male and female, and antenna conspicuously narrowing from antennomere 4–11 (Fig. 35). Within clade C the topologies remain almost the same between the different weighting schemes, except for P. pennicornis and P. rossii (compare Figs 3 and 4). The clade P. pennicornis + P. rossii appears in implied weighting, being supported by one homoplasy (Fig. 4): conspicuous ovate set of fine punctures in median region near the apex of the prosternum of male (Fig. 8). This character The genus Piestus is a monophyletic group, excluding the species P. laevis and P. andinus, both here transferred to Hypotelus (see ‘Taxonomic revision’). Therefore, the genus Piestus contains 43 species, including 13 new species, and is supported by 11 synapomorphies (Fig. 3) and high branch support (Bremer support, >10; Fig. 2). In the current analysis, the most recent and regularly adopted classification of Scheerpeltz (1952) (Table 1) has not been entirely confirmed, as his subgenera are either poly- or paraphyletic or, in two cases, removed from Piestus. Thus, the subgenera established by Scheerpeltz (1952) are not recognised here. The genus Piestus has four evident clades in which most of the species are distributed. The only species excluded from the clades is P. formicinus, which appears to be related to clades C and 500 Invertebrate Systematics D. However, from implied weighting it is possible to suggest that P. formicinus is more closely related to clade D than C (Fig. 4). Taxonomic revision Genus Piestus Gravenhorst, 1806 Piestus Gravenhorst, 1806: 223 (original description); Laporte, 1835: 126 (list of species, characters, notes); Erichson, 1840: 830 (characters, notes); Sharp, 1876: 403 (list of species, characters, notes); Sharp, 1887: 712 (notes); Bernhauer & Schubert, 1910: 6 (list of species); Blackwelder, 1943: 43 (characters, notes); Blackwelder, 1944: 100 (list of species); Blackwelder, 1952: 309 (nomenclatural notes); Scheerpeltz, 1952: 281 (revision); Herman, 2001b: 1787 (bibliography, distribution, notes); Navarrete-Heredia et al., 2002: 207 (characters, notes, list of species); Newton et al., 2005: 37 (list of species from Colombia); Caron et al., 2008: 6 (abdominal defensive glands complex). Type species: Piestus sulcatus Gravenhorst, 1806 (fixed by monotypy). Zirophorus Dalman, 1821: 372 (original description); Laporte, 1835: 126 (synonym of Piestus). Type species: Zirophorus fronticornis Dalman, 1821 (fixed by subsequent designation by Crotch 1870). Piestus (Zirophorus): Scheerpeltz, 1951: 5 (key); Scheerpeltz, 1952: 282 (key); Herman, 2001b: 1787 (bibliography); Navarrete-Heredia et al., 2002: 207 (mention). Irenaeus Latreille, 1829: 438 (cited as a synonym of Zirophorus); Lacordaire, 1854: 127 (synonym of Piestus); Herman, 2001b: 1787 (synonym of Piestus, bibliography). Type species: cited as a synonym of Zirophorus, without type species. Note: nomen nudum (Herman 2001b). Trichocoryne Gray, 1832: 306 (original description); Herman, 2001b: 1788 (synonym of Piestus, bibliography). Type species: Zirophorus penicillatus Dalman, 1821 (fixed by subsequent designation by Blackwelder 1943). Note: not monotypy as cited by Blackwelder (1943, 1952) and Herman (2001b), because Gray also originally included T. striata Gray, which he indicated as a probable synonym of Z. penicillatus. Tricoryna Laporte, 1835: 126 (synonym of Piestus, attributed to Gray, but evident misspelling of Trichocoryne). Type species: Not applicable. Trachypiestus Scheerpeltz, 1952: 291 (as subgenus of Piestus); Herman, 2001b: 1788 (bibliography); Navarrete-Heredia et al., 2002: 207 (mention). Type species: Piestus angularis Fauvel, 1864 (fixed by original designation). Elytropiestus Scheerpeltz, 1952: 294 (as subgenus of Piestus); Herman, 2001b: 1788 (bibliography). Type species: Piestus paradoxus Bernhauer, 1917 (fixed by original designation and monotypy). Redescription BL: 2.6–15.8 mm, BW: 0.8–2.5 mm. Body flat dorsoventrally to somewhat convex (Fig. 6); light brown to entirely black, sometimes with all or only apical half of abdominal segment 7 and segments 8–10 yellowish; antennomere 11 slightly lighter than others; tarsi, maxillary and labial palpi lighter than body; setae on body golden or black. Body integument with undulate microstriae, easy to see on metaventrite (Figs 9, 10); fine to large punctures uniformly distributed, sometimes contiguous; elytral striae punctured or not. Male. Head transverse to subquadrate, front slightly deflected to entirely deflected (Figs 7, 82), some species with a pair of acute and short or long frontal processes (Fig. 22); E. Caron et al. V-shaped frontal sulcus generally conspicuous and complete, curved arms joined medially (Fig. 85); anterior angles curved and generally prominent; eyes somewhat prominent in dorsal view, and some moderately sized punctures with setae on basal half of dorsal margin (Fig. 7). Antennae long, reaching apex of elytra or abdomen (Figs 18, 22); scape the longest or subequal in length to antennomere 3 (Fig. 106); pedicel the shortest, antennomeres 4–11 increasing in length and generally oblong shape; scape slightly the widest, pedicel to antennomere 11 with same width or decreasing in width towards apex; scape with long setae distributed on basal half or on apical half of dorsal face (Fig. 106); pedicel and antennomere 3 with long setae generally distributed on apical half of dorsal face, and antennomeres 4–11 entirely with microsetae and some long dispersed setae; antennomeres 3–10 with apical circular set of long setae. Labrum transverse (Fig. 108), anterior margin emarginate, four to eight long setae medially; each lateral third, four long setae (two apical and two subapical). Epipharynx short or long (Figs 108, 110), deeply emarginate medially, long fringes on internal margin and short fringes on external margin. Mandibles strongly projecting or not (Figs 25, 26); internal border smooth or teethed, symmetrical or asymmetrical; dorsal teeth absent or present, sometimes weakly developed; prostheca developed, almost reaching apex of ventral tooth and ending in sulcus on ventral face (Fig. 114); mola conspicuous, and short tooth on basal half of external margin of mandible (Fig. 113). Maxillary cardo subquadrate and base somewhat rounded (Fig. 130); stipes triangular; lacinia and galea of same length; lacinia with dense short setae and spur-like structure on apex; galea narrower than lacinia, dense short setae on apex and some long setae on external margin near apex; maxillary palpus with palpomere 1 the shortest, 2 and 3 subequal in length and 4 the longest, sometimes longer than 2 and 3 combined (Fig. 131). Mentum subquadrate to 2 times wider than long (Figs 133–135); each side with set of long setae; anterior margin somewhat curved; prementum short and hidden under mentum; ligula emarginate at middle and long setae on anterior margin; labial palpus with palpomere 1 and 3 subequal in length and longer than 2 (Fig. 133). Gular plate narrow, gular sutures narrowly separated (Fig. 8). Pronotum wider than long, PW/PL = 1.25 (P. zischkai; Fig. 69) to 1.56 (P. aper; Fig. 98); anterior angles generally weakly projected; surface sometimes irregular; longitudinal median sulcus conspicuous or inconspicuous (Fig. 5); longitudinal sulcus of posterior angles present in some species and reach the middle of pronotum (Fig. 5); abrupt constriction at basal one-third or one-quarter and, anterior to it, in some species, a short tooth and a slight emargination of each side (Fig. 5); apical half, twothrids or three-quarters curved, sinuous or with parallel sides. Prosternum with anterior margin anteriorly projected in middle (Fig. 8); some species with conspicuous ovate set of fine punctures in median region near apex (Fig. 8); protrochantin visible externally, developed; postcoxal hypomeral projection acute and long, almost reaching apex of procoxa in ventral view but not reaching prosternum or closing procoxal cavities. Scutellum visible, developed and somewhat triangular (Fig. 5). Elytra truncate; nearly as long as wide, EL/BW = 0.90 (P. spinosus; Fig. 19) to 1.17 (P. paradoxus; Fig. 48); each Revision of Piestus with remarks on related genera Invertebrate Systematics Figs 5–11. Piestus sulcatus. 5, Habitus, male, dorsal view; 6, habitus, male, lateral view; 7, head and pronotum, male, dorsal view; 8, head and pronotum, male, ventral view; 9, meso and metaventrite, male, ventral view; 10, metaventrite and base of the abdomen, male, ventral view; 11, apex of the abdomen, male, ventral view. Scale bar (5, 6) = 1 mm; (7–11) = 0.50 mm. 501 502 Invertebrate Systematics E. Caron et al. Figs 12–23. Habitus, dorsal view. 12, Piestus convexus, sp. nov.; 13, P. similis, sp. nov.; 14, P. lacordairei; 15, P. heterocephalus; 16, P. puncticollis; 17, P. capricornis; 18, P. planatus; 19, P. spinosus; 20, P. longipennis; 21, P. zischkai; 22, P. bicornis; 23, P. bicornis. Scale bar = 1 mm. elytron with five complete longitudinal striae and in some species stria 6 on basal half and/or stria 7 complete (Fig. 5); apex with short translucent region; epipleural carina present or not (Fig. 6). Hind wings fully developed, ~3 times longer than wide; anal area with long setae and short setae on posterior margin. Mesoventrite with apical process reaching middle of mesocoxae and carinate (Fig. 9). Metaventrite developed (Fig. 10); each lateral region with two or three long posteriorly directed setae; paracoxal suture Revision of Piestus with remarks on related genera Invertebrate Systematics 503 Figs 24–35. Habitus, dorsal view. 24, Piestus fronticornis, sp. rev.; 25, P. validus; 26, P. rossii, sp. nov.; 27, P. pennicornis; 28, P. pygmaeus; 29, P. extimus; 30, P. buquetii; 31, P. filicornis; 32, P. abdominalis, sp. nov.; 33, P. niger; 34, P. minutus; 35, P. penicillatus. Scale bar = 1 mm. inconspicuous; posterior margin of metacoxa slightly emarginate. Anterior legs more robust than others (Fig. 138); procoxae transverse and contiguous; protrochanter small and subtriangular; profemur robust and slightly flat anteroposteriorly, apical region of anterior margin of ventral face with groove and short or long set of contiguous hard setae 504 Invertebrate Systematics (Figs 138, 139); protibia with apex wider than base, longitudinal row of robust spines on apical half of external margin and at apex, internal margin slightly emarginate and four longitudinal rows of robust setae, robust spine on the internal angle of apex, reaching tarsomere 4; tarsomeres 1–4 subequal in length, long setae at apex, tarsomere 5 longer than 1–4 combined, slightly curved; pretarsus composed of pair of claws, unguitractor plate, empodium and pair of empodial setae (Fig. 142). Mesotrochantin present, visible (Fig. 140); mesocoxae separated, each globose, and metacoxae contiguous (Fig. 10), each transverse and subtrapezoidal; meso- and metatrochanter subtriangular (Fig. 141); meso- and metafemur slightly flat anteroposteriorly and somewhat curved dorsally; meso- and metatibiae subcylindrical and thin, mesotibia notably more curved than metatibia; each apical half with longitudinal rows of robust setae, external rows more robust, apex with two robust spines, internal the longest; tarsus and pretarsus similar to anterior legs. Abdominal segments 3–6 with parallel or slightly curved sides; tergite 1 and 2 weakly sclerotised and hidden under elytra (Fig. 5); tergites 3–7 with pair of paratergites; sternite 1 short and inconspicuous and sternite 2 sclerotised and fused with sternite 3 (Figs 143, 144); sternite 3 with transverse carina developed or not (Figs 10, 143, 144); sternite 6 with very small emargination on each posterolateral area (Fig. 11), external openings of abdominal defensive gland complex (for details see Caron et al. 2008); tergite 8 with apical margin curved (Fig. 134); sternite 8 with apical margin truncate to slightly curved (Fig. 146); tergite 10 separating tergite 9 in two equal parts (Fig. 149) contiguous or weakly separate at base of tergite 10; tergite 9 with ventral struts, short or long (Figs 149, 150); tergite 10 with one or two pairs of long setae near apex; sternite 9 weakly pigmented (Fig. 152), composed of a single median sclerite, symmetrical, 3 times longer than wide, basal margin acute, apical margin rounded, with one or two pairs of short setae on apex. Aedeagus with median lobe curved in lateral view (Fig. 214), sometimes bulbous base in ventral view; lateral lobes developed, in some species exceeding apex of median lobe; internal sclerites of variety format. Female. In general similar to male, except: apex of sternite 8 slightly acute (Fig. 147); tergite 9 without ventral struts; sternite 9 as ovipositor consisting of pair of very weakly pigmented hemisternites and pair of more apical coxites (Fig. 153), short setae and some microsetae on apex, and without stylus. Spermathecal duct short or long, and in some species sclerotised at base (Fig. 262); spermatheca in various formats. Sexual dimorphism Some species are sexually dimorphic in shape of the frontal processes of the head, male converging to the apex and female not (Figs 68, 69); in length of antenna, male longer than female (Figs 106, 107); at antennal scape, long setae and enlarged area only in male; on internal border of mandible, male with teeth and female with only one tooth on the right mandible (Figs 123, 124); at middle region near the apex of prosternum, ovate set of fine punctures only in male (Fig. 8); and at elytra, length of the apical process longer in females than males (Figs 48, 49). E. Caron et al. Distribution Nearctic region: USA (only P. extimus), and Neotropical region: Mexico, Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador (including Galápagos Islands), Peru, Bolivia, Brazil, Paraguay, Argentina, Uruguay and the West Indies (Cuba, Dominican Republic, Puerto Rico, Guadeloupe, Dominica, Martinique, Saint Vincent and the Grenadines, Grenada, and Trinidad and Tobago). Piestus is nearly restricted to the Neotropical region in a strict sense, excluding Chile and southern Argentina. Only one species is found in the Nearctic region, in south-western USA (Arizona). Biological notes The species of Piestus are commonly found associated with organic material in a decomposition process, where they are saprophages and/or mycophages as adults and larvae (based on our observations of gut contents). Piestus has been found on or under bark of decaying logs, in leaf litter, in fungi (‘fungus covered log’), in decaying cacti and sotol (Dasylirion spp.), in fruit and flower falls on the ground, and also on dung of mammals. One species may be associated with termites (P. termitis) and another with ants (P. formicinus), but these associations need to be confirmed. Some species of Piestus have been found occurring together in the same habitat. Piestus species have also been collected by flight intercept and Malaise traps, Berlese and Winkler extraction of leaf litter, and attracted to UV light. Key to species of Piestus 1. Pronotum with lateral tooth before abrupt constriction (Fig. 7); elytra with epipleural carina (Fig. 6)....................................................................2 Pronotum without lateral tooth before abrupt constriction; elytra without epipleural carina ...............................................................................18 2. (1) Head with microstriae uniformly distributed (Fig. 7); pronotum with some fine to moderately sized punctures .............. P. sulcatus (Fig. 5) Head without microstriae uniformly distributed; pronotum with large punctures, contiguous or not..............................................................3 3. (2) Pronotum with large punctures not contiguous, and with conspicuous median longitudinal sulcus (Fig. 88) .................................................4 Pronotum with large punctures contiguous, and with inconspicuous median longitudinal sulcus (Fig. 91) ..............................................................7 4. (3) Head with fine to moderately sized punctures (Fig. 87); elytra with stria 6 developed at the basal half................................ P. gounellei (Fig. 39) Head with large punctures (Fig. 88); elytra with stria 6 absent...................5 5. (4) Head with large punctures not contiguous at the median region of front and vertex (Fig. 88)......................................... P. mexicanus (Fig. 40) Head with large punctures contiguous at the median region of front and vertex (Fig. 89) .............................................................................................6 6. (5) Head with median longitudinal sulcus and a conspicuous fovea at the base of the vertex (Fig. 90) ................ P. foveolatus sp. nov. (Fig. 42) Head without median longitudinal sulcus and fovea at the base of the vertex (Fig. 89) ........................................ P. ecuadorensis sp. nov. (Fig. 41) 7. (3) Pronotum with anterior angles acute and projected (Fig. 98) .............. ..................................................................................P. aper (Fig. 51) Pronotum with anterior angles weakly or not projected ..............................8 8. (7) Mandibles with only one acute tooth on each internal border ...........9 Mandibles with two acute teeth on each internal border, basal anteriorly projected (Fig. 127)..........................................................................12 Revision of Piestus with remarks on related genera 9. (8) Elytra with stria 6 developed at the basal half ..................................... ....................................................................... P. sulcipennis (Fig. 43) Elytra without stria 6..................................................................................10 10. (9) Elytra with apical margin emarginate when closed (Fig. 50); metaventrite with moderately sized punctures on median region ................................................ P. boliviensis sp. nov. (Fig. 50) Elytra with apical margin not emarginate when closed; metaventrite with moderately sized punctures only on lateral region ..........................11 11. (10) Aedeagus with slightly curved apex of median lobe in lateral view, and lateral lobes not narrowing to apex (Fig. 228); spermatheca L-shaped and spermathecal duct long (Fig. 279)................................. ............................................................................ P. costatus (Fig. 45) Aedeagus with curved apex of median lobe in lateral view, and lateral lobes narrowing to apex (Fig. 232); spermatheca spiral shaped and spermathecal duct short (Fig. 281)................ P. nevermanni (Fig. 47) 12. (8) Metaventrite with moderately sized punctures on median region ..13 Metaventrite with moderately sized punctures only on lateral region .......15 13. (12) Elytra with stria 6 developed at the basal half, interstriae 2 and 4 joined at the apex and weakly posteriorly extended in male and strongly in female (Figs 48, 49)................P. paradoxus (Figs 48, 49) Elytra without stria 6 at the basal half, interestriae 2 and 4 not joined at the apex and posteriorly extended ...................................................14 14. (13) Head with V-shaped frontal sulcus incomplete (Fig. 92)................. .........................................................P. imperfectus sp. nov. (Fig. 44) Head with V-shaped frontal sulcus complete (Fig. 94) ................................. ......................................................................P. chiriquensis (Fig. 46) 15. (12) Striae of elytra with large punctures and microgranulate (Fig. 55).. .........................................................................................................16 Striae of elytra with only large punctures (Fig. 52).... P. angularis (Fig. 52) 16. (15) Elytra with interstriae 1 and 4 joined at the apex and weakly posteriorly extended (Fig. 53); tergite 3 hidden by elytra ................... ......................................................... P. acuminatus sp. nov. (Fig. 53) Elytra without interstriae 1 and 4 joined at the apex and weakly posteriorly extended; tergite 3 not hidden by elytra ..........................................17 17. (16) Antennomeres 4–10 oblong shape (male and female) (Fig. 54); spermathecal duct short, about the same length as spermatheca (Fig. 286) .............................................................P. rugosus (Fig. 54) Antennomeres 4–10 bead-like (female) (Fig. 55); spermathecal duct long, ~2 times the length of spermatheca (Fig. 287) .................................... ............................................................... P. elegans sp. nov. (Fig. 55) 18. (1) Pronotum with a slight emargination on each side, anterior to abrupt constriction (Fig. 18)........................................................................19 Pronotum without a slight emargination on each side...............................22 19. (18) Head with pair of frontal processes ..............................................20 Head without frontal processes (Fig. 63).................P. puncticollis (Fig. 16) 20. (19) Head with frontal processes shorter than the antennal scape (Fig. 64); V-shaped frontal sulcus incomplete; dorsal teeth of mandibles with the same length as ventral teeth (Fig. 119) ................ .......................................................................P. capricornis (Fig. 17) Head with frontal processes longer than antennal scape; V-shaped frontal sulcus complete; dorsal teeth of mandibles longer than ventral ......21 21. (20) Head, pronotum and elytra reddish brown and abdomen reddish dark brown to black, sometimes head darker than pronotum and elytra (Fig. 19) ............................................................. P. spinosus (Fig. 19) Body entirely black (Fig. 18)....................................... P. planatus (Fig. 18) 22. (18) Head with frontal processes..........................................................35 Head without pair of frontal processes ......................................................23 23. (22) Antennae short, reaching or exceeding a little the apex of elytra (male and female); pronotum with abrupt constriction on basal third ..................................................................................................24 Antennae long, almost or reaching the apex of abdomen (male longer than female); pronotum with abrupt constriction on basal quarter ..........26 24. (23) Head with V-shaped frontal sulcus incomplete (Fig. 86); elytra with stria 7 complete......................... P. formicinus sp. nov. (Fig. 38) Invertebrate Systematics 505 Head with V-shaped frontal sulcus complete (Fig. 84); elytra with stria 7 only at the apical half....................................................................25 25. (24) Aedeagus with lateral lobes exceeding the apex of median lobe (Fig. 206); spermathecal duct about the same length as spermatheca (Fig. 269) ............................................... P. termitis sp. nov. (Fig. 36) Aedeagus with lateral lobes reaching the apex of median lobe, not exceeding it (Fig. 208); spermathecal duct long, ~3 times the length of spermatheca (Fig. 270) ..................... P. surrufus sp. nov. (Fig. 37) 26. (23) Head with V-shaped sulcus complete (Fig. 80) ...........................27 Head with V-shaped sulcus incomplete (Fig. 74)......................................34 27. (26) Head with transverse carina at base of vertex (Fig. 82) ...............28 Head without transverse carina at base of vertex (Fig. 76) .......................32 28. (27) Pronotum with undulate microstriae uniformly distributed..........29 Pronotum with undulate microstriae only on lateral regions, sometimes also near the median longitudinal sulcus.................................................31 29. (28) Head with transverse carina at base of vertex strongly interrupted at the middle (Fig. 83) ...................................P. penicillatus (Fig. 35) Head with transverse carina at base of vertex not interrupted or weakly interrupted at the middle ..................................................................30 30. (29) Specimens of moderate size, ~8.0 mm long; aedeagus with curved apex of median lobe in lateral view (Fig. 196).. P. filicornis (Fig. 31) Specimens of small size, ~4.5 mm long; aedeagus without curved apex of median lobe in lateral view (Fig. 202).................P. minutus (Fig. 34) 31. (29) Pronotum with conspicuous fine punctures and some moderately sized punctures on the median longitudinal sulcus (Fig. 80); aedeagus with curved apex of median lobe in lateral view (Fig. 198)................ ........................................................P. abdominalis sp. nov. (Fig. 32) Pronotum only with conspicuous fine punctures (Fig. 81); aedeagus without curved apex of median lobe in lateral view (Fig. 200)........... .................................................................................P. niger (Fig. 33) 32. (27) Pronotum with parallel sides (Fig. 76); aedeagus with lateral lobes exceeding the apex of median lobe by one-quarter of its length in lateral view (Fig. 1).......................................... P. pygmaeus (Fig. 28) Pronotum with subparallel sides, very slightly curved on apical half (Fig. 78); aedeagus with lateral lobes exceeding a little the apex of median lobe in lateral view (Figs 192, 194).........................................................33 33. (32) Pronotum with undulate microstriae and inconspicuous fine punctures evenly distributed (Fig. 77); aedeagus with slightly narrowed apex of lateral lobes in lateral view (Fig. 192) .................... ............................................................................. P. extimus (Fig. 29) Pronotum with undulate microstriae only on lateral areas and conspicuous fine punctures evenly distributed (Fig. 78); aedeagus with broad apex of lateral lobes in lateral view (Fig. 194)............ P. buquetii (Fig. 30) 34. (26) Pronotum with curved sides (Fig. 73); aedeagus with curved apex of the median lobe in lateral view (Fig. 186).............................. ...................................................................P. rossii sp. nov. (Fig. 26) Pronotum with parallel sides (Figs 74, 75); aedeagus without curved apex of the median lobe in lateral view (Fig. 188) ...................................... ...................................................................... P. pennicornis (Fig. 27) 35. (22) Head with basal distance between the frontal processes the same or narrower than the basal width of each one (Fig. 67)...................36 Head with basal distance between the frontal processes wider than basal width of each one (Fig. 61)..............................................................40 36. (35) Body entirely dark brown to black (Fig. 20); aedeagus without extended region of median lobe for supporting the base of lateral lobes (Fig. 172) ................................................................................37 Body brown to black with apical half or all the segment 7 and the segments 8–10 of abdomen yellowish to yellowish brown (Fig. 22), or head and pronotum dark brown to black with pronotum and elytra reddish brown, also with apical half of the segment 7 and the segments 8–10 of abdomen yellowish brown (Fig. 23); aedeagus with extended region of median lobe for supporting the base of lateral lobes (Fig. 179) .....................................................................38 506 Invertebrate Systematics 37. (36) Head with frontal processes of male strongly converging towards their apices (Fig. 68) while in female the internal margins are parallel (Fig. 69); tergite 10 with two pairs of short setae on the apex, equal in length; aedeagus with lateral lobes exceeding the apex of median lobe in lateral view (Fig. 176)..................................... P. zischkai (Fig. 21) Head with frontal processes of male and female equally and slightly diverging towards their apices (Fig. 67); tergite 10 with two pairs of long setae on the apex, basal the longest; aedeagus with lateral lobes reaching the apex of median lobe in lateral view (Fig. 172) ............... .......................................................................P. longipennis (Fig. 20) 38. (36) Body brown to black with, in general, all of segment 7 and segments 8–10 of abdomen yellowish (Fig. 24); aedeagus in lateral view very thin and with slightly developed internal sclerite (Fig. 182) ............... ...................................................................... P. fronticornis (Fig. 24) Body brown to black with apical half of segment 7 and segments 8–10 of abdomen yellowish to yellowish brown (Fig. 22), or head and pronotum dark brown to black with pronotum and elytra reddish brown, also with apical half of the segment 7 and the segments 8–10 yellowish brown (Fig. 23); aedeagus different from above.............39 39. (38) Body brown to black with apical half of segment 7 and segments 8–10 of abdomen yellowish (Fig. 25); antennal scape with many long setae on the apical half of the dorsal face and antennomeres 3–6 with many long setae on dorsal face (Figs 25, 72).............................. ..............................................................................P. validus (Fig. 25) Body brown to black with apical half of segment 7 and segments 8–10 of abdomen yellowish brown (Fig. 22), or head and pronotum dark brown to black with pronotum and elytra reddish brown, also with apical half of segment 7 and segments 8–10 of abdomen yellowish brown (Fig. 23); antennal scape with some long setae at the middle of the dorsal face and antennomeres 3–6 with some long setae on dorsal face (Figs 22, 23, 70) .........................P. bicornis (Figs 22, 23) 40. (35) Head with short frontal processes shorter than the antennal scape (Fig. 60) ...........................................................................................41 Head with long frontal processes, longer than the antennal scape (Fig. 61) . .........................................................................................................42 41. (40) Antennae long, almost reaching the apex of abdomen (Fig. 12), and scape with long setae on basal half of exposed dorsal face (Fig. 59).. .............................................................P. convexus sp. nov. (Fig. 12) Antennae short, reaching the apex of elytra (Fig. 13), and scape with only two long setae at the middle of the dorsal face (Fig. 60) .................... ................................................................. P. similis sp. nov. (Fig. 13) 42. (40) Body entirely brown to black, sometimes with the abdomen slightly lighter than the body (Fig. 14); head with ventral face of frontal processes without or with only one tooth near the base ................................................................ P. lacordairei (Fig. 14) Body brown to black with apical half of segment 7 and segments 8–10 of abdomen yellowish (Fig. 15); head with ventral face of frontal processes coarsely serrate ........................ P. heterocephalus (Fig. 15) The species of Piestus are arranged below based on the current phylogenetic study (see ‘Results’, ‘Cladistic analysis’) and to facilitate the discussion. Piestus convexus, sp. nov. (Figs 12, 59, 156, 157) Type material Holotype deposited in SEMC, male: (1) ‘Ecuador: Napo, Cosanga/4.2 km S on Baeza-Tena/Road then 1.5 km W on pipeline/access road, 2150 m/ 0
370 1900 S, 77
500 100 W/5–7 Nov 1999, Z.H. Falin/ECU1F99 123 flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0352227/ KUNHM-ENT’ [white label, printed in black]; (3) ‘<’ [white label, printed in black]. E. Caron et al. Description BL: 9.5 mm, BW: 2.7 mm. Body convex; entirely black (Fig. 12); setae on body black, except antennomeres 10–11, and tarsi golden. Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures (Fig. 59); elytral striae unpunctured; abdominal tergites on anterolateral regions, with moderately sized punctures. Male. Head with pair of broad and short frontal processes, shorter than the scape, and basal distance between processes ~2 times basal width of each one (Fig. 59); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and prominent; eyes with line of five moderately sized punctures with long setae on basal half of dorsal margin, equal in length. Antennae almost reaching apex of abdomen (Fig. 12); scape with long setae on basal half of exposed dorsal face; scape shorter than antennomeres 2 and 3 combined; antennomeres 4–11 oblong shape. Mandibles strongly projected, bifurcate apices; each with dorsal tooth shorter than the ventral; internal border asymmetrical, with one acute tooth at middle and, anterior to it, in right mandible, with two areas that resemble a tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Pronotum wider than long (PW/PL = 1.43) (Fig. 59); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical threequarters with slightly curved sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.04); stria 6 at basal half and stria 7 at apical quarter inconspicuous; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 156); lateral lobes exceed a little apex of median lobe in lateral view; internal sclerites as in Fig. 157. Female. Unknown. Distribution Ecuador (Napo). Biological notes The unique specimen (holotype) was collected in a flight intercept trap. Remarks Piestus convexus differs from P. similis by the broad and short frontal processes (Fig. 59), long antennae almost reaching the apex of the abdomen (Fig. 12), antennal scape with long setae on the basal half of exposed dorsal face, and mandibles with asymmetrical internal border and without external marginal carina on basal half. Piestus similis has acute and short frontal processes (Fig. 60), antennae short, reaching the apex of elytra (Fig. 13), antennal scape with two long setae at the middle of the dorsal face, mandibles with symmetrical internal border and external margin slightly carinate on basal half. Revision of Piestus with remarks on related genera Invertebrate Systematics 507 Figs 36–46. Habitus, dorsal view. 36, Piestus termitis, sp. nov.; 37, P. surrufus, sp. nov.; 38, P. formicinus, sp. nov.; 39, P. gounellei; 40, P. mexicanus; 41, P. ecuadorensis, sp. nov.; 42, P. foveolatus, sp. nov.; 43, P. sulcipennis; 44, P. imperfectus, sp. nov.; 45, P. costatus; 46, P. chiriquensis. Scale bar = 1 mm. 508 Invertebrate Systematics E. Caron et al. Figs 47–58. Habitus, dorsal view. 47, Piestus nevermanni; 48, P. paradoxus, male; 49, P. paradoxus, female; 50, P. boliviensis, sp. nov.; 51, P. aper; 52, P. angularis; 53, P. acuminatus, sp. nov.; 54, P. rugosus; 55, P. elegans, sp. nov.; 56, Hypotelus laevis, comb. nov.; 57, H. andinus, comb. nov.; 58, Eleusis interrupta, comb. rest. Scale bar = 1 mm. Revision of Piestus with remarks on related genera Invertebrate Systematics 509 Figs 59–75. Head and pronotum, dorsal view. 59, Piestus convexus, sp. nov.; 60, P. similis, sp. nov.; 61, P. lacordairei; 62, P. heterocephalus; 63, P. puncticollis; 64, P. capricornis; 65, P. planatus; 66, P. spinosus; 67, P. longipennis; 68, P. zischkai, male; 69, P. zischkai, female; 70, P. bicornis; 71, P. fronticornis, sp. rev.; 72, P. validus; 73, P. rossii, sp. nov.; 74, P. pennicornis, male; 75, P. pennicornis, female. Scale bar = 0.50 mm. 510 Invertebrate Systematics E. Caron et al. Figs 76–91. Head and pronotum, dorsal view. 76, Piestus pygmaeus; 77, P. extimus; 78, P. buquetii; 79, P. filicornis; 80, P. abdominalis, sp. nov.; 81, P. niger; 82, P. minutus; 83, P. penicillatus; 84, P. termitis, sp. nov.; 85, P. surrufus, sp. nov.; 86, P. formicinus, sp. nov.; 87, P. gounellei; 88, P. mexicanus; 89, P. ecuadorensis, sp. nov.; 90, P. foveolatus, sp. nov.; 91, P. sulcipennis. Scale bar = 0.50 mm. Revision of Piestus with remarks on related genera Invertebrate Systematics 511 Etymology Remarks The specific name refers to the body format. Piestus similis differs from P. convexus by the characters cited above (see remarks under P. convexus). Piestus similis, sp. nov. (Figs 13, 60, 158, 159) Type material Etymology The name of this species refers to the similar species, P. convexus. Holotype deposited in SEMC, male: (1) ‘Ecuador: Napo/Sierra Azul, 2300m/0
400 000 S, 77
550 000 W/21 May–6 Jun 1996, P. Hibbs/ECU2H96 004D, ex: Malaise trap’ [white label, printed in black]; (2) ‘bar code/ SM0170320/KUNHM-ENT’ [white label, printed in black]; (3) ‘<’ [white label, printed in black]. Description BL: 9.7 mm, BW: 2.8 mm. Body convex; entirely dark brown (Fig. 13); setae on body black, except antennomeres 10–11 and tarsi golden. Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures (Fig. 60); elytral striae unpunctured; metaventrite on lateral regions, and abdominal tergites on anterolateral regions with moderately sized punctures. Male. Head with pair of acute and short frontal processes, shorter than scape, and basal distance between processes ~6 times basal width of each one (Fig. 60); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and prominent; eyes with one line of five moderately sized punctures with long setae on basal half of dorsal margin, equal in length. Antennae reaching apex of elytra (Fig. 13); scape with two long setae at middle of dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Mandibles strongly projected; bifurcate apices; each with dorsal tooth shorter than ventral; internal border symmetrical, right mandible with one acute tooth at middle; external margin slightly carinate at basal half. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Pronotum wider than long (PW/PL = 1.39) (Fig. 60); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical three-quarters with curved sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the shortest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 158); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 159. Female. Unknown. Distribution Ecuador (Napo). Biological notes The unique specimen (holotype) was collected in a Malaise trap. Piestus lacordairei Laporte (Figs 14, 61, 116, 160, 161, 251) Piestus lacordairei Laporte, 1835: 129 (original description, type locality: ‘Cayenne’); Erichson, 1840: 832 (characters, distribution); Fauvel, 1864: 21 (characters, distribution); Blackwelder, 1944: 100 (distribution, error: Laporte, 1834: 129); Herman, 2001b: 1791 (distribution); Newton et al., 2005: 37 (distribution); Caron et al., 2008: 5 (characters). Piestus (Zirophorus) lacordairei: Bernhauer & Schubert, 1910: 6 (distribution, error: Laporte, 1834: 129); Scheerpeltz, 1951: 7 (characters, distribution, error: Laporte, 1834: 129); Scheerpeltz, 1952: 283 (characters, distribution, error: Laporte, 1834: 129). Zirophorus furcatus Sharp, 1887: 712 (original description, type locality: ‘Panama, Volcan de Chiriqui 2500 to 4000 feet’). New synonym. Piestus (Zirophorus) furcatus: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1951: 7 (characters, distribution); Scheerpeltz, 1952: 283 (characters, distribution). Piestus furcatus Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1791 (distribution). Type material Piestus lacordairei Laporte, 1835. Neotype (here designated) deposited in ZMHB, sex undetermined, with labels: (1) ‘6804’ [white label, printed in black]; (2) ‘Lacordairei/Lap./Cayenne Dej.’ [green label, handwritten] (ZMHB). Note: In the original description Laporte (1835) did not specify how many specimens were observed. The type material was not found and probably it is lost. Zirophorus furcatus Sharp, 1887. Syntype deposited in FMNH, sex undetermined, with labels: (1) ‘Piestus furcatus./D.S./V. de Chiriqui. 2500-4000 ft./Champion.’ [white label, handwritten, together with the specimen]; (2) ‘V. de Chiriqui,/25-4000 ft./Champion.’ [white label, printed in black]; (3) ‘B.C.A. Col. I. 2./Zirophorus/furcatus,/Sharp.’ [white label, printed in black]; (4) ‘Sharp Coll. 1905,-313.’ [white label, printed in black]; (5) ‘Chicago Nat. Hist. Mus./(ex. D. Sharp Colln./by exchange with/ Brit. Mus. Nat. Hist.)’ [white label, printed in black]. Note: In the original description Sharp (1887) specified eight specimens observed. There are seven syntypes (not examined) deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). Additional material See Appendix 2. Redescription BL: 5.1–8.0 mm, BW: 1.4–2.1 mm. Body flattened dorsoventrally; entirely brown to black, sometimes with abdomen lighter than body (Fig. 14). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures (Fig. 61); striae of elytra unpunctured or with inconspicuous fine punctures; 512 Invertebrate Systematics metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes, longer than the scape, ventral face of each frontal process with only one tooth near base, and basal distance between processes ~2 times basal width of each one (Fig. 61); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae almost reaching apex of elytra (Fig. 14); scape with two long setae near apex of dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with four setae medially, equal in length; each lateral third, antero-internal seta with same length as postero-external seta. Mandibles strongly projected (Fig. 116); bifurcate apices; each with dorsal tooth shorter than ventral; internal border asymmetrical, internal border with one acute tooth at middle and, anterior to it, in right mandible with an area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/ PL = 1.27) (Fig. 61); anterior angles not projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical threequarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.05); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 160); lateral lobes exceeding apex of median lobe by one-quarter of its length in lateral view and slightly curved apex on ventral margin; internal sclerites as in Fig. 161. Female. Similar to male. Ovipositor as in P. sulcatus; spermathecal duct ~2 times length of spermatheca, with sclerotised basal region (Fig. 251); spermatheca with basal half suddenly narrower than apical half (Fig. 251). Distribution In the current study, the species is listed from Nicaragua (Rio San Juan), Costa Rica (Guanacaste, Alajuela, Heredia, San José, Cartago, Limón and Puntarenas), Panamá (Bocas del Toro, Chiriquí, Veraguas and Colón), Colombia (Cundinamarca and Huila), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Esmeraldas, Pichincha, Napo and Cotopaxi), Peru (Ucayali and Cusco) and Brazil (Pará and Rondônia). Herman (2001b) listed the species also from Venezuela, Guyana and Suriname, and Newton et al. (2005) from Colombia (Valle del Cauca). E. Caron et al. Remarks Piestus lacordairei is similar to P. heterocephalus but differs mainly by the body being entirely brown to black (Fig. 14), ventral face of frontal processes with only one tooth near the base, labrum with four setae medially and lateral lobes of aedeagus with slightly curved apex on ventral margin (Fig. 160). Piestus heterocephalus has body black with apical half of abdominal segment 7 and all of segments 8–10 yellowish (Fig. 15), ventral face of frontal processes coarsely serrate, six setae medially on labrum and lateral lobes of aedeagus with truncate apex on ventral margin (Fig. 162). Piestus heterocephalus Fauvel, 1902 (Figs 15, 62, 162, 163, 252) Piestus heterocephalus Fauvel, 1902: 22 (original description, type locality: ‘Venezuela: Merida’); Blackwelder, 1944: 100 (distribution, error: Fauvel, 1865: 22); Herman, 2001b: 1791 (distribution); Caron et al., 2008: 3 (redescription, distribution, defensive abdominal gland complex). Piestus (Piestus) heterocephalus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 285 (characters, distribution). Type material Holotype deposited in IRSNB, sex undetermined, with labels: (1) ‘Merida/Venezuela’ [white label, handwritten]; (2) ‘heterocephalus/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et. det. A. Fauvel’ [white label, first and third lines printed in black, and second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in red]. Note: For discussion about the type of P. heterocephalus see Caron et al. (2008). Diagnosis Body flattened dorsoventrally; black with apical half of abdominal segment 7 and all segments 8–10 yellowish (Fig. 15); pair of acute and long frontal processes on head, ventral face of frontal processes coarsely serrate and basal distance between the processes ~2 times the basal width of each one (Fig. 62); V-shaped frontal sulcus incomplete; antennae short, reaching the apex of elytra and scape with two long setae near the apex of the dorsal face; labrum with six setae medially; mandibles with bifurcate apex, internal border asymmetrical; pronotum with inconspicuous fine punctures and parallel lateral sides; lateral lobes of aedeagus with truncate apex on ventral margin (Fig. 162). Distribution Colombia (Valle del Cauca), Venezuela (Mérida) and Ecuador (Pichincha, Cotopaxi and El Oro) (Caron et al. 2008). Biological notes Unknown. Biological notes Remarks This species has been found under bark, associated with fermenting decaying logs. Some specimens were collected at flight intercept traps or by Berlese extraction (forest litter). This species was recently redescribed by Caron et al. (2008) and its abdominal defensive gland complex described and reported for the first time in Piestinae. Revision of Piestus with remarks on related genera Piestus heterocephalus is similar to P. lacordairei, differing from that species by the characters cited above (see remarks under P. lacordairei). Piestus puncticollis Fauvel, 1902 (Figs 16, 63, 117, 118, 164, 165) Piestus puncticollis Fauvel, 1902: 24 (original description, type locality: ‘Amazones: Ega’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1793 (distribution). Piestus (Piestus) puncticollis: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 286 (characters, distribution). Type material Piestus puncticollis Fauvel, 1902. Lectotype (here designated) deposited in IRSNB, male, with labels: (1) ‘Ega’ [white label, handwritten]; (2) ‘puncticollis’ [white label, handwritten]; (3) ‘Ex-Typis’ [white label, printed in red]; (4) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black, and second line handwritten]. Three paralectotypes deposited in IRSNB, males, with labels: (1) ‘Ega’ [white label, handwritten]; (2) ‘Ex-Typis’ [white label, printed in red]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/puncticollis Fvl./Coll. et det. A. Fauvel’ [white label, first and fourth lines printed in black, second and third lines handwritten]. Note: In the original description, Fauvel (1902) did not specify how many specimens he observed. We received from IRSNB four males with type labels. Redescription BL: 5.5–5.6 mm, BW: 1.4–1.5 mm. Body strongly flattened dorsoventrally; entirely light brown to reddish dark brown, with head and abdomen darker (Fig. 16). Dorsal integument of head and pronotum with inconspicuous fine punctures and some moderately sized punctures distributed on pronotal disc, undulate microstriae only on lateral regions (Fig. 63); striae of elytra with moderately sized punctures; metaventrite, on lateral regions, and abdominal tergites on anterolateral regions with moderately sized punctures. Male. Head with front slightly deflected (Fig. 63); V-shaped frontal sulcus complete, sometimes curved arms slightly joined medially; anterior angles curved, prominent and with median fovea at base; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 16); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Mandibles with bifurcate apex (Fig. 117); each with dorsal tooth shorter than ventral, projected dorsally and forming perpendicular angle (Fig. 118); internal border of mandibles symmetrical, each with strong emargination near apex and two acute teeth, basal smaller and projected anteriorly and apical more acute and projected almost posteriorly. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Pronotum wider than long (PW/PL = 1.50) (Fig. 63); anterior angles projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter and, anterior to it, in each side with slight emargination; apical three-quarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.05); stria 6 Invertebrate Systematics 513 at basal half and stria 7 absent or slightly developed only on apex; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 164); lateral lobes exceeding apex of median lobe by one-quarter of its length in lateral view; internal sclerites as in Fig. 165. Female. Unknown. Distribution Brazil (Amazonas). Biological notes Unknown. Remarks Piestus puncticollis is similar to P. capricornis, from which it is easily distinguished by the head without a pair of frontal processes (Fig. 63), antennal scape with a slightly enlarged area and long setae on the basal half of the dorsal face, and mandibles with bifurcate apex, each with dorsal tooth projected dorsally and forming a perpendicular angle with the ventral tooth, internal border symmetrical (Figs 117, 118). Piestus capricornis has a pair of acute and short frontal processes (Fig. 64), antennal scape with two long setae at the middle of the dorsal face, mandibles with internal border asymmetrical and right mandible with trifurcate apex (Fig. 119). Fauvel (1902), when describing P. puncticollis, cited a slight sexual dimorphism of the antennae. However, all type specimens examined from IRSNB are males and probably Fauvel considered a small male as female. Polymorphism in males of Piestus, mainly related to sexual dimorphism, is common and it was observed in other species. The females of P. puncticollis are still unknown, as only the type material has been studied. Piestus capricornis Laporte, 1835 (Figs 17, 64, 109, 119, 166, 167, 253) Piestus capricornis Laporte, 1835: 129 (original description, type locality: ‘Cayenne’); Erichson, 1840: 833 (characters, distribution); Fauvel, 1864: 22 (characters, distribution); Fleutiaux & Sallé, 1889: 382 (distribution); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1789 (distribution). Piestus (Zirophorus) capricornis: Bernhauer & Schubert, 1910: 6 (distribution); Blackwelder, 1943: 50 (characters, distribution, notes); Scheerpeltz, 1951: 8 (characters, distribution); Scheerpeltz, 1952: 284 (characters, distribution). Piestus capricornis variety muticus Fauvel, 1902: 22 (original description, type locality: ‘Para: Benevides, 2; Marco da Legua, 30 ); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1790 (distribution). Piestus (Zirophorus) capricornis variety muticus: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1951: 8 (characters, distribution); Scheerpeltz, 1952: 284 (characters, distribution). 514 Invertebrate Systematics Piestus frontalis Sharp, 1876: 405 (original description, type locality: ‘Ega’). Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1791 (distribution). New synonym. Piestus (Zirophorus) frontalis: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1951: 7 (characters, distribution); Scheerpeltz, 1952: 284 (characters, distribution). Type material Piestus capricornis Laporte, 1835. Syntype deposited in BMNH, sex undetermined [damaged apex of the left antenna], with labels: (1) ‘Piestus capricornis Lap. Type/ex coll. Chevrolat./Guadeloupe. L‘Herminier’ [white label, together with the specimen, handwritten]; (2) ‘Type’ [circle white label with border red, printed in black]; (3) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (4) ‘Piestus/capricornis/Er. p. 833, H. type/Guadalpia/ pointe a pitre D. Sherminier?’ [green label, handwritten]. Note: In the original description Laporte (1835) did not specify how many specimens were observed. We received only one specimen from BMNH with type label, which it is labelled as coming from Guadeloupe, and not from Cayenne (French Guiana), the original type locality. However, we are considering this as type following Blackwelder (1943), that already commented it. Piestus capricornis variety muticus Fauvel, 1902. Lectotype (here designated) deposited in IRSNB, male, with labels: (1) ‘Maroc de/Legua (Para)/30 [white label, handwritten]; (2) ‘v. muticus Fvl.’ [white label, handwritten]; (3) ‘Coll. et det. A. Fauvel/Piestus/capricornis Cast./R I.Sc. N.B. 17.479’ [white label, the first and fourth lines printed in black, second and third lines handwritten]. Note: In the original description Fauvel (1902) did not specify how many specimens he observed, but he listed two localities (see type locality above). We received from IRSNB one male from ‘Maroc de Legua (Para)/30 which is here considered as lectotype. Piestus frontalis Sharp, 1876. Holotype deposited in BMNH, sex undetermined, with labels: (1) ‘Piestus/frontalis/Amazons Type/D.S.’ [white label, together with the specimen, handwritten]; (2) ‘Holo-/type’ [circle white label with border red, printed in black]; (3) ‘Ega.’ [circle green label, handwritten]; (4) ‘S. America:/Brazil.’ [white label, printed in black]; (5) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (6) ‘Holotype/Piestus/frontalis/Sharp, 1876/det. R.G. Booth 2008’ [white label, the four first lines handwritten and the last one printed in black]. Note: In the original description Sharp (1876) specified only one specimen observed. Additional material See Appendix 2. E. Caron et al. face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six or seven setae medially, equal in length; each lateral third, antero-internal seta with same length to postero-external seta (Fig. 109). Mandibles with bifurcate apex in left and trifurcate in right (Fig. 119); dorsal tooth a little shorter or equal to ventral; internal border asymmetrical, each with one acute tooth at middle and, anterior to it, in right mandible with an area that resemble a tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/PL = 1.66) (Fig. 64); anterior angles projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter and, anterior to it, in each side with a slight emargination; apical three-quarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.99); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 166); lateral lobes exceed a little apex of median lobe in lateral view; internal sclerites as in Fig. 167. Female. Similar to male. Ovipositor as in P. sulcatus. Spermathecal duct ~2 times the length of spermatheca, with sclerotised basal region (Fig. 253); spermatheca with basal half suddenly narrower than apical (Fig. 253). Distribution In the current study, the species was examined from Guadeloupe, Peru (Loreto) and Brazil (Amazonas, Pará and Rondônia). Herman (2001b) listed P. capricornis also from Venezuela, Suriname and French Guiana. Biological notes Unknown. Redescription BL: 8.2–9.5 mm, BW: 2.2–2.4 mm. Body strongly flattened dorsoventrally; entirely reddish brown, sometimes with head and abdomen darker (Fig. 17). Dorsal integument of head and pronotum with inconspicuous fine punctures and some moderately sized punctures distributed on pronotal disc, undulate microstriae only on lateral regions (Fig. 64); striae of elytra with moderately sized punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and short frontal processes (Fig. 64), each shorter than scape, and basal distance between processes ~3 times basal width of each one; V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and prominent, and fovea at base; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 17); scape with two long setae at middle of dorsal Remarks Piestus capricornis is easily distinguished from P. puncticollis by the characters under remarks of P. puncticollis. Piestus planatus (Sharp, 1887) (Figs 18, 65, 168, 169, 254) Zirophorus planatus Sharp, 1887: 711 (original description, type localities: ‘Mexico, Cordova; British Honduras, R. Sarstoon; Guatemala, Sinanja; Nicaragua, Chontales; Panama, Volcan de Chiriqui 2000 to 3000 feet’). Piestus (Zirophorus) planatus: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1951: 7 (characters, distribution); Scheerpeltz, 1952: 283 (characters, distribution). Piestus planatus: Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1793 (distribution); Navarrete-Heredia et al., 2002: 209 (distribution). Revision of Piestus with remarks on related genera Type material Syntype deposited in FMNH, sex undetermined, labels: (1) ‘Chontales,/ Nicaragua./Janson.’ [white label, printed in black]; (2) ‘B.C.A. Col. I. 2./ Zirophorus/planatus,/Sharp.’ [white label, printed in black]; (3) ‘Sharp Coll./ 1905.-313’ [white label, printed in black]; (4) ‘Chicago Nat. Hist. Mus./ (ex. D. Sharp Colln./by exchange with/Brit. Mus. Nat. Hist.)’ [white label, printed in black]. Note: There are 11 syntypes (not examined) deposited in BMNH: ‘5 from Chontales, 3 from Rio Sarstoon, 1 from Cordova, 1 from Sinanja, 1 from V. de Chiriqui’ (Dr Roger Booth, BMNH, pers. comm.). Additional material See Appendix 2. Invertebrate Systematics 515 Female. Similar to male. Ovipositor as in P. sulcatus; spermathecal duct ~2 times length of spermatheca, with sclerotised basal region (Fig. 255); spermatheca with basal half suddenly narrower than apical (Fig. 255). Distribution In the current study the species was examined from Mexico (Chiapas, Veracruz, Guerrero and Oaxaca), Guatemala (Alta Verapaz), Nicaragua (Río San Juan), Costa Rica (Alajuela and Limón), Panama (Colón and Darién) and Ecuador (Esmeraldas). Navarrete-Heredia et al. (2002) listed P. planatus also from Belize. Biological notes Redescription BL: 8.2–10.8 mm, BW: 2.4–2.9 mm. Body strongly flattened dorsoventrally; entirely black (Fig. 18). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures, some moderately sized punctures distributed on pronotal disc (Fig. 65); striae of elytra with moderately sized punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes, each longer than scape, weakly divergent to apex, and basal distance between processes ~3 times basal width of each one (Fig. 65); frontal anterior margin with two prominent areas at middle; V-shaped frontal sulcus complete; anterior angles curved and prominent, short median longitudinal sulcus with fovea at base; eyes with two moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae almost reaching apex of abdomen (Fig. 18); scape with two long setae at middle of dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with eight or nine setae medially, equal in length; each lateral third, antero-internal seta with same length to postero-external seta. Mandibles strongly projected anteriorly; with bifurcate apices; each with dorsal tooth longer than ventral; internal border symmetrical, each with one acute tooth at middle. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/PL = 1.54) (Fig. 65); anterior angles projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter and, anterior to it, in each side with a slight emargination; apical three-quarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat shorter than wide (EL/ BW = 0.93); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 168); lateral lobes exceeding apex of median lobe by one-quarter of its length in lateral view; internal sclerites as in Fig. 169. Piestus planatus has been observed only under bark or associated with decaying logs. Remarks This species is very similar to P. spinosus mainly by the dorsal teeth of mandibles longer than the ventral teeth (Figs 65, 66, 120). However, P. planatus differs by the body being entirely black (Fig. 18), frontal processes divergent to the apex (Fig. 65) and front with two conspicuously prominent areas at the middle (Fig. 65). Piestus spinosus has head, pronotum and elytra reddish brown and abdomen reddish dark brown to black (Fig. 19), frontal processes not divergent to the apex (Fig. 66), and front with two inconspicuous prominent areas at the middle (Fig. 66). Piestus spinosus (Fabricius, 1801) (Figs 19, 69, 120, 170, 171, 254) Cucujus spinosus Fabricius, 1801: 93 (original description, type locality: ‘America meridionali’). Piestus spinosus: Laporte, 1835: 128 (characters, distribution); Erichson, 1840: 832 (characters, distribution); Fauvel, 1864: 22 (characters, distribution); Sharp, 1876: 405 (distribution); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1794 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Zirophorus) spinosus: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1933: 993 (mention); Scheerpeltz, 1951: 5 (mention), 7 (characters, distribution); Scheerpeltz, 1952: 283 (characters, distribution). Piestus oxytelinus Perty, 1830: 33 (original description, type locality: ‘Provincia Piauhiensi’). - As synonym of P. spinosus: Erichson, 1840: 832; Fauvel, 1864: 22; Bernhauer & Schubert, 1910: 6; Blackwelder, 1944: 101 (error: Perty, 1834: 33); Herman, 2001b: 1794; Newton et al., 2005: 69. Note: Scheerpeltz (1952: 283) and Scheerpeltz (1960: 66) list this name as valid and attributed it to Laporte, 1835 (error: Laporte, 1834: 129). However, the name oxytelinus sensu Laporte, 1835 is a misidentification of P. bicornis (Herman 2001b). For more details see note under P. bicornis. Type material Cucujus spinosus Fabricius, 1801. Syntype deposited in ZMUC, sex undetermined, with labels: (1) ‘TYPE’ [red label, printed in black]; (2) ‘C?: spinosus/illegible/Cucujus’ [white label, handwritten]. Note: In the original 516 Invertebrate Systematics description Fabricius (1801) did not specify how many specimens were observed. We received from ZMUC only one specimen with type label which is here considered as a syntype. Piestus oxytelinus Perty, 1830. Not found. Additional material See Appendix 2. Redescription BL: 7.8–10.5 mm, BW: 2.2–3.0 mm. Body strongly flattened dorsoventrally; head, pronotum and elytra reddish brown and abdomen reddish dark brown to black, sometimes head darker than pronotum and elytra (Fig. 19). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures, some moderately sized punctures distributed on pronotal disc (Fig. 66); striae of elytra with moderately sized punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes, each longer than scape, and basal distance between processes ~3 times basal width of each one (Fig. 66); frontal anterior margin with two prominent regions at middle, sometimes inconspicuous; V-shaped frontal sulcus complete; anterior angles curved and prominent, and short median longitudinal sulcus with fovea at base; eyes with two moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae almost reaching apex of abdomen (Fig. 19); scape with two long setae at middle of dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six, seven or eight setae medially, equal in length; each lateral third, anterointernal seta with same length to postero-external seta. Mandibles strongly projected; bifurcate apex (Fig. 120); each with dorsal tooth longer than ventral; internal border symmetrical, each with one acute tooth at middle. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times as wide as long. Pronotum wider than long (PW/ PL = 1.50) (Fig. 66); anterior angles projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter and, anterior to it, in each side with slight emargination; apical three-quarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat shorter than wide (EL/BW = 0.90); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 170); lateral lobes exceeding the apex of median lobe by one-quarter of its length in lateral view; internal sclerites as in Fig. 171. Female. Similar to male. Ovipositor as in P. sulcatus; spermathecal duct ~2 times length of spermatheca, with sclerotised basal region (Fig. 255); spermatheca with basal half suddenly narrower than apical (Fig. 255). E. Caron et al. Distribution In the current study, the species was examined from Venezuela (Bolívar), French Guiana (Saint-Laurent-du Maroni and Cayenne), Ecuador (Napo and Pastaza), Peru (Loreto, Huánuco, Lima, Junín, Ucayali and Cusco), Bolivia (Cochabamba and Santa Cruz) and Brazil (Amazonas, Pará, Rondônia and Mato Grosso). Herman (2001b) listed P. spinosus also from Colombia, Suriname and Guyana. Biological notes Piestus spinosus has been found under bark or in leaf litter near decaying logs. One specimen was collected in a flight intercept trap and another was labelled as ‘beating palm branches’. In Bolivia one specimen was observed in Ficus sp. Remarks Piestus spinosus is very similar P. planatus but differs by characters listed under remarks of P. planatus. Piestus longipennis Fauvel, 1864 (Figs 20, 67, 172–175, 256) Piestus longipennis Fauvel, 1864: 20 (original description, type locality: ‘Colombie’); Blackwelder, 1944: 100 (distribution, error: Fauvel 1865: 24); Herman, 2001b: 1792 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Zirophorus) longipennis: Bernhauer & Schubert, 1910: 6 (distribution, error Fauvel, 1865: 24); Scheerpeltz, 1951: 6 (characters, distribution, error: Fauvel, 1865: 25); Scheerpeltz, 1952: 282 (characters, distribution, error: Fauvel, 1865: 25). Type material Holotype deposited in IRSNB, sex undetermined, with labels: (1) ‘Colombie’ [white label, Fauvel’s handwritten]; (2) ‘longipennis/Fvl.’ [white label, Fauvel’s handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black, and second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1864) specified only one specimen observed. Additional material See Appendix 2. Redescription BL: 9.2–10.5 mm, BW: 2.4–2.8 mm. Body flattened dorsoventrally; entirely dark brown to black (Fig. 20). Dorsal integument of head and pronotum with inconspicuous fine punctures, undulate microstriae only on lateral regions (Fig. 67); striae of elytra unpunctured; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes each longer than scape, and basal distance between processes the same or narrower than basal width of each one (Fig. 67); Vshaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin of eyes, equal in length. Antennae almost reaching apex of elytra (Fig. 20); scape with some few long setae at middle of dorsal face; antennomeres 3–6 with some long setae on dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 Revision of Piestus with remarks on related genera oblong shape. Labrum with six setae medially, internal the longest; on each lateral third, antero-internal seta longer than postero-external seta. Mandibles strongly projected with bifurcate apices; each with dorsal tooth shorter than ventral; internal border with one acute tooth at middle and, anterior to it, in right mandible with another acute tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/PL = 1.38) (Fig. 67); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles not developed; abrupt constriction at basal quarter; apical three-quarters with parallel sides. Femur of anterior legs with long hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.08); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; sternite 7 with longitudinal set of long setae near of each lateral margin; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, basal the longest. Median lobe of the aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 172); apex of median lobe strongly emarginate in ventral view (Fig. 174); lateral lobes almost reaching apex of median lobe in lateral view; internal sclerites as in Figs 173 and 175. Female. Similar to male. Ovipositor as in P. fronticornis; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 256); spermatheca as in Fig. 256. Distribution In the current study the species was examined from Colombia (Valle del Cauca) and Ecuador (Carchi, Pichincha, Napo, Cotopaxi and Tungurahua). Herman (2001b) listed this species also from Venezuela. Biological notes Piestus longipennis is only known from under bark of decaying logs. Remarks Piestus longipennis is similar to P. zischkai and easily separated from that by the frontal processes on head of male not converging to the apex (Fig. 67) and lateral lobes of aedeagus never exceeding the apex of median lobe in lateral view (Fig. 172). Piestus zischkai has frontal processes of male converging to the apex (Fig. 68) and the lateral lobes exceeding the apex of median lobe (Fig. 176). Piestus longipennis, as well as P. zischkai, differs from the other closely related species (P. bicornis, P. fronticornis and P. validus) mainly by the body being entirely black and the presence of a bulbous base of the median lobe of aedeagus in ventral view. Piestus zischkai Scheerpeltz, 1951 (Figs 21, 68, 69, 176–178) Piestus (Zirophorus) zischkai Scheerpeltz, 1951: 4 (original description, type locality: ‘Yungas del Palmar’, ‘200 m’); Scheerpeltz, 1952: 282 (characters, distribution). Piestus zischkai: Herman, 2001b: 1795 (distribution). Invertebrate Systematics 517 Type material Piestus (Zirophorus) zischkai Scheerpeltz, 1951. Lectotype (here designated) deposited in NMW, male [damaged specimen: without anterior and left posterior tarsi], with labels: (1) ‘<’ [white label, printed in black]; (2) ‘Bolivia: 2000 m/Yungas del Palmar/15. III. 1951/leg R. Zischka’ [white label, first (except ‘Bolivia’), second and third lines handwritten, ‘Bolivia’ in the first line and fourth line printed in black]; (3) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (4) ‘TYPUS/Piestus/ zischkai/O. Scheerpeltz’ [red label, the first and fourth lines printed in black, second and third handwritten]; (5) ‘Zischkai/Scheerp.’ [green label, handwritten]. Paralectotype deposited in NMW, female [damaged specimen: without last tarsomere of left median leg], with labels: (1) ‘,’ [white label, printed in black]; (2) ‘Bolivia: 2000 m/Yungas del Palmar/15. III. 1951/leg R. Zischka’ [white label, first (except ‘Bolivia’), second and third lines handwritten, ‘Bolivia’ in the first line and fourth line printed in black]; (3) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (4) ‘TYPUS/Piestus/ zischkai/O. Scheerpeltz’ [red label, the first and fourth lines printed in black, second and third handwritten]. Note: In the original description Scheerpeltz (1951) cited two males and one female observed. However, we received from NMW one male and one female with type labels. Additional material See Appendix 2. Redescription BL: 14.5–15.8 mm, BW: 2.3–2.5 mm. Body flattened dorsoventrally; entirely dark brown to black (Fig. 21). Dorsal integument of head and pronotum with inconspicuous fine punctures, undulate microstriae only on lateral regions (Fig. 68); striae of elytra unpunctured; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes each longer than scape, strongly converge to apex, and basal distance between processes the same or narrower than basal width of each one (Fig. 68); V-shaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, equal in length. Antennae reaching apex of elytra (Fig. 21); with some few long setae at middle of dorsal face; antennomeres 3–6 with some long setae on dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the longest; on each lateral third, antero-internal seta with same length to postero-external seta. Mandibles with bifurcate apices; each with ventral tooth anteriorly weakly projected and dorsal tooth shorter than ventral; internal border with one acute tooth at middle and, anterior to it, in right mandible with other acute tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/PL = 1.25) (Fig. 68); anterior angles projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles not developed; abrupt constriction at basal quarter; apical threequarters with parallel sides. Femur of anterior legs with long hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.10); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; 518 Invertebrate Systematics sternite 7 with longitudinal set of long setae near of each lateral margin; tergite 9 with short ventral struts; tergite 10 with two pairs of short setae on apex, equal in length. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 176); apex of median lobe emarginate in ventral view (Fig. 178); lateral lobes exceed little the apex of median lobe in lateral view; internal sclerites as in Fig. 177. Female. Similar to male, except for: frontal processes of head not converging to apex and internal margins are straight. Ovipositor as in P. fronticornis; spermathecal duct with sclerotised basal region (length unknown); spermatheca as in P. longipennis. Distribution In the current study the species was examined from Ecuador (Tungurahua and Zamora-Chinchipe), Peru and Bolivia. Biological notes E. Caron et al. of P. bicornis (‘Cet insecte n’est peut-être qu’une variété du précédent’ Laporte 1835: 128). Piestus oxytelinus: Laporte, 1835: 128. Scheerpeltz, 1952: 283 (error: Laporte, 1834: 129); Scheerpeltz, 1960: 66 (error: Laporte, 1834: 129); Newton et al., 2005: 37 (error: Laporte 1835: 129). Note: the name P. oxytelinus by Laporte (1835) is not an available name, but rather a misidentification of P. oxytelinus Perty, 1830, which is a junior synonym of P. spinosus (see above, P. spinosus). Type material Oxytelus bicornis Olivier, 1811. Neotype (here designated) deposited in FMNH, sex undetermined, with labels: ‘15.544’ [white label, handwritten], ‘Bras.S.Catharina/illegible VIII 10/lg. Luderwaldd.’ [white label, handwritten], ‘bicornis Ol./Ihering./det. Bernh.’ [white label, the first two lines handwritten, third line printed in black], ‘Chicago NHMus/M. Bernhauer/Collection’ [white label, printed in black]. Note: In the original description, Olivier (1811) did not specify how many specimens were observed. The type material has not been found and probably it is lost. Zirophorus striatus Guérin-Méneville, 1829. Not found. There is one specimen of P. zischkai with a note that it was collected using a Malaise trap. Additional material Remarks Redescription This species is very similar to P. longipennis, from which it is easily separated by characters cited under remarks of P. longipennis. BL: 7.3–9.3 mm, BW: 2.1–2.4 mm. Body flattened dorsoventrally; brown to black with apical half of segment 7 and segments 8–10 of abdomen yellowish brown, or head and abdomen dark brown to black with pronotum and elytra reddish brown, also with apical half of segment 7 and segments 8–10 of abdomen yellowish brown (Figs 22, 23). Dorsal integument of head and pronotum with inconspicuous fine punctures, undulate microstriae only on lateral regions; striae of elytra unpunctured (Fig. 70); metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes, each longer than scape, and basal distance between processes the same or narrower than basal width of each one (Fig. 70); Vshaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with a line of three moderately sized punctures with long setae on basal half of dorsal margin, equal in length. Antennae almost reaching apex of elytra (Figs 22, 23); with some long setae at middle of the dorsal face; antennomeres 3–6 with some long setae on dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the longest; each lateral third, antero-internal seta with same length to postero-external seta. Mandibles strongly projected; bifurcate apices; each with dorsal tooth shorter than the ventral; internal border with one acute tooth at middle and, anterior to it, in right mandible with other acute tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/PL = 1.40) (Fig. 70); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles not developed; abrupt constriction at basal quarter; apical three-quarters with parallel sides. Femur of anterior legs with long hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.04); stria 6 at Piestus bicornis (Olivier, 1811) (Figs 22, 23, 70, 150, 179–181, 257) Oxytelus bicornis Olivier, 1811: 615 (original description; type locality: not given). Piestus bicornis: Laporte, 1835, 126 (mention), 128 (characters, distribution); Erichson, 1840: 831 (characters, distribution); Fauvel, 1864: 19 (characters, distribution); Sharp, 1876: 405 (mention, distribution); Fauvel, 1901: 70 (mention); Blackwelder, 1944: 100 (distribution, error: species attributed to Laporte, 1834: 128); Herman, 2001b: 1789 (distribution); Newton et al., 2005: 37, 69 (distribution, synonyms); Caron et al. 2008: 6 (mention). Zirophorus bicornis: Sharp, 1887: 712 (distribution); Sharp & Muir, 1912 (characters of aedeagus). Piestus (Zirophorus) bicornis: Bernhauer & Schubert, 1910: 6 (distribution, error: species attributed to Laporte, 1834: 128); Scheerpeltz, 1951: 6 (characters, distribution, error: species attributed to Laporte, 1834: 128); Scheerpeltz, 1952: 282 (characters, distribution, error: species attributed to Laporte, 1834: 128). Zirophorus striatus Guérin-Méneville, 1829: pl. 9, fig. 12 (type locality: not given). - As synonym of Piestus bicornis: Laporte, 1835: 126, 128; Erichson, 1840: 832; Fauvel, 1864: 20; Bernhauer & Schubert, 1910: 6; Blackwelder, 1944: 100 (note: cited as ‘of authors’); Herman, 2001b: 1789; Newton et al., 2005: 69. Piestus bicornis variety oxytelinus: Laporte, 1835: 128. Erichson, 1840: 832; Fauvel, 1864: 20; Sharp, 1976: 405; Sharp, 1887: 712; Bernhauer & Schubert, 1910: 6 (error: Laporte, 1834: 129); Blackwelder, 1944: 100 (error: Laporte, 1834: 129); Scheerpeltz, 1951: 5, 6 (error: Laporte, 1834: 129); Newton et al., 2005: 37. Note: Laporte, 1835 did not describe the species as new, he attributed it to Perty (1830), but probably it was a misidentification of P. oxytelinus Perty (Herman 2001b). However, Laporte (1835) cited the similarity between P. bicornis and P. oxytelinus and suggested that P. oxytelinus could be a variety See Appendix 2. Revision of Piestus with remarks on related genera basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; sternite 7 with longitudinal set of long setae near of each lateral margin; tergite 9 with long ventral struts (Fig. 150); tergite 10 with two pairs of long setae on apex, basal the longest. Median lobe of aedeagus without bulbous base in ventral view and not curved apex in lateral view (Figs 179, 181); apex of median lobe slightly emarginate in ventral view (Fig. 181); lateral lobes not reaching apex of median lobe in lateral view; internal sclerites as in Fig. 180. Female. Similar to male. Ovipositor as in P. fronticornis; spermathecal duct ~5 times length of spermatheca, without sclerotised basal region (Fig. 257); spermatheca as in Fig. 257. Distribution In the current study, the species was examined from Costa Rica (Cartago and Limón), Panama (Panamá), Colombia (Cundinamarca, Meta and Putumayo), Venezuela (Carabobo), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Sucumbios, Napo, Pastaza and Morona-Santiago), Peru (Loreto, San Martín, Huánuco, Junín, Cusco and Madre de Dios), Bolivia (La Paz, Beni, Cochabamba and Santa Cruz), Brazil (Acre, Amazonas, Pará, Goiás, Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo, Paraná and Santa Catarina) and Paraguay (Guairá). Herman (2001b) listed P. bicornis also from Nicaragua, Suriname, Guyana and Argentina. Biological notes Piestus bicornis has been found under bark or in leaf litter near decaying logs. Some specimens were collected in Malaise trap and others were captured by ‘beating palm branches’. This species also has been noticed occurring together with P. validus, P. fronticornis, P. pygmaeus and P. minutus. The specimens with black pronotum and elytra were only observed in Brazil (Minas Gerais, Espirito Santo, Rio de Janeiro, São Paulo and Paraná e Santa Catarina) and Paraguay (Guairá). The specimens with reddish brown pronotum and elytra were recorded from Costa Rica to Brazil (Acre, Amazonas, Pará, Goiás, São Paulo and Paraná). Invertebrate Systematics 519 tergite 10 with two pairs of long setae on the apex, and apex of median lobe very thin and sclerite of internal sac inconspicuous (Figs 182, 183). Piestus validus has a brown to black body with apical half of segment 7 and segments 8–10 yellowish (Fig. 25), antennal scape with many long setae on the apical half of the dorsal face and antennomeres 3–6 with many long setae on dorsal face, tergite 9 of male with long ventral struts, tergite 10 with one pair of long setae on the apex, and with a well-developed apex of median lobe and sclerite of internal sac (Figs 184, 185). There are two colour patterns in P. bicornis. Specimens with black pronotum and elytra (Fig. 22) or specimens with reddish brown pronotum and elytra (Fig. 23). These patterns were first observed by Laporte (1835), who misidentified the specimens with reddish brown pronotum and elytra as P. oxytelinus Perty, 1830 (junior synonym of P. spinosus) and cited its close similarity to P. bicornis (specimens with black pronotum and elytra), ‘Cet insecte n’est peut-être qu’une variété du précédent’ (Laporte 1835: 128). Since then, many authors have identified the specimens with reddish brown pronotum and elytra as a variety of P. bicornis, P. bicornis variety oxytelinus (Erichson 1840; Fauvel 1864; Sharp 1876, 1887; Scheerpeltz 1951), or as a valid species, P. oxytelinus Laporte, 1835 (Scheerpeltz 1952, 1960; Newton et al. 2005). However, in this study no significant morphological differences between the specimens with the two colour patterns have been found, the morphology of the aedeagus and spermatheca being the same. Different colour patterns in the same species were also observed in P. pennicornis (see below). Piestus fronticornis (Dalman, 1821), sp. rev. (revalidated species) (Figs 24, 71, 110, 121, 122, 130, 133, 143, 154, 182, 183, 258) Zirophorus fronticornis Dalman, 1821: 373 (original description, type locality: ‘Kalkoen’). - As synonym of P. bicornis: Laporte, 1835: 126, 128; Erichson, 1840: 832; Fauvel, 1864: 20; Sharp, 1887: 712; Bernhauer & Schubert, 1910: 6; Blackwelder, 1944: 100; Herman, 2001b: 1789; Newton et al., 2005: 69. Remarks Type material This species is similar to P. fronticornis and P. validus, and can be separated from them by having a body brown to black with apical half of segment 7 and segments 8–10 yellowish brown, or head and abdomen dark brown to black with pronotum and elytra reddish brown and also with apical half of segment 7 and segments 8–10 yellowish brown (Figs 22, 23), antennal scape with a few long setae on the apical half of the dorsal face and antennomeres 3–6 with a few long setae on dorsal face, tergite 9 of male with long ventral struts (Fig. 150), tergite 10 with two pairs of long setae on the apex, and with a welldeveloped apex of median lobe and sclerite of internal sac (Figs 179, 180). Piestus fronticornis has the following characters: body brown to black with all of segment 7 and segments 8–10 yellowish (Fig. 24), antennal scape with long setae on the apical half of the dorsal face and antennomeres 3–6 with long setae on dorsal face, tergite 9 of male with short ventral struts, Syntype deposited in NHRS, sex undetermined [damaged specimen: left antenna without antennomere 6–11; inverted mandibles; without right anterior and median tarsomeres; without two last tarsomeres of the left posterior leg; and without left median leg., with labels: (1) ‘Mus./Pay k’ [white label, printed in black]; (2) ‘fronticornis Dalm.’ [white label, handwritten]; (3) ‘6402/E91 +’ [light blue label, printed in black]. Mouthparts (labrum, maxillae and mentum) are glued on a piece of paper which is pinned separately and labelleds: ‘6401/E91 +’ [light blue label, printed in black]; ‘6400/E91 +’ [light blue label, printed in black]; ‘6399/ E91 +’ [light blue label, printed in black]; ‘6398/E91 +’ [light blue label, printed in black]; ‘buccal parts [handwritten in light green]/det. J. Ferrer [printed in black]’ [white label]. Note: In the original description Dalman (1821) did not specify how many specimens he observed, and he did not know the location of the type locality ‘Kalkoen’. We received from NHRS one specimen, regarded as a syntype. Additional material See Appendix 2. 520 Invertebrate Systematics E. Caron et al. Redescription Biological notes BL: 7.9–10.8 mm, BW: 1.9–2.8 mm. Body flattened dorsoventrally; brown to black with all of segment 7 and segments 8–10 yellowish (Fig. 24). Dorsal integument of head and pronotum with inconspicuous fine punctures, undulate microstriae only on lateral regions (Fig. 71); striae of elytra unpunctured; metaventrite, on lateral regions, and abdominal tergites on anterolateral regions with moderately sized punctures. Male. Head with pair of acute and long frontal processes, each longer than scape, and basal distance between processes same or narrower than basal width of each one (Fig. 71); V-shaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, equal in length. Antennae almost reaching apex of elytra (Fig. 24); scape with some few long setae on apical half of dorsal face; antennomeres 3–6 with some few long setae on dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the longest; each lateral third, antero-internal seta with same length to postero-external seta. Mandibles strongly projected (Figs 71, 121, 122); with bifurcate apices; each with dorsal tooth shorter than ventral; internal border with one acute tooth at middle and, anterior to it, in right mandible with other acute tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than 2 and 3 combined (Fig. 130). Mentum 2 times wider than long (Fig. 133). Pronotum wider than long (PW/PL = 1.41) (Fig. 71); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles not developed; abrupt constriction at basal quarter; apical three-quarters with parallel sides. Femur of anterior legs with long hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.01); stria 6 at basal half sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; sternite 7 with longitudinal set of long setae near of each lateral margin; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, basal the longest. Median lobe of aedeagus without bulbous base in ventral view and not curved apex in lateral view (Fig. 182); lateral lobes almost reaching apex of median lobe in lateral view; internal sclerites as in Fig. 183; apex of median lobe slightly emarginate in ventral view. Female. Similar to male. Ovipositor strongly emarginate at external margin (Fig. 154); spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 258); spermatheca as in Fig. 258. Piestus fronticornis has been found on or under bark of decaying logs. Some specimens were collected in flight intercept traps and others were found on ‘decaying banana, blossoms, leaves’. This species has been noticed occurring together with P. bicornis, P. pygmaeus and P. minutus. Distribution Nicaragua (Granada and Rio San Juan), Guatemala (Guatemala), Costa Rica (Guanacaste, Alajuela, Heredia, San José, Limón and Puntarenas), Panama (Bocas del Toro, Chiriquí, Veraguas, Colón, Panamá and Darién), Colombia (Cundinamarca and Valle del Cauca), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Esmeraldas and Pastaza) and Brazil (Amazonas, Amapá and Pará). Remarks This species is easily separated from those by characters listed in the remarks under P. bicornis. Piestus validus Sharp, 1876 (Figs 25, 72, 184, 185, 259) Piestus validus Sharp, 1876: 404 (original description, type locality: ‘Pebas’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1795 (distribution); Caron et al., 2008: 6 (mention). Piestus (Zirophorus) validus: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1951: 6 (characters, distribution); Scheerpeltz, 1952: 283 (characters, distribution). Type material Three syntypes (not examined) deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). Note: In the original description Sharp (1876) specified three specimens observed. Additional material See Appendix 2. Redescription BL: 8.3–11.3 mm, BW: 2.2–3.2 mm. Body flattened dorsoventrally; brown to black with apical half of segment 7 and segments 8–10 yellowish (Fig. 25). Dorsal integument of dorsum of head and pronotum with inconspicuous fine punctures, undulate microstriae only on lateral regions (Fig. 72); striae of elytra unpunctured; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with pair of acute and long frontal processes, each longer than scape, and basal distance between processes the same or narrower than basal width of each one (Fig. 72); Vshaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with a line of three moderately sized punctures with long setae on basal half of dorsal margin, equal in length. Antennae almost reaching apex of elytra (Fig. 25); scape with many long setae on apical half of the dorsal face; antennomeres 3–6 with many long setae on dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the longest; on each lateral third, the antero-internal seta with same length to postero-external seta. Mandibles strongly projected; with bifurcate apices; each with dorsal tooth shorter than ventral; internal border with acute tooth at middle and, anterior to it, in right mandible with other acute tooth, absent in left mandible. Maxillary palpus with palpomere 4 shorter than the 2 and 3 combined. Mentum 2 times wider than long. Pronotum wider than long (PW/PL = 1.40) (Fig. 72); anterior angles weakly projected; longitudinal median sulcus Revision of Piestus with remarks on related genera conspicuous; longitudinal sulcus of posterior angles inconspicuous; abrupt constriction at basal quarter; apical three-quarters with parallel sides. Femur of anterior legs with long hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.05); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; sternite 7 with longitudinal set of long setae near of each lateral margin; tergite 9 with long ventral struts; tergite 10 with pair of long setae on apex. Median lobe of aedeagus without bulbous base in ventral view and not curved apex in lateral view (Fig. 184); apex of median lobe slightly emarginate in ventral view; lateral lobes not almost reaching apex of median lobe in lateral view; internal sclerites as in Fig. 185. Female. Similar to male. Ovipositor as in P. fronticornis; spermathecal duct ~2 times length of spermatheca, with sclerotised basal region (Fig. 259); spermatheca as in Fig. 259. Distribution In the current study the species was examined from Colombia (Meta), Venezuela (Carabobo, Aragua), Ecuador (Napo, Pastaza and Loja), Peru (Junín, Huánuco, Cusco and Madre de Dios), Bolivia (La Paz and Cochabamba) and Brazil. Biological notes Piestus validus has been found under bark or in litter near decaying logs. Some specimens were collected in flight intercept traps and others were captured by ‘beating palm branches’. This species also has been noticed occurring together with P. bicornis. Remarks This species is very similar to P. bicornis and P. fronticornis but differs by characters cited above (see remarks under P. bicornis). Three syntypes deposited in BMNH were not examined. However, at the moment, the original description (Sharp 1876) and a reasonable quantity of specimens identified from many institutions were considered sufficient for positive identification, but the study of type material would be advisable for confirming the identification of this species. Piestus rossii, sp. nov. (Figs 26, 73, 186, 187) Invertebrate Systematics 521 conspicuous fine punctures equally distributed; striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites on anterolateral regions with moderately sized punctures. Male. Head with front slightly deflected (Fig. 73); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved, weakly prominent and with median fovea at base; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin of eyes, equal in length, and additional set, not line, of five moderately sized punctures with long setae, shorter than and internal to the other ones. Antennae reaching apex of abdomen (Fig. 26); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomere 2 with many long setae on apical half of dorsal face; antennomeres 4–11 oblong shape and slightly narrowing from antennomere 4 to 11. Mandibles with dorsal teeth developed, not forming bifurcate apex, shorter than ventral; internal border asymmetrical, each with one acute tooth at middle and, anterior to it, in right mandible with an area that resembles a tooth, absent in left mandible; external-dorsal margin on basal half of each mandible slightly carinate. Maxillary palpus with palpomere 4 subequal in length to 2 and 3 combined. Pronotum wider than long (PW/PL = 1.48) (Fig. 73); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical three-quarters with slightly curved sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.98); stria 6 at basal half and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 186); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 187. Female. Unknown. Distribution Ecuador (Cotopaxi). Biological notes Unknown. Type material Remarks Holotype deposited in DZUP, male, labels: (1) ‘Ecuador – Cotopaxi/ Canton Sigchos – Las/Pampas – Otonga Natural/Reserve – 25-28, VII,2005/W. Rossi, leg.’ [white label, printed in black]; (2) ‘<’ [white label, printed in black]. Piestus rossii is similar to P. pennicornis, differing from that species by the single line of three moderately sized punctures and an additional set, not in line, of five moderately sized punctures with long setae on basal half of dorsal margin of eyes, conspicuous fine punctures on pronotum and curved lateral sides of pronotum (Fig. 73), and curved apex of median lobe of aedeagus (Fig. 186). Piestus pennicornis has a single line of three moderately sized punctures with long setae, inconspicuous fine punctures and subparallel sides (Figs 74, 75), and uncurved apex of median lobe (Fig. 188). Description BL: 6.8 mm, BW: 1.8 mm. Body flattened dorsoventrally; black with apical half of segment 7 and segments 8–10 yellowish (Fig. 26). Dorsal integument of head with somewhat microgranulate and inconspicuous fine punctures (Fig. 73); pronotum with undulate microstriae only on lateral areas and 522 Invertebrate Systematics Etymology The species is dedicated to Walter Rossi, Italian mycologist, who collected the holotype and made it available. Piestus pennicornis Fauvel, 1864 (Figs 27, 74, 75, 123, 124, 188, 189, 260) Piestus pennicornis Fauvel, 1864: 26 (original description, type locality: ‘Colombie’); Blackwelder, 1944: 101 (distribution, error: Fauvel 1865: 30); Herman, 2001b: 1793 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Piestus) pennicornis: Bernhauer & Schubert, 1910: 7 (distribution, error: Fauvel, 1865: 30); Scheerpeltz, 1952: 289 (characters, distribution, error: Fauvel 1865: 30). Piestus plagiatus Fauvel, 1864: 26 (original description, type locality: ‘Nuovo-Friburgo, Brésil’); Blackwelder, 1944: 101 (distribution, error: Fauvel 1865: 30); Herman, 2001b: 1793 (distribution). New synonym. Piestus (Piestus) plagiatus: Bernhauer & Schubert, 1910: 7 (distribution, error: Fauvel, 1865: 30); Scheerpeltz, 1952: 289 (characters, distribution, error: Fauvel 1865: 30). Piestus rectus Sharp, 1876: 406 (original description, type locality: ‘Ega’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1794 (distribution). New synonym. Piestus (Piestus) rectus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 288 (characters, distribution). Piestus pygialis Fauvel, 1902: 23 (original description, type locality: ‘Brésil: Blumenau’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1793 (distribution). New synonym. Piestus (Piestus) pygialis: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 288 (characters, distribution). Piestus surinamensis Bernhauer, 1928: 287 (original description, type locality: ‘Suriname’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1795 (distribution). Note: In the original description Bernhauer (1928) gives the information about the type locality at the beginning of the paper for all species described in that paper. New synonym. Piestus (Piestus) surinamensis: Scheerpeltz 1933: 994 (distribution); Scheerpeltz, 1952: 289 (characters, distribution). Type material Piestus pennicornis Fauvel, 1864. Lectotype (here designated) deposited in IRSNB, female, with labels: (1) ‘Colombie’ [white label, handwritten]; (2) ‘pennicornis/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/ Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in red]. Note: In the original description Fauvel (1864) specified two specimens, male and female, studied from Colombia (‘Deux exemplaires < ,’). However, we received from IRSNB three specimens with type label (‘Ex-Typis’): one female from ‘Colombie’ (here considered as syntype) and two from ‘Para’, which we consider additional material. Piestus plagiatus Fauvel, 1864. Lectotype (here designated) deposited in IRSNB, female, with labels: (1) ‘N
Friburgo’ [white label, handwritten]; (2) ‘plagiatus/FvL.’ [white label/handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/ Coll. et. det. A. Fauvel’ [white label, printed in black]; (4) ‘Ex-Typis’ [white label, printed in red]. Note: In the original description Fauvel (1864) specified two specimens observed. We received from IRSNB only one specimen with type label. Piestus rectus Sharp, 1876. Syntype deposited in FMNH, female, labels: (1) ‘Piestus rectus/amazons./Ind. typ./D.S.’ [white label, together with the specimen, handwritten]; (2) ‘Ega.’ [green label, handwritten]; (3) ‘S.America: Brazil.’ [white label with transverse green line, printed in black]; (4) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (5) ‘Chicago Nat. Hist. Mus./ E. Caron et al. (ex. D. Sharp Colln./by exchange with/Brit. Mus. Nat. Hist.)’ [white label, printed in black]. Note: In the original description Sharp (1876) specified four specimens, ‘probably all females’, studied from ‘Ega’ and more one male from the same locality. However, we have only seen one syntype deposited in FMNH. The other 4 syntypes (not examined) are deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). Piestus pygialis Fauvel, 1902. Holotype deposited in IRSNB, female, with labels: (1) ‘Blumenau/Brésil’ [white label, handwritten]; (2) ‘pygialis/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third line printed in black and second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1902) specified only one specimen observed. Piestus surinamensis Bernhauer, 1928. Holotype deposited in FMNH, male [damaged specimen: without right antenna and antennomeres 3–11 of left; head separated of body], with labels: (1) ‘Suriname/Uyttenboogaart/ 8-9 1900’ [white label, printed in black]; (2) ‘2/52’ [white label, printed in black]; (3) ‘Surinamen-/sis Bernh./Typ. un.’ [white label, handwritten]; (4) ‘Piestus/Surinamensis/Bernh. Typus un.’ [light yellow label, Bernhauer’s handwritten]; (5) ‘Chicago NHMus/M.Bernhauer/Collection’ [white label, printed in black]. Note: In the original description Bernhauer (1928) specified only one specimen observed. Additional material See Appendix 2. Redescription BL: 5.0–7.1 mm, BW: 1.3–1.6 mm. Body flattened dorsoventrally; entirely reddish brown to black with all or apical half of segment 7 and segments 8–10 yellowish; or head and abdomen dark brown to black with pronotum and elytra reddish brown; or head, pronotum and abdomen dark brown to black and elytra reddish brown with apical, internal and external margin darker (Fig. 27). Integument of dorsum of head with undulate microstriae and inconspicuous fine punctures (Figs 74, 75); pronotum with undulate microstriae only on lateral areas and inconspicuous fine punctures equally distributed; striae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites on anterolateral regions with moderately sized punctures. Male. Head with front slightly deflected (Fig. 74); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved, prominent and with a median fovea at base; eyes slightly prominent from above, and a line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 27); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomere 2 with many long setae on apical half of dorsal face; antennomeres 4–11 oblong shape and conspicuously narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; each lateral third, the antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth developed (Fig. 123), not forming a bifurcate apex, shorter than ventral; internal border asymmetrical, each with one acute tooth at middle and, anterior to it, in the right mandible with area that resemble a tooth, absent in left mandible; external-dorsal margin on basal half slightly carinate. Maxillary palpus with palpomere 4 subequal in length to 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.43) Revision of Piestus with remarks on related genera (Fig. 74); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical three-quarters with subparallel sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.06); stria 6 at basal half, sometimes inconspicuous and stria 7 at apical half, sometimes inconspicuous; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 188); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 189. Female. Similar to male, except for: vertex of head with anterior angles not prominent as on male (Fig. 75); antennae exceeding apex of elytra, but not reaching apex of abdomen; scape as in male, but fewer setae and without enlarged area; antennomere 2 with few setae on dorsal face; internal border of right mandible with only one tooth, broad (Fig. 124); ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, without sclerotised basal region (Fig. 260); spermatheca as in Fig. 260. Distribution In the current study the species was examined from Nicaragua (Matagalpa and Granada), Panama (Colón and Veraguas), Colombia, Venezuela, Suriname, Ecuador (Sucumbios, Napo), Peru (Cusco), Bolivia (Cochabamba), Brazil (Rondônia, Pará, Rio de Janeiro, São Paulo, Paraná and Santa Catarina) and Paraguay (Caazapá). Herman (2001b) listed the species also from Guyana and French Guiana. Biological notes Piestus pennicornis has been found under bark of decaying logs. Some specimens were captured by ‘beating palm branches’. Remarks Piestus pennicornis is similar to P. rossii and differs by some characters listed above (see remarks under P. rossii). Specimens of P. pennicornis have different colour patterns. Colour patterns were also observed in P. bicornis (see above, remarks in P. bicornis). Piestus pygmaeus Laporte, 1835 (Figs 28, 76, 190, 191, 261) Piestus pygmaeus Laporte, 1835: 130 (original description, type locality: ‘Colombie’); Erichson, 1840: 835 (characters, distribution); Fauvel, 1864: 29 (characters, distribution); Sharp, 1876: 407 (distribution); Sharp, 1887: 714 (distribution); Blackwelder, 1943: 49 (characters, distribution, notes); Blackwelder, 1944: 101 (distribution, error: Laporte, 1834: 130); Herman, 2001b: 1793 (distribution); NavarreteHeredia et al., 2002: 209 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Piestus) pygmaeus: Bernhauer & Schubert, 1910: 7 (distribution, error: Laporte, 1834: 130); Scheerpeltz, 1933: 994 (mention); Invertebrate Systematics 523 Scheerpeltz, 1952: 290 (characters, distribution, error: Laporte, 1834: 130). Piestus rufipennis Fleutiaux & Sallé, 1889: 382 (cited as synonym of P. pygmaeus Laporte, 1835, note: manuscript name by Chevrolat). - As synonym of P. pygmaeus: Blackwelder, 1943: 49, 562; Blackwelder, 1944: 100; Newton et al., 2005: 69. Note: Nomen nudum (Newton et al. 2005). Type material Piestus pygmaeus Laporte, 1835. Lectotype (here designated) deposited in BMNH, sex undetermined [damaged apex of the antennae and right anterior leg], with labels: (1) ‘Type’ [circle white label with border red, printed in black]; (2) ‘Pygmaeus/Cart? Colomb.’ [white label, handwritten]; (3) ‘Sharp Coll./1905.-313.’ [white label, printed in black]; (4) ‘Ex mus. Castel-?/nau./probably type of/Piestus pygmaeus’ [white labels, handwritten]; (5) ‘B.C.A. Col. I. 2./Piestus/pygmaeus,/Lap.’ [white label, printed in black]. Paralectotype deposited in BMNH, sex undetermined [damaged apex of the left antenna, right antenna and right anterior leg], and with the same labels of lectotype [lectotype and paralectotype glued on the same card]. In the original description, Laporte (1835) did not specify how many specimens were observed. We received two specimens with type labels. Additional material See Appendix 2. Redescription BL: 3.1–3.8 mm, BW: 0.8–1.00 mm. Body flattened dorsoventrally; head, pronotum and abdomen light brown to black and elytra light brown to reddish, always lighter than body (Fig. 28). Dorsal integument of head with undulate microstriae and inconspicuous fine punctures (Fig. 76); pronotum with undulate microstriae, sometimes absent near longitudinal sulcus, and conspicuous fine punctures equally distribute, some moderately sized punctures on median longitudinal sulcus; striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front slightly deflected (Fig. 76); V-shaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, middle very short and basal the longest. Antennae reaching apex of abdomen (Fig. 28); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and slightly narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with a very weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/ PL = 1.36) (Fig. 76); anterior weakly projected; longitudinal media sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching basal third of pronotum; abrupt constriction at basal quarter; apical three-quarters with parallel 524 Invertebrate Systematics sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/ BW = 1.08); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 190); lateral lobes exceeding apex of median lobe by one-quarter of its length in lateral view, and apical half much wider than basal; internal sclerites as in Fig. 191. Female. Similar to male, except for: antennae exceed a little the apex of elytra; scape with fewer setae; ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 261); spermatheca with basal half suddenly narrower than apical (Fig. 261). Distribution In the current study the species was examined from Mexico (San Luís Potosí, Jalisco, Veracruz, México, Oaxaca, Chiapas and Nayarit), Belize (Orange Walk, Belize and Toledo), Guatemala (Zacapa and Escuintla), Honduras (Santa Bárbara, Colón and Francisco Morazán), Nicaragua (Granada and Río San Juan), Costa Rica (Guanacaste, Heredia, Cartago, Limón and Puntarenas), Panama (Bocas del Toro, Chiriquí, Veraguas, Colón and Panamá), Guadeloupe, Saint Vincent and the Grenadines (Saint Vincent), Grenada, Trinidad and Tobago (Trinidad), Colombia (Magdalena, César, Boyacá, Meta and Putumayo), Venezuela (Bolívar), Guyana (Demerara-Mahaica and Potaro-Siparuni), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Sucumbios, Pichincha, Chimborazo and Pastaza), Peru (Loreto, Cusco and Junin), Bolivia (Beni, Cochabamba and Santa Cruz), Brazil (Pará, Ceará, Pernambuco, Bahia, Mato Grosso, Goiás, Espirito Santo, Minas Gerais, Rio de Janeiro, São Paulo and Santa Catarina), Paraguay (Asunción, Paraguarí, Guairá, Caazapá and Itapúa) and Argentina (Salta, Tucumán and Misiones). Navarrete-Heredia et al. (2002) list P. pygmaeus from some countries above (note: ‘Antillas’) as well as Uruguay and Galápagos Islands (Ecuador), and Newton and Peck (2006) also listed it from the Galápagos Islands (Santa Cruz). In Mexico, Navarrete-Heredia et al. (2002) cite this species also from Sinaloa and Tabasco. Biological notes Piestus pygmaeus has been found under bark, in leaf litter near decaying logs, recently dead or not (‘in dry inner bark of Lecythis’), and on fungi (‘fungus covered log’). Some specimens were collected in flight intercept traps or UV light traps. This species occurs together with P. fronticornis, P. bicornis and P. minutus. Remarks Piestus pygmaeus is similar to P. extimus and P. buquetii but differs mainly by slightly deflected front (Fig. 76), prominent eyes when seen from above, undulate microstriae and conspicuous fine punctures on pronotum, parallel sides of E. Caron et al. pronotum, and very broad apex of lateral lobes of aedeagus (Fig. 190). Piestus extimus and P. buquetii have front suddenly deflected (Figs 77, 78), eyes slightly prominent from above, and subparallel lateral sides of pronotum. Piestus extimus has undulate microstriae and inconspicuous fine punctures equally distributed on pronotum (Fig. 77), and narrower apex of lateral lobes (Fig. 192), while Piestus buquetii has undulate microstriae only on lateral areas on pronotum and conspicuous fine punctures (Fig. 78), and slightly broad apex of lateral lobes (Fig. 194). Piestus extimus Sharp, 1887 (Figs 29, 77, 192, 193, 262) Piestus extimus Sharp, 1887: 713 (original description, type locality: ‘Mexico, Pinos Altos in Chihuahua’); Blackwelder, 1944: 100 (distribution); Newton et al. 2000: 376 (distribution, biological notes); Herman, 2001b: 1791 (distribution); Navarrete-Heredia et al., 2002: 208 (distribution). Piestus (Piestus) extimus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 290 (characters, distribution). Type material Holotype deposited in BMNH, female, with labels: (1) ‘Piestus extimus/ Type D.S./Pinos altos. Chihua/hua. Buch-Hepburn’ [white label, together with the specimen, handwritten]; (2) ‘Holo-/type’ [circle white label with border red, printed in black]; (3) ‘Pinos Altos,/Chihuahua,/Mexico./BuchanHepburn’ [white label, printed in black]; (4) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (5) ‘Holotype/Piestus/extimus/Sharp, 1887/det. R.G. Booth 2008’ [white label, the four first lines handwritten and the last one printed in black]. Note: In the original description Sharp (1887) specified only one specimen observed. Additional material See Appendix 2. Redescription BL: 3.7–5.2 mm, BW: 1.1–1.3 mm. Body flattened dorsoventrally; head, pronotum and abdomen light brown to black and elytra light brown to reddish, always lighter than body (Fig. 29). Dorsal integument of head and pronotum generally with undulate microstriae and inconspicuous fine punctures, sometimes with moderately sized punctures on median longitudinal sulcus (Fig. 77); striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 77); V-shaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae almost reaching apex of abdomen (Fig. 29); scape with strongly enlarged area and long setae on the basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and slightly narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; on each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal Revision of Piestus with remarks on related genera border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with a very weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.41) (Fig. 77); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching basal third of pronotum; abrupt constriction at basal quarter; apical three-quarters with subparallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.98); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 192); lateral lobes exceed a little the apex of median lobe in lateral view; internal sclerites as in Fig. 193. Female. Similar to male, except for: antennae exceed a little apex of elytra; scape with fewer setae; ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 262); spermatheca with basal half suddenly narrower than apical (Fig. 262). Distribution In the current study the species was examined from the United States of America (Arizona) and Mexico (Durango, Nayarit, Jalisco, México, Oaxaca and Guanajuato). In Mexico, Navarrete-Heredia et al. (2002) list P. extimus also from Chihuahua and Sonora. Biological notes Piestus extimus has been found on decaying cacti and sotol (Dasylirion spp.) (Newton et al. 2000), under bark of decaying logs and collected in flight intercept traps. Remarks Piestus extimus is very similar to P. buquetii, from which it is easily separated by undulate microstriae and inconspicuous fine punctures on pronotum, and narrow apex of lateral lobes of aedeagus (in P. buquetii, undulate microstriae only on lateral areas, conspicuous fine punctures equally distributed and slightly broad apex of lateral lobes). Piestus extimus is also similar to P. pygmaeus, from which it differs by characters listed above (see remarks under P. pygmaeus). Piestus buquetii Fauvel, 1864 (Figs 30, 78, 194, 195, 263) Piestus buquetii Fauvel, 1864: 28 (original description, type locality: ‘Cayenne’); Herman, 2001b: 1789 (distribution); Navarrete-Heredia et al., 2002: 208 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Piestus) buqueti: Bernhauer & Schubert, 1910: 7 (error: misspelling and Fauvel 1865: 32, distribution); Blackwelder, 1944: 100 (error: misspelling and Fauvel 1865: 32, distribution); Scheerpeltz, Invertebrate Systematics 525 1952: 291 (error: misspelling and Fauvel 1865: 32, characters, distribution). Type material Holotype deposited in IRSNB, male, with labels: (1) ‘Cayenne’ [white label, handwritten]; (2) ‘Buqueti/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N. B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1864) specified only one specimen observed. Additional material See Appendix 2. Redescription BL: 5.0–5.5 mm, BW: 1.3–1.4 mm. Body flattened dorsoventrally; head, pronotum and abdomen light brown to black and elytra light brown to reddish, always lighter than body (Fig. 30). Dorsal integument of head with undulate microstriae and inconspicuous fines punctures (Fig. 78); pronotum with undulate microstriae only on lateral areas and conspicuous fine punctures equally distributed, some moderately sized punctures on longitudinal sulcus; striae and interstriae of elytra with inconspicuous fine punctures. Metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 78); Vshaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae exceeding apex of abdomen (Fig. 30); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and slightly narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; on each lateral third, antero-internal seta a little shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it in right mandible with a very weak area that resembles a tooth, absent in left mandible. Pronotum wider than long (PW/PL = 1.34) (Fig. 78); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reach basal third of pronotum; abrupt constriction at basal quarter; apical three-quarters with subparallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/ BW = 1.03); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 with parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 194); lateral lobes exceed a little apex of median lobe in lateral view and weakly wide on apex; internal sclerites as in Fig. 195. Female. Similar to male, except for: antennae exceed a little apex of elytra; scape with fewer setae; ovipositor as in P. sulcatus; spermathecal duct ~2 times the length of spermatheca, with 526 Invertebrate Systematics sclerotised basal region (Fig. 263); spermatheca with basal half suddenly narrower than apical (Fig. 263). Distribution In the current study the species was examined from French Guiana (Cayenne), Peru (Cusco) and Bolivia (La Paz and Beni). Navarrete-Heredia et al. (2002) list P. buquetii also from Mexico (Guanajuato), Guyana, Suriname, Brazil, Paraguay and Argentina. However, the distribution of P. buquetii in Mexico is doubtful, as this species is very similar to P. extimus, which occurs in Mexico. Biological notes Piestus buquetii has been found under bark or on fungi on decaying logs (‘fungus covered log’). Remarks Piestus buquetii is very similar to P. extimus, from which it is easily separated by characters of the pronotum and lateral lobes of aedeagus (see remarks under P. extimus). Piestus buquetii is also similar to P. pygmaeus, from which it differs by characters listed above (see remarks under P. pygmaeus). Piestus filicornis Fauvel, 1902 (Figs 31, 79, 125, 196, 197, 264) Piestus filicornis Fauvel, 1902: 23 (original description, type locality: ‘Bolivie: Yuracaris’); Blackwelder, 1944: 100 (distribution, error: Fauvel, 1902: 22); Herman, 2001b: 1791 (distribution). Piestus (Piestus) filicornis: Bernhauer & Schubert, 1910: 7 (distribution, error: Fauvel, 1902: 22); Scheerpeltz, 1952: 287 (characters, distribution, error: Fauvel, 1902: 22). Type material Holotype deposited in IRSNB, male [specimen damaged, left antenna without antennomeres 8–11, right antenna without antennomeres 9–11], with labels: (1) ‘Yuracaris/Bolivie’ [white label, handwritten]; (2) ‘filicornis/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/ Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1902) specified only one specimen observed. Additional material See Appendix 2. Redescription BL: 7.9–8.3 mm, BW: 2.0–2.2 mm. Body flattened dorsoventrally; entirely brown to reddish dark brown (Fig. 31). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures (Fig. 79); striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 79); Vshaped frontal sulcus complete; anterior angles curved and prominent, and transverse carina on base, slightly emarginate at middle; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. E. Caron et al. Antennae reaching apex of abdomen (Fig. 31); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and conspicuously narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; on each lateral third, the antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth developed, not forming a bifurcate apex (Fig. 125), shorter than ventral; internal border asymmetrical, each with one acute tooth at middle (left somewhat deflected) and, anterior to it, in right mandible with a area that resemble a tooth, absent in left mandible; external dorsal margin on basal half carinate. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.48) (Fig. 79); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical three-quarters with subparallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 at the basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 196); lateral lobes exceeding apex of median lobe by onequarter of its length in lateral view, external margin, near the apex, conspicuously curved; internal sclerites as in Fig. 197. Female. Similar to male, except for: antennae exceed a little the apex of elytra; scape with fewer long setae; ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 264); spermathecal capsule with apex somewhat globose and with projection on the base (Fig. 264). Distribution Bolivia (Cochabamba). Biological notes All material of P. filicornis observed in this study (except the holotype) were collected in flight intercept traps. Remarks Piestus filicornis is similar to P. penicillatus and P. minutus but differs by transverse carina slightly emarginate at the middle on vertex, and curved apex of median lobe of aedeagus (Fig. 79) (transverse carina interrupted at the middle and uncurved apex of median lobe in P. penicillatus; Fig. 83; transverse carina continuous at the middle and uncurved apex of median lobe in P. minutus; Fig. 82). Piestus filicornis, as well as P. penicillatus and P. minutus, are also similar to P. abdominalis and P. niger, with which they share the transverse carina on base of vertex. However, these species (Piestus filicornis, P. penicillatus and P. minutus) may be easily separated from them by integument of pronotum with undulate microstriae equally distributed on pronotum (Figs 79, 82, 83) (P. abdominalis and P. niger have only undulate microstriae on Revision of Piestus with remarks on related genera lateral areas; Figs 80, 81). There is also a difference in morphology of the spermatheca, in which the spermatheca has a projection on the base of capsule (P. filicornis, P. penicillatus and P. minutus; Figs 264, 267, 268) or lacks it (P. abdominalis and P. niger; Figs 265, 266). Piestus abdominalis, sp. nov. (Figs 32, 80, 198, 199, 265) Type material Holotype deposited in AMNH, male, labels: (1) ‘Mexico: Chiapas/Union Juarez, n.v. slope/Volcan Tacapa, Rio de/Finca Muxbal (Guat.) ex./pile of nouldy wood/1430m 21 Dec 1976/H. Frania D. Proctor 6’ [white label, printed in black]. 12 Paratypes deposited in following collections: 7 in AMNH, 1 male, 1 female, labels: 1) ‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, on ground/agricultural land, ele./1720 m, 14-26 Dec 1975/115-75’ [white label, printed in black]; (2) ‘H. Frania/Collector’ [white label, printed in black]; 1 female, labels: (1) ‘Mexico, Chiapas/Union Juarez Barranca/Providencia, n.e. slope/Volcan Tacana, shaded/stream, leaf litter/ edge, ele. 1560 m,/16-25 Dec 1975, 119-75’ [white label, printed in black]; ‘H. Frania/Collector’ [white label, printed in black]; 1 female, labels: (1) ‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, (Guat.)/Rio de Finca Muxba, ex./freshly cut stump, much/sap, peeling bark, ele./1430 m, 26 Dec 1975/219-75’ [white label, printed in black]; (2) ‘H. Frania/Collector’ [white label, printed in balck]; 1 female, labels: (1) ‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, (Guat.)/Rio de Finca Muxba, leaf/litter river edge, ele./1430 m, 26 Dec 1975/218-75’ [white label, printed in black]; ‘H. Frania/Collector’ [white label, printed in black]; 1 female, label: (1) ‘Mexico: Chiapas/Union Juarez n.v. slope/Volcan Tacana ex. lear/litter weedy areas under/coffee 1000m 3 Jan 1977/H. Frania 28’ [white label, printed in black]; 1 in SEMC, 1 male, labels: (1) ‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, on ground/agricultural land, ele./1720 m, 14-26 Dec 1975/115-75’ [white label, printed in black]; (2) ‘H. Frania/ Collector’ [white label, printed in black]; 1 in DZUP, male, labels: (1) ‘Mexico, Chiapas/Union Juarez, n.e. slope/Volcan Tacana, (Guat.)/Rio de Finca Muxba, ex./freshly cut stump, much/sap, peeling bark, ele./1430 m, 26 Dec 1975/219-75’ [white label, printed in black]; (2) ‘H. Frania/Collector’ [white label, printed in balck]; 4 in FMNH, 1 sex undetermined, labels: (1) ‘Guatemala:/N.W. Slope V. de/Fuego. V:2:1948/5700’.R.Mitchell’ [white label, printed in black]; (2) ‘on/fungi’ [white label, printed in black]; (3) ‘Piestus/(s. st.) spp./det. A.F. Newton 1987’ [white label, two first lines handwritten, third printed in black]; 1 male, labels: (1) ‘Mun. Yepocapa,/ Chinal, Guat./V-2:48’ [white label, printed in black, except ‘V-2’ manuscript]; (2) ‘La Jolla Grande/(Finca Monserrat)/NW slope Volcan/ Fuego El.5700ft.’ [white label, printed in black]; (3) ‘CNHM Guatemala/ Zool. Exped. (1948)/R.L.Wenzel, leg./Lot. No.’ [white label, printed in black]; (4) ‘ex rotting/banana/stalks’ [white label, printed in black]; 1 sex undetermined, labels: the same two first labels; (3) ‘CNHM Guatemala/Zool. Exped. (1948)/R.L.Wenzel, leg./Lot. No. 75’ [white label, printed in black, except ‘75’ manuscript]; 1 sex undetermined, labels: (1) ‘Mun. Yepocapa,/ Chinal, Guat./V-3:48’ [white label, printed in black]; the same second label; (3) ‘CNHM Guatemala/Zool. Exped. (1948)/R.L. Wenzel leg./Lot. No. 82’ [white label, printed in black, except ‘82’]; 4) ‘under/rotting/banana/stalks’ [white label, printed in black]. Description BL: 5.8–6.7 mm, BW: 1.6–1.8 mm. Body flattened dorsoventrally; head, pronotum and elytra dark brown to black and abdomen light brown to reddish, always lighter than body. Sometimes elytra lighter than head and pronotum, but darker than abdomen (Fig. 32). Dorsal integument of head with undulate Invertebrate Systematics 527 microstriae and inconspicuous fine punctures (Fig. 80); pronotum with undulate microstriae only on lateral regions and conspicuous fine punctures equally distributed and some moderate on longitudinal sulcus; striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 80); Vshaped frontal sulcus complete; anterior angles curved and prominent, and transverse carina on base, continuous at middle; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 32); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and conspicuously narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth developed, not forming a bifurcate apex, shorter than ventral; internal border asymmetrical, each with one acute tooth at middle and, anterior to it, in right mandible with a area that resemble a tooth, absent in left mandible; external dorsal margin on basal half slightly carinate. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.41) (Fig. 80); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles reach basal third of pronotum; abrupt constriction at basal quarter; apical three-quarters with subparallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.97); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 198); lateral lobes exceed a little apex of median lobe in lateral view, and apex somewhat truncate on ventral margin; internal sclerites as in Fig. 199. Female. Similar to male, except for: antennae exceed a little apex of elytra; scape with fewer long setae; ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 265); spermatheca with basal half suddenly narrower than apical (Fig. 265). Distribution Mexico (Chiapas) and Guatemala (Chimaltenango). Biological notes Piestus abdominalis has been found in litter, on fungi or in association with rotting logs. One specimen was found in ‘leaf litter river edge’. Remarks Piestus abdominalis is very similar to P. niger differing by some moderately sized punctures on longitudinal sulcus of pronotum 528 Invertebrate Systematics (Fig. 80), curved apex of median lobe of aedeagus and slightly truncate apex of lateral lobes on ventral margin (Fig. 198). Piestus niger has only inconspicuous fine punctures on pronotum (Fig. 81), uncurved apex of median lobe and nontruncate apex of lateral lobes (Fig. 200). Piestus abdominalis, as well as P. niger, is also similar to P. filicornis, P. penicillatus and P. minutus, from which they differ by characters discussed above (see remarks under P. filicornis). Etymology The species name refers to the colour of the abdomen, which is always lighter than the rest of the body. Piestus niger Fauvel, 1864 (Figs 33, 81, 200, 201, 266) Piestus niger Fauvel, 1864: 24 (original description, type locality: ‘Mexico’); Sharp, 1887: 712 (distribution); Blackwelder, 1944: 100 (distribution, error: Fauvel, 1865: 28); Herman, 2001b: 1792 (distribution); Navarrete-Heredia et al., 2002: 209 (distribution). Piestus (Piestus) niger: Bernhauer & Schubert, 1910: 7 (distribution, error: Fauvel, 1865: 28); Scheerpeltz, 1952: 289 (characters, distribution, error: Fauvel, 1865: 28). Type material Piestus niger Fauvel, 1864. Lectotype (here designated) deposited in IRSNB, male, with labels: (1) ‘Cordova/Mexique’ [white label, handwritten]; (2) ‘niger/Fvl.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/Piestus/ Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in red]. Two paralectotypes deposited in IRSNB, 1 female with the same labels of lectotype, and 1 male, with the same labels of lectotype, except the second label, absent. Note: In the original description Fauvel (1864) did not specify how many specimens he observed. We consider all material from IRSNB with type label ‘Ex-Typis’ as type material. Additional material E. Caron et al. narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth developed, not forming a bifurcate apex, shorter than ventral; internal border asymmetrical, each with acute tooth at middle and, anterior to it, in right mandible with a area that resembles a tooth, absent in left mandible; external dorsal margin on basal half slightly carinate. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.43) (Fig. 81); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles reach basal third of pronotum; abrupt constriction at basal quarter; apical three-quarters with subparallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.99); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 200); lateral lobes exceed a little apex of median lobe in lateral view; internal sclerites as in Fig. 201. Female. Similar to male, except for: antennae exceed a little the apex of elytra; scape with fewer long setae; ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca, with sclerotised basal region (Fig. 266); spermatheca with basal half suddenly narrower than apical half (Fig. 266). Distribution In the current study, the species was examined from Mexico (Veracruz), Guatemala (Guatemala), Costa Rica (Guanacaste) and Panama. Navarrete-Heredia et al. (2002) list this species also from Honduras and Nicaragua. In Mexico, Navarrete-Heredia et al. (2002) cite this species in Oaxaca, Puebla and Veracruz. See Appendix 2. Biological notes Redescription The only information noted for Piestus niger is occurring under bark of decaying logs. BL: 5.0–6.0 mm, BW: 1.4–1.6 mm. Body flattened dorsoventrally; entirely dark brown to black (Fig. 33). Dorsal integument of head with undulate microstriae and inconspicuous fine punctures (Fig. 81); pronotum with undulate microstriae only on lateral regions and inconspicuous fine punctures equally distributed; striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 81); Vshaped frontal sulcus complete; anterior angles curved and prominent, and transverse carina on base, continuous at middle; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 33); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and conspicuously Remarks Piestus niger is very similar to P. abdominalis but differs by characters cited under remarks of P. abdominalis. It is also similar to P. filicornis, P. penicillatus and P. minutus (see discussion under remarks of P. filicornis). Piestus minutus Erichson, 1840 (Figs 34, 82, 202, 203, 267) Piestus minutus Erichson, 1840: 834 (original description, type locality: ‘Columbia’); Fauvel, 1864: 27 (characters, distribution); Sharp, 1876: 407 (note, distribution); Sharp, 1887: 713 (note, distribution); Fauvel, 1901: 71 (distribution); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1792 (distribution); Navarrete-Heredia et al., 2002: 208 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Piestus) minutus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 291 (characters, distribution). Revision of Piestus with remarks on related genera Piestus sulcatus: Laporte, 1835: 129. - As junior synonym of P. minutus: Erichson, 1840: 834; Fauvel, 1864: 27; Fauvel, 1901: 71; Bernhauer & Schubert, 1910: 7 (error: Laporte, 1834: 129); Blackwelder, 1944: 100 (error: Laporte, 1834: 129); Navarrete-Heredia et al., 2002: 209; Newton et al., 2005: 69. Note: Misidentification, Laporte did not describe the species as new, he attributed it to Gravenhorst, 1806 (Herman 2001b). Piestus nigrator Fauvel, 1902: 24 (original description, type locality: ‘Amazones: Pebas’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1792 (distribution). New synonym. Piestus (Piestus) nigrator: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 291 (characters, distribution). Type material Piestus minutus Erichson, 1840. Lectotype (here designated) deposited in ZMHB, male, with labels: (1) ‘6810’ [white label/printed in black]; (2) ‘minutus/Er./Columb. Mor.’ [green label, handwritten]; 3) ‘SYNTYPUS/ Piestus/minutus Erichson, 1840/labelled by MNHUB 2006’ [red label, printed in black]. Four paralectotypes deposited in ZMHB, sex undetermined, with labels: (1) ‘Hist.-Coll. (Coleoptera)/Nr. 6810/Piestus minutus/Erichson, 1840/Columb. Moritz/Zool. Mus. Berlin’ [green label/ printed in black]; (2) ‘SYNTYPUS/Piestus/minutus Erichson, 1840/labelled by MNHUB 2006’ [red label, printed in black]. Note: In the original description Erichson (1840) did not specify how many specimens he observed. We consider all material from ZMHB with type label ‘SYNTYPUS/Piestus/minutus. . .’ as type material. Piestus nigrator Fauvel, 1902. Lectotype (here designated) deposited in IRSNB, 1 female, with labels: (1) ‘Pebas’ [white label, handwritten]; (2) ‘nigrator/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/ Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (4) ‘Ex-Typis’ [white label, printed in red]. Three paralectotypes deposited in IRSNB, 2 males and 1 female, with labels: (1) ‘Pebas’ [white label, handwritten]; (2) ‘Coll. et det. A. Fauvel/ Piestus/nigrator Fauv./R.I.Sc.N.B. 17.479’ [white label, first and fourth lines printed in black, second and third lines handwritten]; (3) ‘Ex-Typis’ [white label, printed in red]. Note: In the original description Fauvel (1902) did not specify how many specimens he observed. We consider all material from IRSNB with type label ‘Ex-Typis’ as type material. We designated a female as lectotype because it is the only specimen with presumably Fauvel’s handwritten label. Additional material See Appendix 2. Redescription BL: 4.2–5.0 mm, BW: 1.2–1.3 mm. Body flattened dorsoventrally; light brown to reddish dark brown, sometimes with elytra reddish and apical margin darker (Fig. 34). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures (Fig. 82); striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 82); V-shaped frontal sulcus complete, sometimes inconspicuous at middle; anterior angles curved and prominent, and a transverse carina on base; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae almost reaching apex of abdomen (Fig. 34); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than Invertebrate Systematics 529 antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and conspicuously narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth developed, not forming a bifurcate apex, shorter than ventral; internal border slightly asymmetrical, each with one acute tooth at middle and, anterior to it, in right mandible with area that resemble a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wider as long. Pronotum wider than long (PW/PL = 1.39) (Fig. 82); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical three-quarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 at the basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 202); lateral lobes exceeding apex of median lobe by one-third of its length in lateral view; internal sclerites as in Fig. 203. Female. Similar to male, except for: antennae reaching apex of elytra; scape with fewer long setae; ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 267); spermathecal capsule with basal half suddenly narrower than apical and with projection on base (Fig. 267). Distribution In the current study, this species was examined from Mexico (Nayarit), Belize (Orange Walk), Guatemala (Alta Verapaz, Izabal and Escuintla), El Salvador (Usulatán), Honduras (Atlántida), Nicaragua (Granada and Río San Juan), Costa Rica (Guanacaste, Heredia, San José, Cartago and Puntarenas), Panama (Chiriquí, Veraguas, Colón, Panamá and Darién), Dominica, Grenada, Trinidad and Tobago (Trinidad), Colombia (Magdalena and Amazonas), Venezuela (Aragua), Guyana, French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Sucumbios, Pichincha, Napo and Pastaza), Peru (Cusco and Madre de Dios), Bolivia (Beni, Cochabamba and Santa Cruz), Brazil (Amazonas, Pará, Rondônia, Goiás, São Paulo, Santa Catarina and Rio Grande do Sul), Paraguay (Guairá) and Argentina (Tucumán). In addition, Navarrete-Heredia et al. (2002) listed this species from Mexico (Tabasco) and Cuba and Newton and Peck (2006) listed it from the Galápagos Islands (Santa Cruz). Biological notes Piestus minutus has been found under bark, in leaf litter near decaying logs, on fermenting fruits (‘berlese in fruit fall’) and on fungi (‘fungusy log’). Some specimens were collected in flight intercept or Malaise traps. This species also has been noticed occurring together with P. fronticornis, P. bicornis and P. pygmaeus. 530 Invertebrate Systematics Remarks This species is very similar to P. penicillatus but differs mainly by transverse carina on base of vertex of head continuous at the middle (Fig. 82), and long lateral lobes of aedeagus (Fig. 202). Piestus penicillatus has transverse carina interrupted at the middle (Fig. 35), and short lateral lobes (Fig. 204). Piestus minutus, as well as P. penicillatus, may be confused with P. filicornis because they have similar integument of pronotum with undulate microstriae and inconspicuous fine punctures. However, they are easily separated by characters cited under P. filicornis. Piestus minutus is also similar to P. abdominalis and P. niger, from which it differs by characters discussed under P. filicornis. Piestus penicillatus (Dalman, 1821) (Figs 35, 83, 204, 205, 268) Zirophorus penicillatus Dalman, 1821: 373 (original description, type locality: ‘insula Guadeloupe’). Piestus paenicillatus: Laporte, 1835: 129 (misspelling, as junior synonym of P. sulcatus); Blackwelder, 1943: 46 (mention, error: Laporte, 1834: 129). Piestus penicillatus: Erichson, 1840: 834 (characters, distribution); Fauvel, 1864: 23 (characters, distribution); Blackwelder, 1943: 46 (characters, distribution, notes); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1793: 373 (distribution); NavarreteHeredia et al., 2002: 209 (distribution). Piestus (Piestus) penicillatus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 288 (characters, distribution). Piestus erythropus Erichson, 1840: 834 (original description, type locality: ‘Cuba’); Fauvel, 1864: 25 (characters, distribution); Sharp, 1887: 713 (note: discussion about distribution); Fauvel, 1902: 26 (note: discussion about distribution); Herman, 2001b: 1790 (distribution). syn. rest. (restored synonym). Piestus (Piestus) erythropus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 290 (characters, distribution). - As synonym of P. penicillatus: Blackwelder, 1943: 46; Blackwelder, 1944: 100; Navarrete-Heredia et al., 2002: 209 (error: Dalman, 1822). Trichocoryne striata Gray, 1832: 306, Fig. 5 (original description, type locality: ‘West Indies’). - As junior synonym of P. erythropus: Erichson, 1840: 834; Fauvel, 1864: 25; Bernhauer & Schubert, 1910: 7; Herman, 2001b: 1790; Navarrete-Heredia et al., 2002: 209. - As junior synonym of P. penicillatus: Blackwelder, 1943: 46; Blackwelder, 1944: 100. Note: Name preoccupied by Zirophorus striatus Guérin-Méneville, 1829 (see P. bicornis above). Type material Zirophorus penicillatus Dalman, 1821. Syntype deposited in NHRS, male, with labels: (1) ‘Guadeloupe/Forsström.’ [white label, printed in black]. Note: In the original description Dalman (1821) did not specify how many specimens he observed. We received from NHRS one specimen, regarded as a syntype. Piestus erythropus Erichson, 1840. Syntype deposited in ZMHB, male, with labels: (1) ‘6807’ [white label, printed in black]; (2) ‘erythropus/Er.?/ Cuba Aiehr?’ [green label, handwritten]; (3) ‘SYNTYPUS/Piestus/ erythropus Erichson, 1840/labelled by MNHUB 2006’ [red label, printed in black]. Note: In the original description Erichson (1840) did not specify E. Caron et al. how many specimens he observed. We received from ZMHB one specimen, regarded as a syntype. Trichocoryne striata Gray, 1832. Not found. Blackwelder (1943) cited the type as ‘unknown’. Additional material See Appendix 2. Redescription BL: 4.6–6.7 mm, BW: 1.1–1.5 mm. Body flattened dorsoventrally; light brown to reddish dark brown (Fig. 35). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures (Fig. 83); striae and interstriae of elytra with inconspicuous fine punctures; metaventrite, on lateral regions, and abdominal tergites, on anterolateral regions, with moderately sized punctures. Male. Head with front suddenly deflected (Fig. 83); Vshaped frontal sulcus complete; anterior angles curved and prominent, and transverse carina on base, interrupted at middle; eyes with line of three moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 35); scape with strongly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape and conspicuously narrowing from antennomere 4 to 11. Labrum with six setae medially, equal in length; on each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth developed, not forming a bifurcate apex, shorter than ventral; internal border asymmetrical, each with one acute tooth at middle and, anterior to it, in right mandible with a area that resemble a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/ PL = 1.31) (Fig. 83); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles slightly developed and reaching basal quarter of pronotum; abrupt constriction at basal quarter; apical threequarters with parallel sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.07); stria 6 at basal half, sometimes inconspicuous, and stria 7 absent; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view (Fig. 204); lateral lobes exceed a little apex of median lobe in lateral view; internal sclerites as in Fig. 205. Female. Similar to male, except for: antennae exceed a little apex of elytra; scape with fewer long setae; ovipositor as in P. sulcatus; spermathecal duct ~1.5 times length of spermatheca, with sclerotised basal region (Fig. 268); spermathecal capsule with basal half suddenly narrower than apical and with a projection on the base (Fig. 268). Distribution In the current study, the species was examined from Mexico, Cuba (Pina del Río, La Habana, Matanzas, Santiago de Cuba and Revision of Piestus with remarks on related genera Guantánamo), Dominican Republic, Puerto Rico and Guadeloupe. The distribution in Mexico is doubtful (Sharp 1887; Fauvel 1902; Navarrete-Heredia et al. 2002). We observed one old specimen, female, from ‘Sierra de Durango, Mexique’ deposited in IRSNB. Biological notes Piestus penicillatus occurs on silk cotton tree, under bark of logs, in rotten coconut husks, in old banana stalks and in rotten cocoa pods (Blackwelder 1943). Remarks This species is very similar to P. minutus, from which it is easily distinguished by characters cited above (see remarks under P. minutus). Piestus penicillatus, as well as Piestus minutus, may be confused with P. filicornis because they share the integument of pronotum with undulate microstriae and inconspicuous fine punctures. However, they can be separated by characters cited above (see remarks under P. filicornis). Piestus penicillatus is also similar to P. abdominalis and P. niger, which differ by characters discussed above (see remarks under P. filicornis). Piestus termitis, sp. nov. (Figs 36, 84, 206, 207, 269) Type material Holotype deposited in SEMC, male, labels: (1) ‘Ecuador: Napo, mid./Rio Tiputini, Yasumi Res./Stn. 0
40.50 S, 76
240 W/28 July 1999. AKT#105’ [white label, printed in black]; (2) ‘Ex. Arboreal carton/termitarium TE#4/ A. Tishechkin’ [white label, printed in black]. 6 paratypes distributed: 4 in SEMC: 1 female, the same labels of holotype; 1 sex undetermined, the same first label, (2) ‘bar code/SM0023850/KUNHM-ENT’ [white label, printed in black]; 1 female, labels: (1) ‘Ecuador: Sucumbios/Sacha Lodge, 0.5
S,/ 76
.5W, 270m, 27-VIII-10-IX/1994, Hibbs, ex: malaise’ [white label, printed in black]; (2) ‘bar code/SM0023742/KUNHM-ENT’ [white label, printed in black]; 1 male, labels: (1) ‘Ecuador: Sucumbios/Sacha Lodge, 0.5
S,/76
.5W, 270m, 31-X-10-XI/1994, Hibbs, ex: malaise’ [white label, printed in black], ‘bar code/SM0023085/KUNHM-ENT’ [white label, printed in black]; 1 in FMNH: sex undetermined, labels: (1) ‘Peru: Madre de Dios/Cocha Cashu Bio. Stn./Manu National Park, 350m/11
530 45’S, 71
240 24’W/17–19 Oct 2000; R.Brooks/PERU1B00 042/ex: flight intercept trap’ [white label, printed in black], ‘bar code/SM0257917/ KUNHM-ENT’ [white label, printed in black]; 1 in DZUP: male, labels: (1) ‘Ecuador: Sucumbios/Sacha Lodge, 0.5
S,/76
.5W, 270m, 16-29-VIII-/ 1994, Hibbs, ex: malaise’ [white label, printed in black]; (2) ‘bar code/ SM0024457/KUNHM-ENT’ [white label, printed in black]. Description BL: 4.2–4.3 mm, BW: 1.2–1.3 mm. Body somewhat convex; head, pronotum and abdomen black and elytra reddish (Fig. 36). Dorsal integument of head with undulate microstriae and moderately sized punctures, not contiguous (Fig. 84); pronotum with large punctures, sometimes contiguous, there are some areas not punctured and undulate microstriae only on lateral areas; striae of elytra with inconspicuous fine punctures and interstriae with undulate microstriae; abdominal tergites with moderately sized punctures uniformly distributed. Invertebrate Systematics 531 Male. Head with front slightly deflected (Fig. 84); V-shaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with one moderately sized punctures with long setae on basal half of dorsal margin of eyes. Antennae exceed a little apex of elytra (Fig. 36); scape with one long setae at middle of dorsal face; scape subequal to antennomere 2 and 3 combined in length; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the shortest; each lateral third, antero-internal seta shorter than postero-external seta. Epipharynx as visible dorsally with two lobes at apex. Mandibles without dorsal teeth developed; internal border symmetrical, smooth. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.48) (Fig. 84); anterior angles not projected; on basal half a slightly transverse depression; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching the middle of pronotum; abrupt constriction at basal third; and, anterior to it, in each side with a slight emargination; apical two-thirds with sinuous sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.94); stria 6 absent and stria 7 incomplete, only on apical half; striae narrower than interstriae. Metaventrite with conspicuous median depression near apex. Abdominal segments 3 and 4 slightly narrower than 5–7; tergite 9 with long ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 206); lateral lobes exceeding the apex of median lobe by one-quarter of its length in lateral view; internal sclerites as in Fig. 207. Female. Similar to male, except for: ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca, without sclerotised basal region (Fig. 269); spermatheca as in Fig. 269. Distribution Ecuador (Sucumbios and Napo) and Peru (Madre de Dios). Biological notes Piestus termitis has been collected in Malaise and flight intercept traps. It was also cited occurring in a termitarium, but its association with termites needs to be confirmed. Remarks Piestus termitis is very similar to P. surrufus because they share some characters, such as: complete V-shaped frontal sulcus (Figs 84, 85), prominent eyes from above, antenna short (male and female), two lobes at the apex of epipharynx (Fig. 111), symmetrical internal border of mandibles (Fig. 126), incomplete stria 7 on elytra and undulate microstriae on interstriae, and abdominal segment 3 and 4 narrower than 5–7 (Figs 36, 37). However, P. termitis may be easily separated by having one long setae on dorsal margin of the eyes, one long setae on scape (male and female), large punctures on pronotum (Fig. 84), long lateral lobes of aedeagus, exceeding the apex of median lobe (Fig. 206), and short spermathecal duct, about the same length as spermatheca (Fig. 269). 532 Invertebrate Systematics Piestus surrufus has two long setae on dorsal margin of the eyes, slightly enlarged area and long setae on scape of male and only two long setae on female, inconspicuous fine punctures and some moderately sized punctures dispersed on pronotum (Fig. 85), short lateral lobes of aedeagus, reaching the apex of median lobe (Fig. 208), and long spermathecal duct, ~3 times the length of spermatheca (Fig. 270). Etymology The specific name is derived from the Latin word termes (termite), in allusion to the presumed association of this species with termites, as some type specimens were found in an arboreal carton termitarium. Piestus surrufus, sp. nov. (Figs 37, 85, 111, 126, 208, 209, 270) Type material Holotype deposited in SEMC, female, labels: (1) ‘Nicaragua: Río San Juan/ Dept., 8 km SE EI Castillo/Refugio Bartole, 30m, 10
58.60 N/84
20.40 W, 23-31-V-2002/S.Peck 02-09, ex. flight/intercept trap, NIC1P02 002’ [white label, printed in black]; (2) ‘bar code/SM0561263/KUNHM-ENT’ [white label, printed in black]. 5 paratypes deposited in the following collections: in SEMC, 1 male, the same first label of holotype, (2) ‘bar code/SM0561144/ KUNHM-ENT’ [white label, printed in black]; 1 female, the same first label, (2) ‘bar code/SM0561230/KUNHM-ENT’ [white label, printed in black]; 1 male, labels: (1) ‘Nicaragua: Río San Juan Dept./60 km SE San Carlos, Refúgio/Bartola, 100m 10
58.400 N, 84
20.300 W/25-28-V-2002, R.Brooks, Z.Falin,/S.Chatzimanolis ex. flight intercept/trap, NIC1BFC02 112’ [white label, printed in black]; (2) ‘bar code/SM539779/KUNHM-ENT’ [white label, printed in black]; in FMNH, 1 female, labels: (1) ‘Ecuador: Pastaza Prov.;/Ashuara indian village on/Rio Macuma nr. Rio Morona./Leg: B. Malkin VII.11-16.1971’ [white label, printed in black]; (2) ‘Beating/ dry/foliage’ [white label, printed in black]; in ZMHB, 1 sex undetermined, label: (1) ‘Peru/VI’ (ZMHB); in DZUP, 1 female, the same first label of holotype, (2) ‘bar code/SM0561230/KUNHM-ENT’ [white label, printed in black]. Description BL: 4.4–4.8 mm, BW: 1.1–1.2 mm. Body somewhat convex; head, pronotum and abdomen reddish brown to reddish dark brown and elytra reddish with external and apical margins region darker (Fig. 37). Dorsal integument of head and pronotum with undulate microstriae and inconspicuous fine punctures, some moderately sized punctures on pronotum near base of longitudinal median sulcus and on longitudinal sulcus of posterior angles (Fig. 85); striae of elytra with inconspicuous fine punctures and interstriae with undulate microstriae; abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 85); V-shaped frontal sulcus complete; anterior angles curved and weakly prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae exceed a little apex of elytra (Fig. 37); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape subequal to antennomere 2 and 3 combined in length; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the shortest (Fig. 111); on each E. Caron et al. lateral third, antero-internal seta shorter than postero-external seta. Epipharynx as visible dorsally with two lobes at apex (Fig. 111). Mandibles without dorsal teeth (Fig. 126); internal border symmetrical, smooth. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.43) (Fig. 85); anterior angles not projected; on basal half a slightly transverse depression; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with a slight emargination; apical two-thirds with slightly sinuous sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 at the basal half and stria 7 incomplete, only on apical two-thirds; striae narrower than interstriae. Metaventrite with slightly median depression near apex. Abdominal segments 3 and 4 slightly narrower than 5–7; tergite 9 with long ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 208); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 270. Female. Similar to male, except for antennal scape without enlarged area and with two long setae at middle of dorsal face; ovipositor as in P. sulcatus; spermathecal duct long, ~3 times length of spermatheca, without sclerotised basal region (Fig. 270); spermatheca as in Fig. 270. Distribution Nicaragua (Río San Juan), Ecuador (Pastaza) and Peru. Biological notes Piestus surrufus has been collected in flight intercept traps. One specimen from Ecuador was collected by beating dry foliage. Remarks Piestus differs from P. termitis by characters cited under remarks of P. termitis. Etymology The name refers to the colour of the elytra. Piestus formicinus, sp. nov. (Figs 38, 86, 210, 211, 271) Type material Holotype deposited in SEMC, male, labels: (1) ‘Guiana: Region 8/ Iwokrama Forest, 1 km W Kurupukari/Iwokrama Field Stn., 60m/ 4
400 19’N, 58
410 4’W, 21 May 2001, R.Brooks, Z.Falin/GUY1BF01 004/ ex: fogging fungusy logs’ [white label, printed in black]; (2) ‘bar code/ SM0226160/KUNHM-ENT’ [white label, printed in black]. 25 paratypes distributed: 15 SEMC, 3 AMNH, 3 DZUP, 3 FMNH and 1 MZSP, which: 1 female, the same first label as holotype, (2) ‘bar code/SM0266113/KUNHMENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226214/KUNHM-ENT’ [white label, printed in black] (FMNH); 1 sex undetermined, the same first label, (2) ‘bar code/ SM0226229/KUNHM-ENT’ [white label, printed in black] (DZUP); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226230/KUNHM- Revision of Piestus with remarks on related genera Invertebrate Systematics 533 ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226231/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/ SM0226232/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226233/KUNHMENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226216/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226205/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/ SM0226212/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226224/KUNHMENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226223/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226228/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/ SM0226221/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226220/KUNHMENT’ [white label, printed in black] (MZSP); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226219/KUNHM-ENT’ [white label, printed in black] (AMNH); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226218/KUNHM-ENT’ [white label, printed in black] (AMNH); 1 sex undetermined, the same first label, (2) ‘bar code/ SM0226213/KUNHM-ENT’ [white label, printed in black] (FMNH); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226193/ KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226210/KUNHMENT’ [white label, printed in black] (DZUP); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226209/KUNHM-ENT’ [white label, printed in black] (DZUP); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226208/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/ SM0226207/KUNHM-ENT’ [white label, printed in black] (SEMC); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226206/KUNHMENT’ [white label, printed in black] (FMNH); 1 sex undetermined, the same first label, (2) ‘bar code/SM0226226/KUNHM-ENT’ [white label, printed in black] (AMNH). developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it in right mandible with a very weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.35) (Fig. 38); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles gently developed and reaching the basal third of pronotum; abrupt constriction at basal third; apical two-thirds with curved sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.94); stria 6 absent and stria 7 complete; striae narrower than interstriae. Abdominal segments 3–6 parallel sides; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and slightly curved apex in lateral view (Fig. 210); lateral lobes not reaching the apex of median lobe in lateral view; internal sclerites as in Fig. 211. Female. Similar to male, except for: antennae not reaching apex of elytra; scape without enlarged area and with two long setae at middle of the dorsal face; ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca, without sclerotised basal region (Fig. 271); spermatheca as in Fig. 271. Additional material Remarks See Appendix 2. Piestus formicinus is easily separated from the other species of Piestus by incomplete V-shaped frontal sulcus (Fig. 86), slightly prominent eyes from above, slightly enlarged area with long setae on scape only on male, undulate microstriae and moderately sized punctures equally distributed on pronotum. Description BL: 2.6–3.6 mm, BW: 0.8–0.9 mm. Body somewhat convex; light brown to reddish dark brown (Fig. 38). Dorsal integument of head and pronotum with undulate microstriae and moderately sized punctures (Fig. 86); striae and interstriae of elytra with conspicuous fine punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed; dorsal teeth of mandibles weakly developed. Male. Head with front slightly deflected (Fig. 86); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and weakly prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin of eyes, basal the longest. Antennae exceed a little apex of elytra (Fig. 38); scape with slightly enlarged area and long setae on the basal half of exposed dorsal face; scape subequal to antennomere 2 and 3 combined in length; antennomeres 4–11 oblong shape. Labrum with six setae medially, internal the longest; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly Distribution Guyana (Upper Demerara-Berbice). Biological notes Piestus formicinus may be associated with ants of the genus Acromyrmex, or at least with its refuse material (‘ex: Acromyrmex hystrix refuse pile’). The type series was captured by fogging fungus logs. Etymology The specific name is derived from the Latin word formica (ant), in allusion the presumed association of this species with ants, as some specimens were found in refuse piles of the genus Acromyrmex Mayr, 1865. Piestus sulcatus Gravenhorst, 1806 (Figs 5–10, 106–108, 113–115, 131, 134, 139, 149, 153, 212–214, 272) Piestus sulcatus Gravenhorst, 1806: 224 (original description, type locality: ‘Brasilia’); Erichson, 1840: 835 (characters, distribution); Fauvel 1864: 30 (note, distribution); Sharp, 1876: 407 (note, distribution); Blackwelder, 1943: 44 (characters, distribution, notes); Blackwelder, 1944: 101 (distribution); Blackwelder, 1952: 309 (mention); Herman, 2001b: 1795 (distribution). 534 Invertebrate Systematics Piestus (Piestus) sulcatus: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 286 (distribution). Piestus sanctae-catharinae Bernhauer, 1906: 193 (original description, type locality: ‘Brasilia (Santa Catharina)’); Blackwelder, 1944: 101 (distribution). New synonym. Piestus (Piestus) sanctae-catharinae: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 286 (characters, distribution). Piestus sanctaecatharinae: Herman, 2001b: 1794 (distribution). Piestus condei Wendeler, 1955: 187 (original description, type locality: ‘Brasilien, Espirito Santo, Sta. Theresa’); Herman, 2001b: 1790 (distribution). New synonym. Type material Piestus sulcatus Gravenhorst, 1806. Lectotype (here designated) deposited in ZMHB, male, with labels: (1) ‘Para/Sieber/Nr. 6812’ [green label, handwritten]; (2) ‘Typus’ [red label, printed in black]; (3) ‘Zool. Mus./ Berlin’ [light yellow label, printed in black]; (4) ‘SYNTYPUS/Piestus/ sulcatus Gravenhorst, 1806/labelled by MNHUB 2006’ [red label, printed in black]. Three paralectotypes deposited in ZMHB, 2 females, with the same labels of lectotype, and 1 male [specimen damaged, without abdomen], with labels: (1) ‘6810’ [white label, printed in black]; (2) ‘sulcatus/Gr. x/Pará Sieb’ [green label, handwritten]; (3–5) [third to fifth labels are the same of second to fourth of lectotype]. Note: In the original description Gravenhorst (1806) did not specify how many specimens he used for description. We received 4 specimens with type labels ‘Typus’ from ZMHB from which we are considering as type material. Piestus sanctae-catharinae Bernhauer, 1906. Lectotype (here designated) deposited in NMW, male, with labels: (1) ‘S.Catharina/Brasil. Klimsch’ [white label/printed in black]; (2) ‘Piestus Sanctae/Catharinae Bh.’ [white label, handwritten]; (3) ‘Klimsch/donavit’ [white label, handwritten]; (4) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (5) ‘COTYPUS/Piestus/ Sanctae Catharin/Bernhauer.’ [rose label, first line printed in black and the others handwritten]; (6) ‘Sanctae-/catharinae Bh.’ [green label, handwritten]. Two paralectotypes, 1 female deposited in NMW with the same labels of lectotype, except for: (2) ‘Sanctae Catharinae/Bernh./det. Bernhauer’ [white label, two first lines handwritten and third line printed in black], and 1 female deposited in FMNH with the same labels of lectotype, except for: (2) ‘Sanctae Cathari’/nae Bernh./Typus.’ [light yellow label, Bernhaeur’s handwritten]; (3) ‘Chicago NHMus/M.Bernhauer/Collection’ [white label, printed in black]. Note: In the original description Bernhauer (1906) did not specify how many specimens he used for description. We received 3 specimens with type labels from NMW and FMNH: we are considering all three as type material. Piestus condei Wendeler, 1955. Holotype deposited in ZMHB, male, with labels: (1) ‘EspiritoSanto/Sta. Theresa/Jan.29.Conde’ [white label, printed in black]; (2) ‘,’ [white label, printed in black]; (3) ‘Piestus s.str./condei n.sp. Wdlr.’ [white label, Wendeler’s handwritten]; (4) ‘Holotypus’ [red label, printed in black]; (5) ‘Sammlung/Wendeler’ [white label, printed in black]; (6) ‘condei */Wdlr.’ [green label, handwritten]. Note: In the original description Wendeler (1955) specified one female studied; however, the type observed (ZMHB) is male. E. Caron et al. metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 7); V-shaped frontal sulcus incomplete (gently complete in some species); anterior angles curved and weakly prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae almost reaching apex of abdomen (reach in some species) (Fig. 5); scape with slightly enlarged area and long setae on basal half of exposed dorsal face (Fig. 106); scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length (Fig. 108); each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed (Fig. 113); internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it in right mandible with weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined (Fig. 130). Mentum ~1.5 times as wide as long (Fig. 134). Pronotum wider than long (PW/PL = 1.49) (Fig. 7); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with curved sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex (Fig. 8). Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.97) (Fig. 7); stria 6 at basal quarter; stria 7 complete; striae narrower than interstriae; epipleural carina developed (Fig. 6). Metaventrite with slightly median depression near apex (Fig. 9). Abdominal segments 3–6 subparallel sides, segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina (Fig. 10); tergite 9 with short ventral struts (Fig. 149); tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Figs 212, 214); lateral lobes exceeding apex of median lobe by one-quarter of its length in lateral view; internal sclerites as in Fig. 213. Female. Similar to male, except for: antennae reaching apex of elytra; scape without enlarged area and with two long setae at middle of the dorsal face (Fig. 107); prosternum without ovate set of fine punctures; ovipositor with curved external margin (Fig. 153); spermathecal duct ~1.5 times length of spermatheca, without sclerotised basal region (Fig. 272); spermatheca somewhat elongate (Fig. 272). Additional material Distribution See Appendix 2. In the current study, the species was examined from Nicaragua (Rio San Juan), Costa Rica (Guanacaste, Alajuela, Heredia, Cartago, Limón and Puntarenas), Panama (Veraguas, Colón, Panamá and Darién), Guadeloupe, Dominica (Saint David), Martinique, Saint Vincent and the Grenadines (Saint Vincent), Trinidad and Tobago (Trinidad), Colombia (Choco), Venezuela (Aragua), Guyana (Upper Demerara-Berbice), Suriname (Marowijne), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Esmeraldas, Sucumbios, Pichincha, Napo, Cotopaxi and Pastaza), Peru (Loreto, San Martín, Huancavelica, Redescription BL: 3.8–5.8 mm, BW: 1.1–1.6 mm. Body flattened dorsoventrally; light brown to reddish dark brown (Fig. 6). Dorsal integument of head and pronotum with undulate microstriae and fine to moderately sized punctures, not contiguous (Fig. 7); striae of elytra with moderately sized punctures and interstriae with conspicuous fine punctures; Revision of Piestus with remarks on related genera Cusco and Madre de Dios), Bolivia (Cochabamba and Santa Cruz), and Brazil (Amazonas, Pará, Bahia, Goiás, Distrito Federal, Rio de Janeiro, São Paulo, Paraná and Santa Catarina). Herman (2001b) listed P. sanctaecatharinae (junior synonym of P. sulcatus) from Paraguay too; Newton and Peck (2006) also report P. sulcatus from the Galápagos Islands (Isabela and Santa Cruz). Biological notes Piestus sulcatus has been found in leaf litter, fruit falls or under bark of decaying logs. Six specimens from Panamá were observed in ‘7 day old tapir dung’. This species also has been collected in flight intercept and Malaise traps, Berlese extraction of leaf litter, and at light trap. Blackwelder (1943) noted some similar habitats. Remarks Piestus sulcatus is similar to P. gounellei and P. mexicanus differing from those species by incomplete V-shaped frontal sulcus (gently complete in some specimens) (Fig. 7), undulate microstriae and moderately sized punctures on head and pronotum, developed stria 6 on elytra (Fig. 5), lateral lobes of aedeagus exceeding the apex of median lobe by one-quarter of its length (Fig. 212), and spermatheca elongate (Fig. 272). Piestus gounellei has a complete V-shaped frontal sulcus (Fig. 87), moderate to large punctures on head and pronotum, undulate microstriae only on lateral areas, developed stria 6, short lateral lobes exceeding a little the apex of median lobe (Fig. 215), and globose apex of spermatheca (Fig. 273). Piestus mexicanus has a complete V-shaped frontal sulcus (Fig. 88), large punctures on head and pronotum, undulate microstriae only on lateral areas, no stria 6, short lateral lobes exceeding a little the apex of median lobe (Fig. 218), and somewhat elongate spermatheca (Fig. 274). Piestus gounellei Fauvel, 1902 (Figs 39, 87, 215–217, 273) Piestus gounellei Fauvel, 1902: 24 (original description, type locality: ‘Brésil: Minas Geraes, Caraça’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1791 (distribution). Piestus (Piestus) gounellei: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1933: 994 (mention); Scheerpeltz, 1952: 285 (characters, distribution). Piestus wasmanni Fauvel, 1902: 25 (original description, type locality: ‘Rio de Janeiro, Colonia Alpina’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1795 (distribution). New synonym. Piestus (Piestus) wasmanni: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 285 (characters, distribution). Type material Piestus gounellei Fauvel, 1902. Holotype deposited in IRSNB, male, with labels: (1) ‘Caraça(Minas Geraes)/Brésil/E.Gounelle 1.2.1885’ [white label, printed in black]; (2) ‘gounellei/Fvl.’ [handwritten]; (3) ‘R.I.Sc.N.B. 17.479 [printed in black]/Piestus [handwritten]/Coll. et det. A. Fauvel [printed in black]’; (4) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1902) specified only one specimen studied. Piestus wasmanni Fauvel, 1902. Holotype deposited in IRSNB, female, with labels: (1) ‘Col. Alpina/Rio de Jan.’ [white label, Invertebrate Systematics 535 handwritten]; (2) ‘Wasmanni/FvL.’ [white label, handwritten]; (3) ‘R.I.Sc. N.B. 17.479/Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (4) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1902) specified only one specimen studied. Additional material See Appendix 2. Redescription BL: 4.4–5.9 mm, BW: 1.2–1.6 mm. Body flattened dorsoventrally; reddish brown to reddish dark brown (Fig. 39). Dorsal integument of head with moderately sized punctures, not contiguous (Fig. 87); pronotum with moderate to large punctures, sometimes contiguous, there are some areas not punctured and, undulate microstriae only on lateral areas; striae of elytra with moderately sized punctures and interstriae with inconspicuous fine punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 87); V-shaped frontal sulcus complete; anterior angles curved and prominent, slightly median fovea at base; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 39); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it in right mandible with weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 as wide as long. Pronotum wider than long (PW/PL = 1.41) (Fig. 87); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching the middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with a short tooth and slight emargination; apical two-thirds with curved sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/ BW = 1.04); stria 6 at basal half and stria 7 complete; striae narrower than interstriae; epipleural carina developed. Metaventrite with slightly median depression near apex. Abdominal segments 3–6 subparallel sides, segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view, curved apex in lateral view, and projected apex in ventral view (Figs 215, 217); lateral lobes exceeding a little the apex of median lobe in lateral view; internal sclerites as in Fig. 273. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape without enlarged area and with some long setae on 536 Invertebrate Systematics basal half of exposed dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca, without sclerotised basal region (Fig. 273); spermatheca globose at apex (Fig. 273). Distribution In the current study, the species was examined from Brazil (Espirito Santo, Rio de Janeiro, São Paulo, Paraná and Santa Catarina), Paraguay (Caazapá and Itapúa) and Argentina (Misiones). Biological notes Piestus gounellei has been found in litter or on or under bark of decaying logs. This species also has been collected in flight intercept traps. Remarks This species is similar to P. sulcatus and P. mexicanus, but differs by characters cited above (see remarks under P. sulcatus). Piestus mexicanus Laporte, 1835 (Figs 40, 88, 218, 219, 274) Piestus mexicanus Laporte, 1835: 130 (original description, type locality: ‘Mexique’); Erichson, 1840: 836 (characters, distribution); Fauvel, 1864: 30 (characters, distribution); Sharp, 1887: 714 (characters, distribution); Blackwelder, 1944: 100 (distribution, error: Laporte, 1834: 30); Herman, 2001b: 1792 (distribution); Navarrete-Heredia et al., 2002: 208 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Piestus) mexicanus: Bernhauer & Schubert, 1910: 7 (distribution, error: Laporte, 1834: 30); Scheerpeltz, 1952: 287 (distribution, error: Laporte, 1834: 30). Piestus alternans Sharp, 1887: 714 (original description, type locality: ‘Panama, David in Chiriqui’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1788 (distribution). New synonym. Piestus (Piestus) alternans: Bernhauer & Schubert, 1910: 6 (distribution); Scheerpeltz, 1952: 287 (distribution). Type material Piestus mexicanus Laporte, 1835. Syntype deposited in BMNH, female [damaged apex of antennae and abdomen], with labels: (1) ‘Piestus mexica-/ nus Type ex./coll. Chevrolat./Mexico. Lesneut.?’ [white label, together with the specimen, handwritten]; (2) ‘Type’ [circle white label with border red, printed in black]; (3) ‘Mexico.’ [white label, printed in black]; (4) ‘Mexico’ [white label, handwritten]; (5) ‘B.C.A. Col. I. 2./Piestus/mexicanus,/Lap.’ [white label, printed in black]; (6) ‘27.?’ [light red label, handwritten]; (7) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (8) ‘Piestus/ mexicanus Lap. Er./836 11.? type tolp?/Mexico Oaxaca? D. Lesneur.’ [green label, handwritten, note: ‘Colombia’ handwritten underside]. Note: In the original description Laporte (1835) did not specify how many specimens he observed. We received from BMNH only one specimen labelled as type. Piestus alternans Sharp, 1887. Holotype deposited in BMNH, male [broken right antenna], with labels: (1) ‘Piestus alternans/ Type D.S./David Panama/Champion’ [white label, together with the specimen, handwritten]; (2) ‘Holo-/type’ [circle white label with border red, printed in black]; (3) ‘David,/Panama./Champion.’ [white label, printed in black]; (4) ‘B.C.A. Col. i. 2./Piestus/alternans, Sharp.’ [white label, printed in black]; (5) ‘Sharp Coll./ 1905-313.’ [white label, printed in black]; (6) ‘Holotype/Piestus/alternans/ Sharp, 1876/det. R.G. Booth 2008’ [white label, the four first lines E. Caron et al. handwritten and the last one printed in black]. Note: In the original description Sharp (1887) specified only one specimen (male) observed. Additional material See Appendix 2. Redescription BL: 3.8–5.4 mm, BW: 1.1–1.5 mm. Body flattened dorsoventrally; light brown to reddish dark brown (Fig. 40). Dorsal integument of head and pronotum with large punctures (Fig. 88), sometimes contiguous, there are some areas not punctured and, undulate microstriae only on lateral areas; striae of elytra with large punctures and interstriae with inconspicuous fine punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 88); V-shaped frontal sulcus complete; anterior angles curved and prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 40); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–10 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than posteroexternal seta. Mandibles with dorsal teeth weakly developed; internal borders slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 as wide as long. Pronotum wider than long (PW/PL = 1.54) (Fig. 88); anterior angles weakly projected; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with curved sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 absent and stria 7 complete; striae narrower than interstriae; epipleural carina developed. Metaventrite with slight median depression near apex. Abdominal segments 3–6 subparallel sides, segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 218); lateral lobes exceeding little apex of median in lateral view; internal sclerites as in Fig. 219. Female. Similar to male, except for: antennae reaching apex of elytra; scape without enlarged area and with two long setae at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca (or ~2 times, specimens from Nicaragua, Costa Rica and Panama), without sclerotised basal region (Fig. 274); spermatheca somewhat elongate (Fig. 274). Revision of Piestus with remarks on related genera Distribution In the current study, the species was examined from Mexico (Tamaulipas, Durango, San Luís Potosí, Hidalgo, Veracruz, Oaxaca and Chiapas), Belize (Cayo), Guatemala (Zacapa and Escuintla), Nicaragua (Granada), Costa Rica (Guanacaste, Alajuela, Heredia, San José, Cartago and Puntarenas), Panama (Chiriquí and Panamá) and Colombia (Boyacá). Herman (2001b) cites this species from some of the countries listed above, plus Honduras, Brazil and the West Indies. Navarrete-Heredia et al. (2002) consider Brazil and West Indies as doubtful distributions. Biological notes Piestus mexicanus has been found in leaf litter near streams and under bark of decaying logs. Some species have been observed in ‘fruit fall’, ‘flower fall’ or ‘mushrooms rotting vegetation’. The species was also collected in flight intercept traps. Remarks Piestus mexicanus is similar to P. sulcatus and P. gounellei, but differs by characters cited under remarks of P. sulcatus. Piestus ecuadorensis, sp. nov. (Figs 41, 89, 220, 221, 275) Type material Holotype deposited in AMNH, male, label: (1) ‘Ecuador: Pichincha Prov.:/W of Alluriquin, Tinalandia,/2600-2800 ft, May 19-20/1993, Lee Herman #2724-/2725; litter nr. stream [white label, printed in black]. 6 paratypes deposited in following collections: in AMNH: 1 female and 1 sex undetermined, the same label of holotype; 1 female, labels: (1) ‘Ecuador: Pichincha Prov.:/W of Alluriquin, 3.3–5.3 km/SW road to Cooperativa/ Bolivar, near Tinalandia,’ [white label, printed in black]; (2) ‘3100-3500, V-20-93; L. Herman/#2733-2734, litter nr stream’ [white label, printed in black]; in DZUP: 1 sex undetermined, label: (1) ‘Ecuador: Pichincha: 17/ km SE Sto. Domingo de/Colorados, Tinalandia/3000’X-16-21-88/litter, leg. L. Herman’ [white label, printed in black]; in SEMC: 1 female, labels: (1) ‘Ecuador: Pichincha/Tinalandia, Santo Domingo/16 km E, 750m/0
160 53’S, 79
30 39’W/26–27 Mar 1999, R. Brooks, D. Brzoska/ECU1B99 048 ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0158037/ KUNHM-ENT’ [white label, printed in black]; 1 male, the first same label, (2) ‘bar code/SM0158147/KUNHM-ENT’ [white label, printed in black]. Invertebrate Systematics 537 face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.49) (Fig. 89); anterior angles weakly projected; longitudinal median sulcus conspicuous, broad and deep; longitudinal sulcus of posterior angles strongly developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with curved sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.00); stria 6 absent and stria 7 complete; striae narrower than interstriae; epipleural carina developed. Metaventrite with slightly median depression near apex. Abdominal segments 3–6 subparallel sides, segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with one pair of long setae on apex, sometimes with one inconspicuous seta near long setae. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 220); lateral lobes exceed little apex of median lobe in lateral view; internal sclerites as in Fig. 221. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape without enlarged area and with two long setae at middle of the dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca, without sclerotised basal region (Fig. 275); spermatheca somewhat elongate (Fig. 275). Distribution Ecuador (Pichincha). Biological notes Piestus ecuadorensis has been collected in flight intercept traps and found in leaf litter near streams. Description BL: 4.0–4.6 mm, BW: 1.2–1.3 mm. Body flattened dorsoventrally; reddish dark brown (Fig. 41). Dorsal integument of dorsum of head, except anterior angles, with large punctures contiguous (Fig. 89); almost all pronotum with large punctures contiguous, there are few some areas not punctured; striae of elytra with large punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 89); V-shaped frontal sulcus complete; anterior angles curved and prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 41); scape with slightly enlarged area and long setae on basal half of exposed dorsal Remarks Piestus mexicanus is very similar to P. foveolatus because they share contiguous large punctures on the head and a broad and deep longitudinal sulcus on the pronotum (Figs 89, 90). However, P. ecuadorensis differs from P. foveolatus by the absence of a median longitudinal sulcus plus fovea at the base on vertex (Fig. 89), short lateral lobes exceeding slightly the apex of median lobe (Fig. 220), and short spermathecal duct of the same length as spermatheca (Fig. 275). Piestus foveolatus has a median longitudinal sulcus with a fovea at the base of vertex (Fig. 90), long lateral lobes exceeding by one-quarter of its length the apex of median lobe (Fig. 222), and long spermathecal duct, ~3 times the length of spermatheca (Fig. 276). 538 Invertebrate Systematics Piestus ecuadorensis, as well as P. foveolatus, is easily separated from P. gounellei and P. mexicanus by contiguous large punctures on the head and a broad and deep longitudinal sulcus on the pronotum (Figs 89, 90) (non-contiguous punctures on head and not broad and slightly deep longitudinal sulcus on pronotum in P. gounellei and P. mexicanus; Figs 87, 88). Etymology The name is derived from Ecuador, the country where the type material was collected. Piestus foveolatus, sp. nov. (Figs 42, 90, 222, 223, 276) Type material Holotype deposited in FMNH, male, labels: (1) ‘Panamá: Bocas d. Toro,/ Fortuna-Chiriquí Grande/road, 8
470 N, 82
110 W,/800m, 14–16.vii.1987’ [white labels, printed in black]; (2) ‘D.M. Olson #566,/premontane rain forest,/sifting litter/Field Museum N.H.’ [white label, printed in black]; (3) ‘David M. Olson,/Staphylinidae: species#82’ [white label, printed in black, except ‘82’]. 40 paratypes: in FMNH, 2 sex undetermined, the same label of holotype (FMNH), one of them with additional label: (4) ‘Piestus/ (Trachypiestus)/det. Newton 1993’ [white label, the two first lines manuscript, third printed in black]; in AMNH, 1 female, 10 sex undetermined, labels: (1) ‘Panamá: Chiriquí Prov./4 km NW Volcán, 1450m/8
490 19’N, 82
400 31’W/XII-24–01, L. Herman/litter near stream’ [white label, printed in black]; in SEMC, 1 sex undetermined, labels: (1) ‘Costa Rica: San Jose/Cerros de Escazu/2 km, S. San Antonio, 1650m/ 9
530 30’N, 84
90 0’W/13 Jun 1997, R.Anderson/CR1A97 013E/ex: berlese forest litter’ [white label, printed in black]; (2) ‘bar code/SM0119958/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119965/KUNHM-ENT’ [white label, printed in black]; 1 male, the same first label, (2) ‘bar code/SM0119943/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119960/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/ SM0119939/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119937/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139469/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119842/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119845/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/ SM0119833/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119836/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119837/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139464/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139443/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/ SM0139444/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139446/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139448/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139467/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139458/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/ SM0139453/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139452/KUNHM- E. Caron et al. ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139451/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139450/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0139468/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, labels, (1) ‘Costa Rica: San Jose/ Cerros de Escazu/2 km, S. San Antonio, 1650m/9
530 30’N, 84
90 0’W/13 Jun 1997, R.Anderson/CR1A97 013H/ex: berlese forest litter’ [white label, printed in black]; (2) ‘bar code/SM0136291/KUNHM-ENT’ [white label, printed in black]; in DZUP, 1 sex undetermined, labels: (1) ‘Costa Rica: San Jose/Cerros de Escazu/2 km, S. San Antonio, 1650m/9
530 30’N, 84
90 0’W/13 Jun 1997, R.Anderson/CR1A97 013E/ex: berlese forest litter’ [white label, printed in black], (2) ‘bar code/SM0119926/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0119931/KUNHM-ENT’ [white label, printed in black]. Additional material See Appendix 2. Description BL: 3.4–5.3 mm, BW: 1.1–1.4 mm. Body flattened dorsoventrally convex; reddish dark brown (Fig. 42). Dorsal integument of head, except anterior angles, with large punctures contiguous (Fig. 90); almost all pronotum with large punctures contiguous, there are few some areas not punctured; striae of elytra with large punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 90); V-shaped frontal sulcus complete; anterior angles curved and prominent, and median longitudinal sulcus with fovea at base; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae reaching apex of abdomen (Fig. 42); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with a very weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.50) (Fig. 90); anterior angles not projected; longitudinal median sulcus conspicuous, broad and deep; longitudinal sulcus of posterior angles strongly developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with curved sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 1.04); stria 6 absent and stria 7 complete; striae narrower than interstriae; interstriae not carinate; epipleural carina developed. Metaventrite with conspicuous median depression near apex. Abdominal segments 3–6 subparallel sides, segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 Revision of Piestus with remarks on related genera with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 222); lateral lobes exceeding apex of median lobe by one-quarter of its length in lateral view; internal sclerites as in Fig. 223. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape without enlarged area and with two long setae at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct long, ~3 times length of spermatheca, without sclerotised basal region (Fig. 276); spermatheca as in Fig. 276. Distribution Costa Rica (Alajuela, Heredia, San José, Cartago and Puntarenas), Panama (Bocas del Toro and Chiriquí) and Colombia (Valle del Cauca). Biological notes Piestus foveolatus has been found in leaf litter, sometimes near streams (‘litter near stream’). This species was also collected in flight intercept traps and by Berlese extraction of leaf litter. Remarks Piestus foveolatus is very similar to P. ecuadorensis, and also to P. gounellei and P. mexicanus, but differs by characters cited under remarks of P. ecuadorensis. Etymology The specific name refers to the conspicuous fovea on the median base of head. Piestus sulcipennis Scheerpeltz, 1952 (Figs 43, 91, 224, 225, 277) Piestus sulcipennis Scheerpeltz, 1952: 299 (original description, type locality: ‘St. Catharina-Brasilien’); Herman, 2001b: 1795 (distribution). Type material Piestus sulcipennis Scheerpeltz, 1952. Lectotype (here designated) deposited in NMW, male [damaged, left antenna incomplete], with labels: (1) ‘St.-Catharina/Brasilien’ [white label, printed in black]; (2) ‘ex coll./ Scheerpeltz’ [blue label, printed in black]; (3) ‘Typus/Piestus (Trachy-/ piestus) sulci-/pennis/O. Scheerpeltz [printed in black]’ [red label, first and fifth lines printed in black, second, third and fourth lines handwritten]; (4) ‘sulcipennis/Scheerp.’ [green label, handwritten]. Paralectotype deposited in NMW, sex undetermined, with the same labels of lectotype. Note: In the original description Scheerpeltz (1952) specified a male and female observed. However, only one specimen was dissected and its sex confirmed (male) (lectotype). The sex of the other specimen (paralectotype) is still undetermined. Invertebrate Systematics 539 Dorsal integument of head, except anterior angles, and almost all pronotum with large punctures contiguous, there are some few areas unpunctured (Fig. 91); striae of elytra with large punctures; abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 91); V-shaped frontal sulcus complete; anterior angles curved and prominent; eyes with two moderately sized punctures with long setae on basal half of dorsal margin, basal the longest. Antennae exceeding apex of elytra, but not reaching apex of abdomen; scape with two long setae at middle of dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape (Fig. 43). Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal borders slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with weak area that resembles a tooth, absent in the left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider than long (PW/PL = 1.47) (Fig. 91); anterior angles weakly projected; the apical half with slightly somewhat longitudinal median depression and on basal half a transverse depression; longitudinal sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle or pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra as long as wide (EL/BW = 0.94); stria 6 at basal half and stria 7 complete; striae narrower than interstriae; interstriae not carinate; epipleural carina developed. Abdominal segments 3–6 subparallel sides, segments 3 and 4 slightly narrower than 5–7; sternite 3 with conspicuous transverse carina; tergite 9 with long ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 224); lateral lobes exceed a little apex of median lobe in lateral view; internal sclerites as in Fig. 225. Female. Similar to male, except for: prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct about same length of spermatheca, with sclerotised basal region (Fig. 277); spermatheca with globose apex and accessory gland on base (Fig. 277). Distribution Brazil (Santa Catarina). Biological notes Unknown. Additional material Remarks See Appendix 2. Piestus sulcipennis is similar to P. aper, differing from that species by complete V-shaped frontal sulcus (Fig. 91), two long setae on dorsal margin of the eyes, two long setae on scape, slightly asymmetrical internal borders of mandibles with one small acute tooth, stria 6 developed at the basal half Redescription BL: 3.8–4.0 mm, BW: 1.1–1.1 mm (types). Body flattened dorsoventrally; reddish brown to reddish dark brown (Fig. 43). 540 Invertebrate Systematics E. Caron et al. Figs 92–105. Head and pronotum, dorsal view. 92, Piestus imperfectus, sp. nov.; 93, P. costatus; 94, P. chiriquensis; 95, P. nevermanni; 96, P. paradoxus; 97, P. boliviensis, sp. nov.; 98, P. aper; 99, P. angularis; 100, P. acuminatus, sp. nov.; 101, P. rugosus; 102, P. elegans, sp. nov.; 103, Eleusis interrupta, comb. rest.; 104, Hypotelus laevis, comb. nov.; 105, Hypotelus andinus, comb. nov. Scale bar = 0.50 mm. Revision of Piestus with remarks on related genera of elytra, subparallel sides of abdomen, segments 3 and 4 slightly narrower than 5–7, and accessory gland at the base of spermatheca (Fig. 277). Piestus aper has an incomplete V-shaped frontal sulcus (Fig. 98), one short seta on dorsal margin of the eyes, one long seta on scape, smooth internal borders of mandibles, stria 6 absent on elytra, abdominal segment 5 wider than others, and accessory gland at apical third of spermatheca (Fig. 283). Piestus imperfectus, sp. nov. (Figs 44, 92, 226, 227, 278) Type material Holotype deposited in SEMC, male, with labels: (1) ‘Ecuador: Pichincha/ Nanegalito, 7 km S, 1500m/0
00 23’N, 78
400 36’W/28 Oct 1999, R. Anderson/ECU1A99 212B/ex: riparian mont. evergreen litter’ [white label, printed in black]; (2) ‘bar code/SM0359240/KUNHM-ENT’ [white label, printed in black]. 36 paratypes deposited in following collections: in SEMC: 1 female, labels: (1) ‘Ecuador: Esmeraldas/Bilsa, 0
200 0’S, 79
430 0’W/28 Apr-10 May 1996/ECU1H96 015, P. Hibbs/ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0088817/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/Maquipucuna For. Res./50 km NW Quito, 1660m/ 21 Dec. 1991, C. Carlton/R. Leschen #22’ [white label, printed in black]; (2) ‘Piestus/(Tachypiestus)/sp./det. J.S. Ashe 19’ [white label, the first three lines handwritten, fourth line printed in black]; 3) ‘near P./(Tachypiestus)/ chiriquensis Shp./det. J.S. Ashe 1992’ [white label, first three lines handwritten, fourth printed in black]; 2 sex undetermined, with the same first label: (1) ‘Ecuador: Pichincha/Maquipucuna For. Res./50 km NW Quito, 1660m/21 Dec. 1991, C. Carlton/R. Leschen #22’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/45 km NNW Quito/Macquipucuna Station/1600-1650m/3-18 Apr 1996/ECU1H96 012, P. Hibbs/ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0090869/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0090885/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/45 km NNW Quito/Macquipucuna Station/16001650m/18 Apr-5 May 1996/ECU1H96 013, P. Hibbs/ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0090741/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/Mindo, 10.6 km W, Mindo Road/0
40 23’S, 78
450 14’W, 1460m/26–29 Mar 1999, R. Brooks, D. Brzoska/ECU1B99 056 ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/ SM0157957/KUNHM-ENT’ [white label, printed in black]; 1 female, labels: (1) ‘Ecuador: Pichincha/Mindo, 10.6 km W, Mindo Road/ 0
40 23’S, 78
450 14’W, 1445m/26–29 Mar 1999, R. Brooks, D. Brzoska/ ECU1B99 064 ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0157373/KUNHM-ENT’ [white label, printed in black]; 1 female, labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological Station/ 0
70 0’N, 78
380 6’W 1200m/27 Oct 1999, R. Anderson/ECU1A99 208B/ex: montane evergreen forest litter’ [white label, printed in black]; (2) ‘bar code/ SM0362386/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, labels: 1) ‘Ecuador: Pichincha/Maquipucuna Biological Station/0
60 25’N, 78
370 18’W 1480m/27 Oct 1999, R. Anderson/ ECU1A99 209A/ex: montane evergreen forest litter’ [white label, printed in black]; (2) ‘bar code/SM0363280/KUNHM-ENT’; ‘Piestus/spp./det. J.S. Ashe 2000’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological Station/River Trail, 1200m/0
70 34’N, 78
370 57’W/27 Oct 1999, Z.H. Falin/ECU1F99 025/ex: under bark’ [white label, printed in black]; (2) ‘bar code/ SM0350256/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological Invertebrate Systematics 541 Station/0
70 0’N, 78
380 6’W 1200m/27 Oct 1999, R. Anderson/ECU1A99 208C/ex: montane evergreen forest litter’ [white label, printed in black]; (2) ‘bar code/SM0363664/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0363666/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Pichincha/Maquipucuna Biological Station 1275m/0
70 22’N, 79
390 0’W/27–29 Oct 1999, Z.H. Falin/ECU1F99 048/ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0355819/KUNHMENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0355818/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0355820/ KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0355821/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/ SM0355839/KUNHM-ENT’ [white label, printed in black]; 1 female, labels: (1) ‘Ecuador: Pichincha/Bellavista Reserve, Ridge Trail/12 km S Nanegalito, 2250m/0
00 54’S, 78
400 56’W/28 Oct 1999, R. Anderson/ ECU1A99 211H/ex: cloud forest litter’ [white label, printed in black]; (2) ‘bar code/SM0357088/KUNHM-ENT’ [white label, printed in black]; 1 sex undetermined, labels: (1) ‘Ecuador: Napo, 2300m/Sierra Azul, Hacienda Arargon/0
400 0’S, 77
550 0’W/17 Feb-26 Mar 1996/ECU1H96 009, P. Hibbs/ex: flight intercept trap’ [white label, printed in black]; (2) ‘bar code/SM0087583’ [white label, printed in black]; 1 sex undetermined, the same first label, (2) ‘bar code/SM0087727’ [white label, printed in black]; 1 female, labels: (1) ‘Ecuador: Napo, Cosanga/4.2 km S on Baeza-Tena/Road then 2.9 km W on pipeline/access road, 2150m/0
370 19’S, 77
500 1’W/6 Nov 1999, Z.H. Falin/ECU1F99 113 ex: under bark’ [white label, printed in black]; (2) ‘bar code/SM0352655/KUNHM-ENT’ [white label, printed in black]; in AMNH: 3 sex undetermined, label: (1) ‘Ecuador: Pichincha:/ Palmeras, old Sto. Dgo./rd, km 59, 43 km NE/Alluriquin, X-15-88/6400’, light, L. Herman’ [white label, printed in black]; 1 female, 2 sex undetermined, label: (1) ‘Ecuador: Pichincha:/18–20 km NE Alluriquin/ old Quito-Sto. Domingo rd/X-21-88 4,700–4,900’/litter, L. Herman’ [white label, printed in black]; 1 female, 3 sex undetermined, labels: (1) ‘Ecuador: Pichincha Prov./W of Alluriquin, 3.3–5.3 km/SW road to Cooperativa/ Bolivar, near Tinalandia,’ [white label, printed in black]; (2) ‘3100-3500, V-20-93; L. Herman/#2733-2734, litter nr stream’ [white label, printed in black]; in DZUP, 1 male, 1 sex undetermined, label: 1) ‘Ecuador: Pichincha/ Maquipucuna For. Res./50 km NW Quito, 1660m/21 Dec. 1991, C. Carlton/ R. Leschen #22’ [white label, printed in black]; in FMNH, 2 sex undetermined, label: (1) ‘Ecuador: Pichincha/Maquipucuna For. Res./ 50 km NW Quito, 1660m/21 Dec. 1991, C. Carlton/R. Leschen #22’ [white label, printed in black]. Description BL: 4.5–5.8 mm, BW: 1.4–1.6 mm. Body somewhat convex; reddish brown to reddish dark brown (Fig. 44). Dorsal integument of head, except anterior angles, and all pronotum with large punctures contiguous (Fig. 89); striae of elytra with large punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 92); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and prominent; eyes with a moderately sized puncture with long seta on basal half of dorsal margin. Antennae reaching apex of abdomen; scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape (Fig. 44). Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with ventral 542 Invertebrate Systematics teeth curved, not projected; dorsal teeth weakly developed; internal borders symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as wide as long. Pronotum wider than long (PW/PL = 1.45) (Fig. 92); anterior angles weakly projected; on apical half with slightly somewhat longitudinal median depression and basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with slightly sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.07); stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae not carinate; epipleural carina developed. Metaventrite with conspicuous median depression near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with pair of long setae on apex. Median lobe of aedeagus with bulbous base in ventral view and slightly curved apex in lateral view, tube longer than base (Fig. 226); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 227. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape without enlarged area and with one long setae at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct long, ~3 times length of spermatheca, without sclerotised basal region (Fig. 278); spermatheca L-shaped (Fig. 278). Distribution Ecuador (Esmeralda, Pichincha and Napo). Biological notes This species has been found in litter, under bark and also collected in flight intercept traps. Remarks Piestus imperfectus is similar to P. chiriquensis, but differs mainly by having an incomplete V-shaped frontal sulcus (Fig. 92), curved apex of median lobe of aedeagus and tube longer than base (Fig. 226), long spermathecal duct, ~3 times the length of spermatheca and an L-shaped spermatheca (Fig. 278). Piestus chiriquensis has a complete V-shaped frontal sulcus (Fig. 94), slightly curved apex of median lobe and short tube with the same length as base (Fig. 230), short spermathecal duct, about the same length as spermatheca, and sinuous shaped spermatheca (Fig. 280). Piestus imperfectus, as well as P. chiriquensis, may be easily separated from P. nevermanni and P. costatus by slightly enlarged area with long setae on scape (somewhat as a tuft) and two acute teeth on symmetrical internal borders of mandibles (only a tuft of long setae on scape of male and only one acute tooth E. Caron et al. on internal borders of mandibles in P. nevermanni and P. costatus). Etymology The name refers to the incomplete V-shaped frontal sulcus on the median of head. Piestus costatus Sharp, 1887 (Figs 45, 93, 228, 229, 279) Piestus costatus Sharp, 1887: 715 (original description, type locality: ‘Panama, Volcan de Chiriqui 2000 to 3000 feet’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1790. Piestus (Piestus) costatus: Bernhauer & Schubert, 1910: 7 (distribution). Piestus (Trachypiestus) costatus: Scheerpeltz, 1952: 294 (characters, distribution). Type material Holotype deposited in BMNH, female, with labels: (1) ‘Piestus costatus/ Type D.S./V. de Chiriqui/2-3000ft. Champion’ [white labels, together the specimen, handwritten]; (2) ‘Holo-/Type’ [circle white label with border red, printed in black]; (3) ‘B.C.A. Col. I. 2./Piestus/costatus,/Sharp.’ [white label, printed in black]; (4) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (5) ‘Holotype/Piestus/costatus/Sharp, 1887/det. R.G. Booth 2008’ [white label, the four first lines handwritten, the last one printed in black]. Note: In the original description Sharp (1887) specified only one specimen (female) observed. Additional material See Appendix 2. Redescription BL: 3.8–5.2 mm, BW: 1.1–1.4 mm. Body somewhat convex; reddish brown to reddish dark brown (Fig. 45). Dorsal integument of head, except anterior angles, and all pronotum with large punctures contiguous (Fig. 93); striae of elytra with inconspicuous inclinate microstriae and large punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 93); V-shaped frontal sulcus complete, sometimes inconspicuous at middle; anterior angles curved and prominent; eyes with a moderately sized puncture with long seta on basal half of dorsal margin. Antennae almost reaching apex of abdomen (Fig. 45); scape with a few long setae on basal half of exposed dorsal face (somewhat as a tuft); scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with weak area that resembles a tooth, absent in the left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as long as wide. Pronotum wider than long (PW/PL = 1.28) (Fig. 93); anterior angles not projected; apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; Revision of Piestus with remarks on related genera abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with slightly sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.05), sometimes occulting tergite 3; stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; epipleural carina developed. Metaventrite with conspicuous median depression near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with pair of long setae on apex. Median lobe of aedeagus with bulbous base in ventral view and slightly curved apex in lateral view, tube longer than base (Fig. 228); lateral lobes not reaching apex of median lobe in lateral view; internal sclerites as in Fig. 229. Female. Similar to male, except for: antennae reaching apex of elytra; scape without enlarged area and with one long setae at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct long, ~5 times length of spermatheca, without sclerotised basal region (Fig. 279); spermatheca L-shaped (Fig. 279). Distribution Costa Rica (San José, Cartago and Puntarenas) and Panama (Bocas del Toro and Chiriquí). Biological notes Piestus costatus has been found in litter, near streams or associated with decaying logs. Some specimens were collected in flight intercept traps and by Berlese extraction of leaf litter. Remarks Piestus costatus is very similar to P. nevermanni but differs by having a long spermathecal duct, ~5 times the length of spermatheca, without sclerotised basal region, and spermatheca L-shaped (Fig. 279) (short spermathecal duct, about the same length as spermatheca, and spiral-shaped spermatheca, in P. nevermanni; Fig. 281). Piestus costatus is also similar to P. imperfectus and P. chiriquensis, from which it differs by characters cited above (see remarks under P. imperfectus). Piestus chiriquensis Sharp, 1887 (Figs 46, 94, 230, 231, 280) Piestus chiriquensis Sharp, 1887: 715 (original description, type locality: ‘Panama, Volcan de Chiriqui 2000 to 4000 feet’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1790 (distribution). Piestus (Piestus) chiriquensis: Bernhauer & Scheerpeltz, 1910: 7 (distribution). Piestus (Trachypiestus) chiriquensis: Scheerpeltz, 1952: 294 (characters, distribution). Type material Syntype deposited in FMNH, sex undetermined, labels: (1) ‘Piestus chiriquen/sis D.S./V. de Chiriqui 2500/4000ftt. Champion’ [white label, together with the specimen, handwritten]; (2) ‘V. de Chiriquí,/2-3000 ft/ Champion’ [white label, printed in black]; (3) ‘B.C.A. Col. I. 2./Piestus/ Invertebrate Systematics 543 chiriquensis/Sharp.’ [white label, printed in black]; (4) ‘Chicago Nat. Hist. Mus./(ex. D. Sharp Colln./by exchange with/Brit. Mus. Nat. Hist.)’ [white label, printed in black]. Note: There are 4 syntypes (not examined) deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). Additional material See Appendix 2. Redescription BL: 4.2–5.8 mm, BW: 1.2–1.6 mm. Body somewhat convex; light brown to reddish dark brown (Fig. 46). Dorsal integument of head, except anterior angles, and all pronotum with large punctures contiguous (Fig. 94); striae of elytra with large punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 94); V-shaped frontal sulcus complete; anterior angles curved and prominent; eyes with a moderately sized puncture with long seta on basal half of dorsal margin. Antennae reaching apex of abdomen; scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape (Fig. 46). Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as wide as long. Pronotum wider than long (PW/PL = 1.45) (Fig. 94); anterior angles weakly projected; on apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with slightly sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.04), occulting tergite 3; stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; epipleural carina developed. Metaventrite with conspicuous median depression near apex. Abdominal segments 3–6 with subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and slightly curved apex in lateral view, tube short with same length of base (Fig. 230); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 231. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape without enlarged area and with one long seta at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct short, about same length of spermatheca, without sclerotised basal region (Fig. 280); spermatheca with sinuous shape (Fig. 280). 544 Invertebrate Systematics Distribution In the current study the species was examined from Nicaragua (Río San Juan), Costa Rica (Guanacaste, Alajuela, Heredia, San José, Cartago, Limón and Puntarenas), Panama (Bocas del Toro, Chiriquí, Panamá and Darién), Colombia (Cundinamarca and Nariño), Venezuela (Lara), Ecuador (Esmeraldas, Sucumbios, Pichincha, Napo, Cotopaxi and Tungurahua), Peru (Huánuco, Cusco) and Bolivia (Cochabamba). Remarks Piestus chiriquensis is very similar to P. imperfectus and also to P. costatus and P. nevermanni, from which it is easily separated by characters cited above (see remarks under P. imperfectus). Biological notes Piestus chiriquensis has been found in litter, near streams or associated with decaying logs. Some specimens were collected in flight intercept and Malaise traps and by Berlese extraction of leaf litter. Piestus nevermanni Scheerpeltz, 1952 (Figs 47, 95, 232, 233, 281) Piestus (Trachypiestus) nevermanni Scheerpeltz, 1952: 294, 302 (original description, type locality: ‘Costarica’). Piestus nevermanni: Herman, 2001b: 1792 (distribution). Type material Holotype deposited in NMW, female [damaged specimen: without right anterior and median legs], with labels: (1) ‘<’ [white label, printed in black]; (2) ‘Costarica/leg. nevermann’ [white label, handwritten]; (3) ‘Dr M Bernhauer/8.11. donavit 1933’ [white label, first line printed in black, second line handwritten, except the word ‘donavit’ printed in black]; (4) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (5) ‘TYPUS/ Piestus (Trachy-/piestus) costa-/ricensis/O. Scheerpeltz’ [red label, first line and fifth printed in black, the others handwritten]; (6) ‘nevermanni’ [light green label, handwritten]. Note: In the original description Scheerpeltz (1952) specified one male specimen observed. However, the observed type from NMW is female. E. Caron et al. third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in the right mandible with weak area that resembles a tooth, absent in the left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as long as wide. Pronotum wider than long (PW/PL = 1.48) (Fig. 95); anterior angles weakly projected; on apical half with a somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with a short tooth and slight emargination; apical two-thirds with slightly sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.06); stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; epipleural carina developed. Metaventrite with conspicuous median depression near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with long ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view, tube longer than base (Fig. 232); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 233. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape without enlarged area and with one long seta at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct short, about same length of spermatheca, without sclerotised basal region (Fig. 281); spermatheca with spiral shape (Fig. 281). Distribution Costa Rica and Panama (Chiriqui). Additional material Biological notes See Appendix 2. This species has been collected in flight intercept traps. Redescription BL: 6.0 mm, BW: 1.7 mm. Body somewhat convex; brown to reddish dark brown (Fig. 47). Dorsal integument of head, except anterior angles, and all pronotum with large punctures contiguous (Fig. 95); striae of elytra with large punctures; metaventrite, only on lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 95); V-shaped frontal sulcus complete; anterior angles curved and prominent; eyes with a moderately sized puncture with long seta on basal half of dorsal margin. Antennae reaching apex of abdomen; scape with one tuft of long setae on basal half of exposed dorsal face; scape shorter than the antennomere 2 and 3 combined; antennomeres 4–11 oblong shape (Fig. 47). Labrum with six setae medially, equal in length; each lateral Remarks Piestus nevermanni is very similar to P. imperfectus and also to P. costatus and P. chiriquensis, differing from these species by characters cited above (see remarks under P. imperfectus). Piestus paradoxus Bernhauer, 1917 (Figs 48, 49, 96, 127, 234, 235, 282) Piestus paradoxus Bernhauer, 1917: 45 (original description, type locality: ‘Caracas’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1793 (distribution). Piestus (Piestus) paradoxus: Scheerpeltz, 1952: 994 (distribution). Piestus (Elytropiestus) paradoxus: Scheerpeltz, 1952: 294 (characters, distribution). Revision of Piestus with remarks on related genera Type material Holotype deposited in FMNH, female [damaged specimen: without left antennomeres 3–11], with labels: (1) ‘Caracas/Bang. Haas.’ [white label, handwritten]; (2) ‘paradoxus/Bernh./Typus unic.’ [light yellow label, handwritten]; (3) ‘Chicago NHMus/M.Bernhauer/Collection’ [white label, printed in black]. Note: In the original description Bernhauer (1917) specified only one specimen observed. Additional material See Appendix 2. Invertebrate Systematics 545 Female. Similar to male, except for: antennae almost reaching apex of abdomen; scape without enlarged area and with two long setae at middle of dorsal face; prosternum without ovate set of fine punctures; each elytron with interstriae 2 and 4 joined at apex and strongly posteriorly extended, forming a spine; ovipositor as in P. sulcatus; spermathecal duct short, ~1.5 times length of spermatheca, without sclerotised basal region (Fig. 282); spermatheca with somewhat globose apex (Fig. 282). Distribution Venezuela (Aragua). Redescription BL: 5.0–5.7 mm, BW: 1.5–1.6 mm. Body somewhat convex; reddish brown to reddish dark brown (Figs 46, 47). Dorsal integument of head, except anterior angles, and all pronotum with undulate microstriae and large punctures contiguous (Fig. 96); striae of elytra with large punctures; metaventrite, lateral regions, and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 96); V-shaped frontal sulcus complete, sometimes inconspicuous at the middle; anterior angles curved and prominent, and slightly median longitudinal sulcus; eyes with a moderately sized puncture with long seta on basal half of dorsal margin. Antennae exceeding apex of abdomen (Fig. 48); scape with slightly enlarged area and long setae on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal shorter than postero-external seta. Mandibles with dorsal teeth weakly developed (Fig. 127); internal border symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as wide as long. Pronotum wider than long (PW/PL = 1.45) (Fig. 96); anterior angles not projected; on apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with a short tooth and slight emargination; apical two-thirds with slightly sinuous sides. Prosternum with conspicuous ovate set of fine punctures in a median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra together longer than wide (EL/BW = 1.17), occulting tergite 3 and basal half of tergite 4; stria 6 at basal half and stria 7 complete; striae with same width of interstriae; interstriae carinate; interstriae 2 and 4 joined at apex and weakly posteriorly extended; epipleural carina developed. Metaventrite with conspicuous median fovea near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with one pair of long setae on apex, sometimes with one of inconspicuous seta near long setae. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 234); lateral lobes exceed a little apex of median lobe in lateral view; internal sclerites as in Fig. 235. Biological notes This species has been noted only as ‘under bark’ and on ‘fungusy log’. Remarks Piestus paradoxus is easily separated from the other species of Piestus, except P. acuminatus, by the following character: interstriae 2 and 4 of each elytron joined at the apex and weakly posteriorly extended on males and strongly on females (Figs 48, 49). Piestus paradoxus may be confused with P. acuminatus because they also have interstriae posteriorly extended on elytra. However, P. paradoxus has very long antennae, exceeding the apex of abdomen in males and almost reaching the apex of abdomen in females (Figs 48, 49), only large punctures on striae of elytra, conspicuous median fovea near the apex of metaventrite, long tube of median lobe of aedeagus and lateral lobes exceeding the apex of median lobe (Fig. 234), and somewhat globose apex of spermatheca (Fig. 282). Piestus acuminatus has shorter antennae, almost reaching the apex of elytra in males and reaching the apex of elytra in females (Fig. 53), microgranulate texture and large punctures on striae of elytra, slightly median depression near the apex of metaventrite, short tube of median lobe of aedeagus (Fig. 242) and lateral lobes reaching the apex of median lobe, and very elongate spermatheca (Fig. 285). Piestus boliviensis, sp. nov. (Figs 50, 97, 236, 237) Type material Holotype deposited in SEMC, male, labels: (1) ‘Bolivia: La Paz, Coroico/Cerro Uchumachi, 2150m/16
12.160 S, 67
43.330 W/27-I-2001, R.A. Anderson,/2nd growth cloud forest litter/BOL1A01-013’ [white label, printed in black]; (2) ‘bar code/SM0454517/KUNHM-ENT’ [white label, printed in black]; (3) ‘Piestus/sp./Det: J.S. Ashe, 2004’ [white label, first two lines handwritten, third printed in black]. Description BL: 5.3 mm, BW: 1.4 mm. Body flattened dorsoventrally; reddish dark brown (Fig. 50). Dorsal integument of head, except anterior angles, and all pronotum with large punctures contiguous (Fig. 97); striae of elytra with large punctures; 546 Invertebrate Systematics metaventrite and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 97); V-shaped frontal sulcus complete; anterior angles curved and prominent; eyes with a moderately sized puncture with short seta on basal half of dorsal margin. Scape antennal with long setae on basal half of exposed dorsal face; scape subequal to antennomere 2 and 3 combined (Fig. 50); antennomeres 4–5 oblong shape (material damaged, not possible to verify antennomeres 6–11). Mandibles with dorsal teeth weakly developed; internal border slightly asymmetrical, each with one small and acute tooth at middle and, anterior to it, in right mandible with weak area that resembles a tooth, absent in left mandible. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Pronotum wider than long (PW/PL = 1.42) (Fig. 97); anterior angles weakly projected; on apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with slightly sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra together shorter than wide (EL/BW = 0.91), apical margin strongly emarginate at middle; stria 6 absent and stria 7 complete; striae with same width of interstriae; interstriae carinate; epipleural carina developed. Metaventrite with a conspicuous median depression near apex. Abdominal segments 3–6 with subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with two pairs of setae on apex (material damaged, not possible to verify length of setae). Median lobe of aedeagus with bulbous base in ventral view and slightly curved apex in lateral view (Fig. 236); lateral lobes exceed little the apex of median lobe in lateral view; internal sclerites as in Fig. 237. Female. Unknown. E. Caron et al. Piestus aper Sharp, 1876 (Figs 51, 98, 238, 239) Piestus aper Sharp, 1876: 408 (original description, type locality: ‘St. Paulo’); Blackwelder, 1944: 100 (distribution); Herman, 2001a: 12 (mention); Herman, 2001b: 1788 (distribution). Piestus (Piestus) aper: Bernhauer & Schubert, 1910: 6 (distribution). Piestus (Trachypiestus) asper Scheerpeltz, 1952: 292 (emendation of P. aper Sharp, 1876, characters, distribution). Piestus asper: Herman, 2001a: 12 (note: unjustified emendation by Scheerpeltz (1952)); Herman, 2001b: 1788 (mention). Piestus (Trachypiestus) schadei Scheerpeltz, 1952: 295 (original description, type locality: ‘Villa Rica (Paraguay)’). New synonym. Piestus schadei: Herman, 2001b: 1794 (distribution). Type material Piestus aper Sharp, 1876. Lectotype deposited in BMNH, sex undetermined, with labels: (1) ‘Piestus/aper./Amazons/Type D.S.’ [white label, together with the specimen, handwritten]; (2) ‘Syn-/Type’ [circle white label with border light blue, printed in black]; (3) ‘Type’ [circle white label with border red, printed in black]; (4) ‘Amaz./Sr. Paula?’ [circle green label, handwritten]; (5) S. America:/Brazil.’ [white label, printed in black]; (6) ‘Sharp Coll/1905-313.’ [white label, printed in black]; (7) ‘Syntype/Piestus apes?/Sharp, 1876/det. R.G. Booth 2008’ [white label, the three first lines handwritten, the last one printed in black]. Note: In the original description Sharp (1876) specified two specimens of sex undetermined. We received from BMNH only one specimen. Another specimen is deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). Piestus (Trachypiestus) schadei Scheerpeltz, 1952. Lectotype deposited in NMW, sex undetermined, with labels: (1) ‘Villa Rica/ Paraguay’ [white label, printed in black]; (2) ‘Fr. Schade leg./8. XII. 1924’ [white label, first line printed in black, second line handwritten, except ‘1924’ printed in black]; (3) ‘ex coll./Scheerpeltz’ [blue label, printed in black]; (4) ‘TYPUS/Piestus (Trachy-/piestus) Schadei/O. Scheerpeltz’ [red label, the first and fourth lines printed in black, the second and third handwritten]; (5) ‘Schadei/Scheerp.’ [light green label/handwritten]. Note: In the original description Scheerpeltz (1952) specified a male and female observed. However, we received from NMW only one specimen. Additional material See Appendix 2. Distribution Redescription Bolivia (La Paz). BL: 3.2–5.2 mm, BW: 1.0–1.4 mm. Body flattened dorsoventrally; light brown to reddish dark brown (Fig. 51). Dorsal integument of dorsum of head and all pronotum with large punctures contiguous (Fig. 98); striae of elytra with large punctures; metaventrite and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 98); V-shaped frontal sulcus incomplete, curved arms not joined medially; anterior angles curved and prominent; eyes with a moderately sized puncture with short seta on basal half of dorsal margin. Antennae exceed little apex of elytra (Fig. 51); scape with one long seta at middle of dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta longer than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border smooth. Maxillary palpus with palpomere 4 longer than to 2 and 3 combined. Mentum ~1.5 times as wide as long. Pronotum wider Biological notes The holotype was collected in ‘2nd growth cloud forest litter’. Remarks Piestus boliviensis is easily separated from the other species of Piestus by the slightly asymmetrical internal borders of mandibles, only one small and acute tooth in each one, and elytra together shorter than wide with apical margin strongly emarginate at the middle (Fig. 50). Etymology The species name is derived from the name of Bolivia, where the holotype was collected. Revision of Piestus with remarks on related genera than long (PW/PL = 1.56) (Fig. 98); anterior angles strongly projected; apical half with a somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles strongly developed and exceed middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with a short tooth and slight emargination; apical two-thirds with sinuous sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra together as long as wide (EL/BW = 0.99); stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; basal half of interstriae 2 and 5 and apical half of interstria 4 prominent; epipleural carina developed. Abdominal segments 3–6 with subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with long ventral struts; tergite 10 with one pair of long setae on apex, sometimes with one pair of inconspicuous setae near long setae. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 238); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 239. Female. Similar to male. Ovipositor as in P. sulcatus; spermathecal duct ~2 times length of spermatheca, without sclerotised basal region (Fig. 283); spermatheca with globose apex and accessory gland at apical third (Fig. 283). Invertebrate Systematics Piestus (Piestus) crassicornis: Bernhauer & Schubert, 1910: 7 (distribution). Piestus (Trachypiestus) crassicornis: Scheerpeltz, 1952: 292 (characters, distribution). Type material Piestus angularis Fauvel, 1864. Holotype deposited in IRSNB, female, with labels: (1) ‘Brésil/Sa Catharina’ [white label, handwritten]; (2) ‘Caracas’ [white label, handwritten]; (3) ‘Mexique’ [white label, handwritten]; (4) ‘crassicornis Shp./parait u? même’ [white label, handwritten]; (5) ‘angularis/FvL.’ [white label, handwritten]; (6) ‘R.I.Sc.N.B. 17.479/ Piestus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black and second line handwritten]; (7) ‘Type’ [red label, printed in black]. Note: In the original description Fauvel (1864) specified only one specimen observed. Piestus crassicornis Sharp, 1887. Holotype deposited in BMNH, female, with labels: (1) ‘Piestus crassi-/cornis Type D.S./V. de Chiriqui 2500-/4000ft. Champion’ [white label, together with the specimen, handwritten]; (2) ‘Holo-/ Type’ [circle white label with border red, printed in black]; (3) ‘Sp. figured.’ [white label, printed in black]; (4) ‘V. de Chiriqui,/25-4000 ft./Champion.’ [white label, printed in black]; (5) ‘B.C.A. Col. I. 2./Piestus/crassicornis,/ Sharp.’ [white label, printed in black]; (6) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (7) ‘Holotype/Piestus/crassicornis/Sharp, 1887/det. R.G. Booth 2008’ [white label, the four first lines handwritten, the last one printed in black]. Note: In the original description Sharp (1887) specified only one specimen observed. Distribution Additional material In the current study the species was examined from Panama (Bocas del Toro and Colón), Colombia (Amazonas), Suriname (Marowijne), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Peru (Loreto, Cusco and Madre de Dios), Bolivia, Brazil (Amazonas, Bahia and Mato Grosso) and Paraguay (Guairá and Itapúa). Scheerpeltz (1952) also listed P. aper from north Argentina. See Appendix 2. Biological notes Piestus aper has been found in litter, and associated with fruit falls, flower falls or decaying logs. Some specimens were collected in flight intercept traps, or by Berlese and Winkler extraction of leaf litter. Remarks This species is similar to P. sulcipennis but differs by characters cited under remarks of P. sulcipennis. Piestus angularis Fauvel, 1864 (Figs 52, 99, 240, 241, 284) Piestus angularis Fauvel, 1864: 31 (original description, type locality: ‘Brésil (Santa-Catharina’); Blackwelder, 1944: 100 (distribution, error: Fauvel, 1865: 35); Herman, 2001b: 1788 (distribution); NavarreteHeredia et al., 2002: 208 (distribution). Piestus (Piestus) angularis: Bernhauer & Schubert, 1910: 6 (distribution, error: Fauvel, 1865: 35). Piestus (Trachypiestus) angularis: Scheerpeltz, 1952: 292 (characters, distribution, error: Fauvel, 1865: 35). Piestus crassicornis Sharp, 1887: 716 (original description, type locality: ‘Panama, Volcan de Chiriqui 4000 to 6000 feet’); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1790 (distribution). New synonym. 547 Redescription BL: 4.6–6.8 mm, BW: 1.4–1.9 mm. Body somewhat convex; reddish brown to reddish dark brown (Fig. 52). Dorsal integument of head, except anterior angles, and all pronotum with large punctures contiguous (Fig. 99); striae of elytra with large punctures; metaventrite and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 99); V-shaped frontal sulcus complete; anterior angles curved and prominent, and conspicuous median longitudinal sulcus with fovea at the base; eyes with moderately sized puncture with short seta on basal half of dorsal margin. Antennae reaching apex of abdomen (Fig. 52); scape with one long seta at middle of dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta as long as postero-external seta. Mandibles with dorsal teeth weakly developed; internal border symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than to 2 and 3 combined. Mentum subquadrate, as wide as long. Pronotum wider than long (PW/PL = 1.31) (Fig. 99); anterior angles not projected; on apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles strongly developed and exceed middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and a slight emargination; apical twothirds with sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median concavity near the apex. Femur of anterior legs with short, hard setae on groove of ventral margin. 548 Invertebrate Systematics Elytra together as long as wide (EL/BW = 0.98); stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; basal half of interstriae 2 and 5 prominent; epipleural carina developed. Metaventrite with conspicuous median fovea near apex. Abdominal segments 3–6 with subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with short ventral struts; tergite 10 with one pair of long setae on apex, sometimes with one pair of inconspicuous setae near long setae. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 240); lateral lobes exceed little apex of median lobe in lateral view; internal sclerites as in Fig. 241. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; prosternum without ovate set of fine punctures and without median concavity near apex; ovipositor as in P. sulcatus; spermathecal duct long, ~5 times length of spermatheca, without sclerotised basal region (Fig. 284); spermatheca somewhat L-shaped (Fig. 284). Distribution In the current study the species was examined from Mexico (Veracruz), Belize (Cayo), Guatemala (Zacapa), Costa Rica (Alajuela, Heredia, San José, Limón and Puntarenas), Panama (Chiriquí), Suriname (Marowijne and Para), French Guiana (Saint-Laurent-du-Maroni and Cayenne), Ecuador (Esmeraldas, Napo and Zamora-Chinchipe), Peru (Cusco), Bolivia (Cochabamba), Brazil (Santa Catarina) and Paraguay (Alto Paraná). Navarrete-Heredia et al. (2002) list P. angularis also from Honduras, Nicaragua and Colombia. Biological notes Piestus angularis has been found in litter, near streams or associated with decaying logs. Some specimens were collected by flight intercept and Malaise traps. Remarks This species is easily distinguished from the other species of Piestus by the conspicuous median longitudinal sulcus with a fovea at the base of the vertex (Fig. 99), two acute teeth on symmetrical internal borders of mandibles, and only one long seta on scape (male and female). Piestus acuminatus, sp. nov. (Figs 53, 100, 135, 144, 242, 243, 285) Type material Holotype deposited in SEMC, male, labels: (1) ‘Ecuador: Napo/El Chaco, 4.8 km NW on road/to Oyacachi, 1750m/0
180 22’S, 77
500 38’W/7 Nov 1999, Z.H. Falin/ECU1F99 119 ex: pyrethrum/fogging mildewy downed tree’ [white label, printed in black]; (2) ‘bar code/SM0351487/KUNHMENT’ [white label, printed in black]. 3 paratypes distributed: 2 in SEMC: 1 female, the same first label of holotype, (2) ‘bar code/SM0351486/KUNHMENT’ [white label, printed in black]; 1 male, labels: (1) ‘Ecuador: Napo/ Baeza-Tena Road, 15 km/W of Cosanga, Reserva Sierra/Azul, 2350m, 0
400 55’S, 77
560 9’W/5 Nov 1999, Z.H. Falin/ECU1F99 097 ex: rotting cut lumber’ [white label, printed in black]; (2) ‘bar code/SM0355660/ E. Caron et al. KUNHM-ENT’ [white label, printed in black]; 1 in FMNH, 1 male, labels: (1) ‘Ecuador: Napo/Cosanga, 4.2 km S on Baeza-Tena/Road then 2.9 km W on pipeline/access road, 2150m/0
370 19’S, 77
500 1’W/7 Nov 1999, Z.H. Falin/ECU1F99 120 ex: on/under/bark downed logs’ [white label, printed in black]; (2) ‘bar code/SM0152154/KUNHM-ENT’ [white label, printed in black]. Description BL: 5.0 mm, BW: 1.6 mm. Body convex; reddish brown to reddish dark brown (Fig. 53). Dorsal integument of head and all pronotum with large punctures contiguous; striae of elytra with microgranulate texture and large punctures (Fig. 100); metaventrite and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 100); Vshaped frontal sulcus complete; anterior angles curved and prominent, and with median fovea at base; eyes prominent with moderately sized puncture with short seta on basal half of dorsal margin. Antennae almost reaching apex of abdomen (Fig. 53); scape with long seta on basal half of exposed dorsal face; scape shorter than antennomere 2 and 3 combined; antennomeres 4–11 oblong shape. Labrum with six setae medially, equal in length; each lateral third, antero-internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as wide as long (Fig. 135). Pronotum wider than long (PW/PL = 1.48) (Fig. 100); anterior angles weakly projected; on apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus conspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra somewhat longer than wide (EL/BW = 1.05), occulting tergite 3; stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; interstriae 1 and 4 joined at apex and weakly posteriorly extended (Fig. 53); epipleural carina developed. Metaventrite with a conspicuous median depression near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina (Fig. 144); tergite 9 with short ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of the aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 242); lateral lobes reaching apex of median lobe in lateral view; internal sclerites as in Fig. 243. Female. Similar to male, except for: antennae reaching the apex of elytra; scape with two long setae at the middle of the dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct very short, about the half of length of spermatheca, without sclerotised basal region (Fig. 285); spermatheca elongate (Fig. 285). Distribution Ecuador (Napo). Revision of Piestus with remarks on related genera Biological notes This species was collected under bark and cited from ‘rotting cut lumber’. Remarks Piestus acuminatus is similar to P. rugosus and P. elegans because they share microgranulate texture and large punctures on striae of elytra. However, P. acuminatus is easily separated by interstriae 1 and 4 of each elytron joined at the apex and weakly posteriorly extended (male and female) (Fig. 53). Piestus rugosus and P. elegans do not have interstriae posteriorly projected. Piestus acuminatus is distinguished from P. paradoxus by characters cited under remarks for P. paradoxus. Etymology The name refers to the sharp point on the apex of each elytron. Piestus rugosus Sharp, 1876 (Figs 54, 101, 244, 245, 286) Piestus rugosus Sharp, 1876: 407 (original description, type locality: ‘Ega’); Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1794 (distribution). Piestus (Piestus) rugosus: Bernhauer & Schubert, 1910: 7 (distribution). Piestus (Trachypiestus) rugosus: Scheerpeltz, 1952: 294 (characters, distribution). Type material Holotype deposited in BMNH, female, with labels: (1) ‘Piestus rugosus/ Amazons/Type/D.S.’ [white label, together with the specimen, handwritten]; (2) ‘Holo-/type’ [circle white label with border red, printed in black]; (3) ‘Ega.’ [circle green label, handwritten]; (4) ‘S. America:/Brazil.’ [white label, printed in black]; (5) ‘Sharp Coll./1905-313.’ [white label, printed in black]; (6) ‘Holotype/Piestus/rugosus/Sharp, 1876/det. R.G. Booth 2008’ [white label, the four first lines handwritten and the last one printed in black]. Note: In the original description Sharp (1876) specified only one specimen observed. Invertebrate Systematics 549 internal seta shorter than postero-external seta. Mandibles with dorsal teeth weakly developed; internal border symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Mentum subquadrate, as wide as long. Pronotum wider than long (PW/ PL = 1.49) (Fig. 101); anterior angles not projected; on apical half with somewhat longitudinal median depression and on basal half a transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with short tooth and slight emargination; apical two-thirds with sinuous sides. Prosternum with conspicuous ovate set of fine punctures in median region near apex. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra together as long as wide (EL/ BW = 0.97); stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; epipleural carina developed. Metaventrite with a conspicuous depression near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina; tergite 9 with long ventral struts; tergite 10 with two pairs of long setae on apex, apical the longest. Median lobe of aedeagus with bulbous base in ventral view and curved apex in lateral view (Fig. 244); lateral lobes not reaching apex of median lobe in lateral view; internal sclerites as in Fig. 245. Female. Similar to male, except for: antennae exceeding apex of elytra, but not reaching apex of abdomen; scape with two long setae at middle of dorsal face; prosternum without ovate set of fine punctures; ovipositor as in P. sulcatus; spermathecal duct short, about same length of spermatheca, without sclerotised basal region (Fig. 286); spermatheca somewhat elongate (Fig. 286). Distribution Guyana, Ecuador (Sucumbios), (Cochabamba) and Brazil (Pará). Peru (Cusco), Bolivia Additional material Biological notes See Appendix 2. This species has been noted occurring under bark and it was also collected in flight intercept traps. Redescription BL: 4.0–6.0 mm, BW: 1.3–1.6 mm. Body somewhat convex; reddish brown to reddish dark brown (Fig. 54). Dorsal integument of head and all pronotum with large punctures contiguous (Fig. 101); striae of elytra with microgranulate texture and large punctures; metaventrite and abdominal tergites with moderately sized punctures uniformly distributed. Male. Head with front slightly deflected (Fig. 101); Vshaped frontal sulcus complete; anterior angles curved and prominent, and median longitudinal sulcus with fovea at base; eyes with moderately sized puncture with short seta on basal half of dorsal margin. Antennae almost reaching apex of abdomen; scape with few long setae on basal half of exposed dorsal face (as a tuft); scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 oblong shape (Fig. 101). Labrum with six setae medially, equal in length; each lateral third, antero- Remarks Piestus rugosus is similar to P. acuminatus and P. elegans because they share microgranulate texture and large punctures on striae of elytra. However, P. rugosus differs from P. acuminatus by characters cited under P. acuminatus, and it differs from P. elegans by antennomeres 4–11 oblong shape (Fig. 54), and short spermathecal duct about the same length as spermatheca (Fig. 286). Piestus elegans has antennomeres 4–11 somewhat bead-like in females (Fig. 55), and long spermathecal duct, ~2 times the length of spermatheca (Fig. 287). Piestus elegans, sp. nov. (Figs 55, 102, 287) 550 Invertebrate Systematics E. Caron et al. Type material Etymology Holotype deposited in SEMC, female, labels: (1) ‘Peru: Tambopata Prov./ Madre de Dios Dpto./15 km NE Puerto’ [white label, printed in black]; (2) ‘Maldonado Reserva/Cuzco Amazónico/12
330 S, 69
030 W/200m, Z2 E15’ [white label, printed in black]; (3) ‘26 June 1989, D. Silva/R. A. Leschen #285/ex., flight intercept’ [white label, printed in black]; (4) ‘Piestus/Peru sp. 3/det J.S. Ashe 19’ [white label, first two lines handwritten, third printed in black]. The specific name refers to the elegant, necklace-shaped antennomeres 4–11. Additional taxonomic notes Subfamily PIESTINAE Erichson, 1839 Description BL: 4.3 mm, BW: 1.2 mm. Body somewhat convex; reddish brown (Fig. 55). Dorsal integument of head and all pronotum with large punctures contiguous (Fig. 102); striae of elytra with microgranulate texture and large punctures; metaventrite and abdominal tergites with moderately sized punctures uniformly distributed. Female. Head with front slightly deflected (Fig. 102); Vshaped frontal sulcus complete; anterior angles curved and prominent, and with slightly median fovea at base. Antennae exceed base of elytra, but not reaching apex of elytra; scape with one short seta at middle of dorsal face; scape subequal to antennomere 2 and 3 combined; antennomeres 4–11 somewhat bead-like (Fig. 55). Mandibles with dorsal teeth weakly developed; internal border symmetrical, each with two acute teeth, basal anteriorly projected. Maxillary palpus with palpomere 4 longer than 2 and 3 combined. Pronotum wider than long (PW/PL = 1.53) (Fig. 102); anterior angles weakly projected; on apical half with slightly longitudinal median depression and on basal half transverse depression; longitudinal median sulcus inconspicuous; longitudinal sulcus of posterior angles developed and reaching middle of pronotum; abrupt constriction at basal third and, anterior to it, in each side with a short tooth and slight emargination; apical two-thirds with sinuous sides. Femur of anterior legs with short, hard setae on groove of ventral margin. Elytra together as long as wide (EL/ BW = 1.00); stria 6 absent and stria 7 complete; striae wider than interstriae; interstriae carinate; epipleural carina developed. Metaventrite with conspicuous median depression near apex. Abdominal segments 3–6 subparallel sides; sternite 3 with conspicuous transverse carina; ovipositor as in P. sulcatus; tergite 10 with two pairs of long setae on apex, apical the longest. Spermathecal duct ~2 times length of spermatheca, without sclerotised basal region (Fig. 287); spermatheca somewhat elongate (Fig. 287). Male. Unknown. Distribution Peru (Madre de Dios). Biological notes The holotype was collected in a flight intercept trap. Remarks Piestus elegans is similar to P. acuminatus and P. rugosus but differs by characters listed under remarks for P. acuminatus and P. rugosus. Hypotelus Erichson, 1839 Hypotelus Erichson, 1839: 31. For review of literature of the genus, see Herman (2001b). Antropiestus Bernhauer, 1917: 45 (as subgenus of Piestus); Scheerpeltz, 1952: 295 (subgenus of Piestus); Herman, 2001b: 1788 (subgenus of Piestus). Type-species. Piestus (Antropiestus) andinus Bernhauer, 1917 (fixed by original designation and monotypy). New synonym. Eccoptopiestus Scheerpeltz, 1952: 295 (as subgenus of Piestus); Herman, 2001b: 1788 (subgenus of Piestus). Type species. Piestus laevis Solsky, 1872 (fixed by original designation and monotypy). New synonym. Remarks Eccoptopiestus and Antropiestus, previously considered as subgenera of Piestus, are here placed as junior synonyms of Hypotelus, since their type species, P. (E.) laevis Solsky, 1872 and P. (A.) andinus Bernhauer, 1917, are transferred to Hypotelus based on some adult characters observed in type material (see remarks under each species below). Hypotelus laevis (Solsky, 1872), comb. nov. (Figs 56, 104, 128, 136, 148, 246, 247) Piestus laevis Solsky, 1872: 311 (original description: ‘Monte-Rico (Pérou)’); Blackwelder, 1944: 100 (distribution, error: Solsky, 1871); Herman, 2001b: 1791 (distribution). Piestus (Piestus) laevis: Bernhauer & Schubert, 1910: 7 (distribution, error: Solsky, 1871). Piestus (Eccoptopiestus) laevis: Scheerpeltz 1952: 295 (characters, distribution, error: Solsky, 1871). Type material Neotype (here designated) deposited in IRSNB, sex undetermined, with labels: (1) ‘Yuracaris/Bolivie’ [white label, handwritten]; (2) ‘laevis Solsky/var?’ [white label, handwritten]; (3) ‘R.I.Sc.N.B. 17.479/ Hypotelus/Coll. et det. A. Fauvel’ [white label, first and third lines printed in black, second line handwritten]. Note: In the original description Solsky (1872) did not specify how many specimens he observed. However, type material from Peru was not found in ZIN (Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia) where Solsky’s collection is deposited, and probably any types there are lost. However, we received from IRSNB (Fauvel’s collection) one specimen, which is here designated as neotype, and possibly it is an authentic specimen sent by Solsky to Fauvel. There is another case where this happened, Aleochara sareptana Solsky 1874 (Dr Didier Drugmand, IRSNB, pers. comm.). In this way, we are designating a neotype with the purpose of clarifying the taxonomic status of the taxon (Article 75.3.1, ICZN 1999). Additional material See Appendix 2. Revision of Piestus with remarks on related genera Redescription BL: 3.0–3.6 mm, BW: 0.8–0.9 mm. Body flattened dorsoventrally; light brown to dark brown with one-quarter apical of elytra darker (Fig. 56). Dorsal integument of head and pronotum with fine punctures and undulate microstriae; elytra with fine punctures disperse and only one longitudinal striae finely punctate, closely to internal margin. Male. Head with pair of broad and short pointed frontal processes, as long as scape, and basal distance between processes narrower than basal width of each one (Fig. 104); V-shaped frontal sulcus incomplete; anterior angles curved and weakly prominent; eyes with one moderately sized puncture with long setae on basal half of dorsal margin. Antennae short, almost reaching apex of elytra (Fig. 56); with one short seta near apex on the dorsal face; antennomeres 5–11 entirely with microsetae and some long setae dispersed; scape the longest, subequal to antennomere 2 and 3 combined; antennomere 4 shortest and 5–11 somewhat oblong shape. Labrum with six setae medially, external longest; each lateral third, three long setae, two apical and one subapical, the shortest. Mandibles with bifurcate apex, dorsal teeth shorter than ventral (Fig. 128); internal border with one acute tooth at middle; prostheca well developed. Maxillary palpus with palpomere 3 wider than long and 4 longer than 2 and 3 combined. Mentum 2 times wider than long and anterior angles emarginate (Fig. 136); ligula slightly emarginate and two pairs of conspicuous long setae on anterior margin, one pair near median sclerite and other on apex of median sclerite. Gular sutures joined. Pronotum wider than long (PW/PL = 1.33) (Fig. 104); anterior angles not projected; longitudinal median sulcus conspicuous; gradual constriction at basal third; apical three-quarters with slightly curved sides. Prosternum with anterior margin truncate. Elytra somewhat longer than wide (EL/BW = 1.18), covering tergite 3; when together slightly emarginate in middle; apical margin with short setae. Abdominal segments 3–6 parallel sides; segments 3–6 with two pair of paratergites, segment 7 with only one pair; sternite 7 with apical margin pointed; tergite 9 with short ventral struts; tergite 10 with some long setae on apex and apical margin with short fringes; sternite 9 with two pairs of long setae and truncate apex. Median lobe of aedeagus with bulbous base in ventral view and not curved apex in lateral view; lateral lobes almost reaching apex of median lobe in lateral view (Fig. 246); internal sclerites as in Fig. 247. Female. Similar to male except for: pair of frontal process less developed than on male; sternite 8 with short setae on apical margin (Fig. 148); ovipositor as in P. andinus. Distribution Bolivia and Peru (Cusco). Remarks The species is excluded from the genus Piestus based on the current phylogenetic study (see ‘Results’, ‘Cladistic analysis’) and here transferred to the genus Hypotelus, with which it shares the following characters: emarginate anterior angles of mentum, pair of conspicuous setae at the apex of the median sclerotised plate of the ligula (Fig. 137), joined gular sutures, Invertebrate Systematics 551 only one complete longitudinal stria on each elytron (Fig. 56), and short setae on apical margin of sternite 8 of female (Fig. 148). Hypotelus laevis differs from H. pusillus and H. andinus by a pair of broad and short pointed frontal processes on the head (Fig. 104) and well-developed dorsal teeth of mandibles, forming a bifurcate apex (Fig. 128). Hypotelus pusillus and H. andinus do not have a pair of frontal processes on the head (Fig. 105), or dorsal teeth on the mandibles (Fig. 129). Biological notes Hypotelus laevis has been collected on rotten palm and by Winkler extraction of leaf litter. Hypotelus andinus (Bernhauer, 1917), comb. nov. (Figs 57, 105, 112, 129, 132, 137, 151, 155, 248–250) Piestus (Antropiestus) andinus Bernhauer, 1917: 45 (original description, type locality: ‘West-Kolumbien: Umgebung von Cali am Rio Cauca’); Scheerpeltz, 1933: 993 (distribution); Scheerpeltz, 1952: 295 (characters, distribution). Piestus andinus: Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1788 (distribution); Newton et al., 2005: 37 (distribution). Piestus (Antropiestus) strigipennis Bernhauer, 1921: 65 (original description, type locality; ‘Bolivien: Yuracaris’); Scheerpeltz, 1933: 993 (distribution); Scheerpeltz, 1952: 295 (characters, distribution). New synonym. Piestus strigipennis: Blackwelder, 1944: 101 (distribution); Herman, 2001b: 1795 (distribution). Type material Piestus (Antropiestus) andinus Bernhauer, 1917. Lectotype deposited in FMNH, male [damaged specimen: without right anterior leg] with labels: (1) ‘<’ [white label, handwritten, together with the specimen]; (2) ‘Columbia occ/ Cali. Fassl’ [white label/printed in black]; (3) ‘Antropiestus/andinus/Brnh. Typus’ [light yellow label, handwritten]; (4) ‘Chicago NHMus/M.Bernhauer/ Collection’ [white label, printed in black]. 1 deposited in NMW with labels: (1) ‘<’ [white label, printed in black]; (2) ‘Columbia occ/Cali. Fassl’ [white label/printed in black]; (3) ‘Piestus (Subg./Antropiestus)’ [white label, handwritten]; (4) ‘andinus Bernh./Cotyp.’ [white label, handwritten]; (5) ‘ex coll./Klima’ [blue label, printed in black]; (6) ‘ex coll./Scheerpeltz’ [light blue label, printed in black]; (7) ‘COTYPUS/Piestus/(Antropiestus)/andinus/ Bernhauer’ [rose label, the first line printed in black, the others handwritten]; (8) ‘andinus/Bh.’ [green label, handwritten]. Paraletotype deposited in NMW, male, with the same labels of lectotype. Note: In the original description Bernhauer (1917) did not specify how many specimens he observed. We received from FMNH and NMW two specimens with type labels, which here we are considering as type material. Piestus (Antropiestus) strigipennis Bernhauer, 1921. Syntype deposited in FMNH, female, with labels: (1) ‘Yuracaris,/A. Fauvel’ (white label, printed in black]; (2) ‘Bolivien/determ.’ [white label, printed in black]; (3) ‘Piestus cavi?-/collis Fauvel?/Mus. Hamburg’ [light blue label, handwritten]; (4) ‘strigipennis/Bernh./Typus.’ [light yellow label, handwritten]; ‘Chicago NHMus/M.Bernhauer/Collections’ [white label, printed in black]. Note: In the original description Bernhauer (1921) did not specify how many specimens he observed. Additional material See Appendix 2. 552 Invertebrate Systematics E. Caron et al. Figs 106–119. 106–108, Piestus sulcatus. 106, Antenna, male; 107, antenna, female; 108, labrum, left epipharynx and right setae removed; 109, P. capricornis, labrum, left epipharynx and right setae removed; 110, P. fronticornis, sp. rev., labrum, left epipharynx and right setae removed; 111, P. surrufus, sp. nov., labrum, left epipharynx and right setae removed; 112, Hypotelus andinus, comb. nov., labrum, left epipharynx and right setae removed. 113–115, Piestus sulcatus, mandibles. 113, Dorsal view, left prostheca removed; 114, right mandible, ventral view; 115, right mandible, lateral view; 116, P. lacordairei, apex of the mandibles, dorsal view, setae and left prostheca removed. 117, 118, Piestus puncticollis, apex of the mandibles, setae and left prostheca removed. 117, Dorsal view; 118, right mandible, lateral view; 119, P. capricornis, apex of the mandibles, setae and left prostheca removed. Scale bar Figs 106, 107, 115, 117, 118 = 0.5 mm; Figs 108–111, 116, 119 = 0.25 mm; Fig. 112 = 0.15 mm. Revision of Piestus with remarks on related genera Invertebrate Systematics Figs 120–137. 120, Piestus spinosus, apex of the mandibles, setae and left prostheca removed; 121, 122, P. fronticornis, sp. rev., apex of the mandibles; 121, dorsal view, left prostheca removed; 122, right mandible, lateral view; 123, P. pennicornis, male, apex of the mandibles, setae and left prostheca removed; 124, P. pennicornis, female, apex of the mandibles, setae and left prostheca removed; 125, P. filicornis, apex of the mandibles, setae and left prostheca removed; 126, P. surrufus, sp. nov., apex of the mandibles, setae and left prostheca removed; 127, P. paradoxus, apex of the mandibles, setae and left prostheca removed; 128, Hypotelus laevis, comb. nov., mandibles, left prostheca removed; 129, Hypotelus andinus, comb. nov., apex of the mandibles, setae and left prostheca removed; 130, P. fronticornis, sp. rev., maxilla; 131, P. sulcatus, maxillary palpus (palpomeres 2–4); 132, H. andinus, maxillary palpus (palpomeres 2–4); 133, P. fronticornis, sp. rev., labium, left setae of the ligula and right setae of the mentum removed; 134, P. sulcatus, mentum, right setae removed; 135, P. acuminatus, sp. nov., mentum, right setae removed; 136, H. laevis, mentum, right setae removed; 137, H. andinus, comb. nov., ligula, right palpus removed. Scale bar Figs 120, 123–130, 133–134 = 0.25 mm; Figs 121, 122 = 0.50 mm; Figs 131, 132, 137 = 0.15 mm; Fig. 136 = 0.10 mm. 553 554 Invertebrate Systematics E. Caron et al. Figs 138–148. 138–142, Piestus sulcatus. 138, Anterior leg, posterior view; 139, anterior leg, anterior view; 140, median leg, posterior view; 141, posterior leg, posterior view; 142, pre-tarsus; 143, P. fronticornis, sp. rev., sternite 1–3; 144, P. acuminatus, sp. nov., sternite 1–3. 145–147, Piestus sulcatus. 145, Tergite 8, male, right setae removed; 146, sternite 8, male, right setae removed; 147, sternite 8, female, right setae removed; 148, Hypotelus laevis, comb. nov., sternite 8, female, right setae removed. Scale bar Figs 138–141, 143, 144 = 0.50 mm; Fig. 142 = 0.10 mm; Figs 145–148 = 0.25 mm. Redescription BL: 6.1 mm, BW: 1.3 mm. Body somewhat flattened dorsoventrally; entirely black with legs a little lighter (Fig. 57). Dorsal integument of head and pronotum with fine punctures and microgranulate sculptures (Fig. 105); elytra with one longitudinal striae finely punctate and closely to internal margin, conspicuous only on basal half; metaventrite and abdomen with fine punctures and microgranulate sculptures. Revision of Piestus with remarks on related genera Invertebrate Systematics 555 Figs 149–155. 149, Piestus sulcatus, tergite 9 and 10, male, right setae removed; 150, P. bicornis, tergite 9 and 10, male, right setae removed; 151, Hypotelus andinus, comb. nov., tergite 10, female, right setae removed. 152, 153, Piestus sulcatus. 152, Sternite 9, male; 153, ovipositor, female; 154, P. fronticornis, sp. rev., ovipositor, female; 155, H. andinus, comb. nov., ovipositor, female. Scale bar Figs 149, 152–154 = 0.25 mm; Fig. 150 = 0.50 mm; Figs 151, 155 = 0.15 mm. Male. Head with front slightly deflected (Fig. 105); Vshaped frontal sulcus incomplete; anterior angles curved and weakly prominent; eyes slightly prominent from above, and line with five moderately sized punctures with long setae on basal half of dorsal margin. Antennae short, almost reaching apex of elytra (Fig. 57); with three long setae near the apex on dorsal face; antennomeres 5–11 entirely with microsetae and some long setae disperse; scape the longest, subequal to antennomere 2 and 3 combined; antennomere 4 shortest and 5–11 somewhat quadrate. Labrum with six setae medially, internal the shortest (Fig. 112); each lateral third, four long setae, two apical and two subapical. Mandibles symmetrical, one acute tooth on each internal border (Fig. 129); without dorsal teeth. Maxillary palpus with palpomere 3 wider than long and 4 longer than 2 and 3 combined (Fig. 132). Mentum 2 times wider than long and anterior angles emarginate; ligula slightly emarginate and two pair of conspicuous long setae on anterior margin, one pair near median sclerite and other on apex of median sclerite (Fig. 137). Gular sutures joined. Pronotum wider than long (PW/PL = 1.25) (Fig. 105); anterior angles not projected; longitudinal median sulcus conspicuous; conspicuous and deep large depression at middle; gradual constriction at basal third; apical two-thirds with slightly curved sides. Prosternum with anterior margin truncate. Elytra somewhat longer than wide (EL/BW = 1.21), covering tergite 3; when together slightly emarginate on the middle; apical margin with short setae. Abdominal segments 3–6 with parallel sides; segments 3–6 with two pair of paratergites, segment 7 with only one pair; sternite 8 with apical margin truncate; tergite 9 with short ventral struts; tergite 10 with 556 Invertebrate Systematics Figs 156–171. Aedeagus, for each species with lateral view and apex in dorsal view respectively. 156–157, Piestus convexus, sp. nov.; 158–159, P. similis, sp. nov.; 160–161, P. lacordairei; 162–163, P. heterocephalus; 164–165, P. puncticollis; 166–167, P. capricornis; 168–169, P. planatus; 170–171, P. spinosus. Scale bar = 0.25 mm. E. Caron et al. Revision of Piestus with remarks on related genera Invertebrate Systematics Figs 172–189. Aedeagus. 172–175, Piestus longipennis. 172, Lateral view; 173, apex in dorsal view; 174, apex in ventral view; 175, apex in lateral view (everted internal sac). 176–178, Piestus zischkai. 176, Lateral view; 177, apex in dorsal view; 178, apex in ventral view. 179–181, Piestus bicornis. 179, Lateral view; 180, apex in dorsal view; 181, ventral view. 182–183, Piestus fronticornis, sp. rev. 182, Lateral view; 183, apex in dorsal view. 184– 185, Piestus validus. 184, Lateral view; 185, apex in dorsal view. 186–187, Piestus rossii, sp. nov. 186, Lateral view; 187, apex in dorsal view. 188–189, Piestus pennicornis. 188, Lateral view; 189, apex in dorsal view. Scale bar Figs 172, 176, 179, 181, 182, 184 = 0.50 mm; Figs 173–175, 177, 178, 180, 183, 185, 186–189 = 0.25 mm. 557 558 Invertebrate Systematics E. Caron et al. Figs 190–205. Aedeagus, for each species with lateral view and apex in dorsal view respectively. 190–191, Piestus pygmaeus; 192–193, P. extimus; 194–195, P. buquetii; 196–197, P. filicornis; 198–199, P. abdominalis, sp. nov.; 200–201, P. niger; 202–203, P. minutus; 204–205, P. penicillatus. Scale bar = 0.25 mm. some long setae on apex and apical margin with short fringes, basal half not divided longitudinally; sternite 9 with two pairs of long setae and truncate apex. Median lobe of aedeagus with bulbous base in ventral view and somewhat globose apex in lateral view, tube closed in dorsal view; lateral lobes exceed little apex of median lobe and curved on apex in lateral view (Figs 248, 249); internal sclerites as in Fig. 250. Female. Similar to male except for: elytra a little shorter, covering only basal half of tergite 3; abdominal sternite 8 with Revision of Piestus with remarks on related genera Invertebrate Systematics Figs 206–223. Aedeagus. 206–207, Piestus termitis, sp. nov. 206, Lateral view; 207, apex in dorsal view. 208–209, Piestus surrufus, sp. nov. 208, Lateral view; 209, apex in dorsal view. 210–211, Piestus formicinus. 210, Lateral view; 211, dorsal view. 212–214, Piestus sulcatus. 212, Lateral view; 213, apex in dorsal view; 214, ventral view. 215–216, Piestus gounellei. 215, Lateral view; 216, apex in dorsal view; 217, apex in ventral view. 218–219, Piestus mexicanus. 218, Lateral view; 219, apex in dorsal view. 220–221, Piestus ecuadorensis, sp. nov. 220, Lateral view; 221, apex in dorsal view. 222–223, Piestus foveolatus, sp. nov. 222, Lateral view; 223, apex in dorsal view. Scale bar = 0.25 mm. 559 560 Invertebrate Systematics Figs 224–239. Aedeagus, for each species with lateral view and apex in dorsal view respectively. 224–225, Piestus sulcipennis; 226–227, P. imperfectus, sp. nov.; 228–229, P. costatus; 230–231, P. chiriquensis; 232–233, P. nevermanni; 234–235, P. paradoxus; 236–237, P. boliviensis, sp. nov.; 238–239, P. aper. Scale bar = 0.25 mm. E. Caron et al. Revision of Piestus with remarks on related genera Invertebrate Systematics 561 Figs 240–250. 240–241, Piestus angularis, aedeagus. 240, Lateral view; 241, apex in dorsal view. 242–243, Piestus acuminatus, sp. nov. 242, Lateral view; 243, apex in dorsal view. 244–245, Piestus rugosus. 244, Lateral view; 245, apex in dorsal view. 246–247, Hypotelus laevis, comb. nov. 246, Lateral view; 247, dorsal view. 248–250, Hypotelus andinus, comb. nov. 248, Lateral view; 249, dorsal view; 250, ventral view. Scale bar = 0.25 mm. pointed apex and short setae on apical margin; tergite 10 divided longitudinally on basal half (Fig. 151); ovipositor consisting of pair of weakly pigmented hemisternites and pair of more apical coxites, and with many long setae on apex (Fig. 155). Spermatheca unknown. Distribution Colombia (Boyacá), Ecuador and Bolivia. Biological notes Hypotelus andinus has been collected in a Malaise trap. Remarks Hypotelus andinus is excluded from the genus Piestus based on the current phylogenetic study (see ‘Results’, ‘Cladistic analysis’) and here transferred to the genus Hypotelus, with which it shares the same characters cited above (see remarks under H. laevis). The species H. andinus differs from H. pusillus by having a pronotum with a conspicuous and deep large depression at the middle (Fig. 105) and by tergite 10 of the female being longitudinally divided on basal half (Fig. 151). Hypotelus andinus differs from H. laevis by characters listed above under remarks for H. laevis. 562 Invertebrate Systematics E. Caron et al. Figs 251–268. Spermatheca. 251, Piestus lacordairei; 252, P. heterocephalus; 253, P. capricornis; 254, P. planatus; 255, P. spinosus; 256, P. longipennis; 257, P. bicornis; 258, P. fronticornis, sp. rev.; 259, P. validus; 260, P. pennicornis; 261, P. pygmaeus; 262, P. extimus; 263, P. buquetii; 264, P. filicornis; 265, P. abdominalis, sp. nov.; 266, P. niger; 267, P. minutus; 268, P. penicillatus. Scale bar = 0.25 mm; except Fig. 257 = 0.50 mm. Revision of Piestus with remarks on related genera Invertebrate Systematics Figs 269–287. Spermatheca. 269, Piestus termitis, sp. nov.; 270, P. surrufus, sp. nov.; 271, P. formicinus, sp. nov.; 272, P. sulcatus; 273, P. gounellei; 274, P. mexicanus; 275, P. ecuadorensis, sp. nov.; 276, P. foveolatus, sp. nov.; 277, P. sulcipennis; 278, P. imperfectus, sp. nov.; 279, P. costatus; 280, P. chiriquensis; 281, P. nevermanni; 282, P. paradoxus; 283, P. aper; 284, P. angularis; 285, P. acuminatus, sp. nov.; 286, P. rugosus; 287, P. elegans, sp. nov. Scale bar = 0.25 mm. 563 564 Invertebrate Systematics Subfamily OSORIINAE Erichson, 1839 Tribe ELEUSININI Sharp, 1887 Eleusis Laporte, 1835 Eleusis Laporte, 1835: 131. For review of literature of the genus and a complete list of synonyms, see Herman (2001b). Lissopiestus Scheerpeltz, 1952: 295 (as subgenus of Piestus); Herman, 2001b: 1788 (subgenus of Piestus). Type species. Isomalus interruptus Erichson, 1840 (fixed by original designation and monotypy). New synonym. Remarks Lissopiestus, previously considered as a subgenus of Piestus, is here synonymised with Eleusis, as its type species P. (L.) interruptus (Erichson, 1840) is transferred to Eleusis based on morphological study of the holotype (see remarks under Eleusis interrupta). Eleusis interrupta (Erichson, 1840), comb. rest. (restored combination) (Figs 58, 103) Isomalus interruptus Erichson, 1840: 839 (original description, type locality: ‘Carthagenae in Columbia’); Fauvel, 1864: 35 (characters, distribution). Eleusis interrupta: Bernhauer & Schubert, 1910: 7 (distribution); Newton et al. 2005: 29 (distribution). Piestus (Piestus) interruptus: Scheerpeltz, 1933: 994 (distribution). Piestus interruptus: Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1791 (distribution). Piestus (Lissopiestus) interruptus: Scheerpeltz, 1952: 295 (characters, distribution). Type material Syntype deposited in ZMHB, female, with labels: (1) ‘6819’ [white label, printed in black]; (2) ‘Type’ [red label, printed in black]; (3) ‘interruptus/E./Carthag. Gory’ [green label, handwritten]; (3) ‘Eleusis/ Cast./1835’ [white label, handwritten]; (4) ‘Zoll. Mus/Berlin’ [white label, printed in black]; (5) ‘Holotypus/Isomalus/interruptus Erichson, 1840/ labelled by MNHUB 2008’ [red label, printed in black]. Note: In the original description Erichson (1840) did not specify how many specimens he observed. Distribution Brazil, Colombia, French Guiana, Ecuador, Peru, Bolivia and Argentina (Herman 2001b; Newton et al. 2005). Remarks The specific name ‘interruptus’ is an adjective and is changed again to the feminine spelling to agree with the gender of the generic name (Article 34.2; ICZN 1999). The species is transferred again to the genus Eleusis based on: head with slight groove on dorsal edge of eye, pronotum strongly narrowed at base and wider than long (Fig. 58), and abdomen without paratergites, each abdominal segment with only a membranous suture between tergum and sternum (Fig. 103). The syntype of Eleusis interrupta is very similar to specimens of E. humilis identified and deposited in FMNH. However, some E. Caron et al. specimens were dissected and some slight differences were observed in aedeagi of those specimens, suggesting that a careful morphological study is necessary before any conclusion about the specific identity of Eleusis interrupta. Eleusis interrupta was not used in the present cladistic analysis because there is no topological correspondence in character states among some observed characters. Species Inquirendae (uncertain status) Piestus longicornis (Lacordaire, 1833) Zirophorus longicornis Lacordaire, 1833: 65 (original description, type locality: ‘Cayenne’). Piestus longicornis: Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1792 (distribution). Type material Not found. Distribution French Guiana (Herman 2001b). Biological notes Unknown (Lacordaire 1833). Remarks Lacordaire (1833) described briefly P. longicornis based only on some characters from the adult and from one larva. The brief adult description (flat body, red elytra and long antennae) and the type locality (Cayenne, French Guiana) of P. longicornis suggest this species is similar to P. spinosus. However, the type material of P. longicornis was not found and its taxonomic status is still doubtful. Piestus longicornis is listed as a valid species only in Blackwelder (1944) and Herman (2001b), but has not been otherwise used since its original description, and thus it should be considered a nomen dubium. Piestus fulvipes Erichson, 1840 Piestus fulvipes Erichson, 1840: 833 (original description, type locality: ‘Guadeloupe’); Fauvel, 1864: 23 (characters, distribution); Blackwelder, 1943: 45 (characters, distribution, notes); Blackwelder, 1944: 100 (distribution); Herman, 2001b: 1791 (distribution). Piestus (Piestus) fulvipes: Bernhauer & Schubert, 1910: 7 (distribution); Scheerpeltz, 1952: 288 (characters, distribution). Type material Not examined. Syntype deposited in BMNH, sex undetermined ‘One example in the British Museum from the Chevrolat collection. Collected by F. de L’Herminier’ (Blackwelder, 1943: 46). Note: ‘One syntype labelled as Type by D. Sharp from the Chevrolat collection’, with label: ‘Piestus fulvipes Er. Type ex. coll. Chevrolat. Guadeloupe L’Herminier.’ and deposited in BMNH (Dr Roger Booth, BMNH, pers. comm.). In the original description Erichson (1840) did not specify how many specimens he observed. Distribution West Indies (Herman 2001b). Revision of Piestus with remarks on related genera Biological notes Unknown (Blackwelder 1943). Remarks All specimens previously identified in collections as P. fulvipes were considered as P. penicillatus. Blackwelder (1943) redescribed P. fulvipes and P. penicillatus, noting a little difference on pronotum integument between the two species. As the type material of P. fulvipes was not observed in the current study, the species is placed as uncertain status, but we suggest that it is a junior synonym of P. penicillatus. Acknowledgements We would like to thank Sonia A. Casari (MZSP), José Ricardo M. Mermudes (Universidade Federal do Rio de Janeiro, UFRJ, Brazil), Albino M. Sakakibara (UFPR), Lucia M. de Almeida (UFPR) and three anonymous reviewers for valuable suggestions on the manuscript. We are also grateful to Margaret K. Thayer (FMNH), Torsten Dikow (FMNH), Dave J. Clarke (FMNH) and Angélico Asenjo (UFPR) for providing helpful comments that improved this research. We thank M. K. Thayer for the use of the Microptics imaging system provided by US National Science Foundation collaborative grant, AToL: Assembling the Beetle Tree of Life (EF-0531768, subcontract 144–439) and the TAXon line – ‘Rede Paranaense de Coleções Biológicas (UFPR) for the photos. This research was supported by MCT/CNPq Proc.no. 477520/2007-3, 473847/2009-4 and ‘Conselho Nacional de Desenvolvimento Científico e Tecnológico’. Finally, we are grateful to curators and institutions listed for loan of types and other specimens used in this study. This paper is part of the Ph.D. project of the first author, and contribution n
1864 of the Departamento de Zoologia, Universidade Federal do Paraná, Brazil. References Bernhauer, M. (1906). Neue Staphyliniden aus Südamerika. Deutsche Entomologische Zeitschrift 1906, 193–202. Bernhauer, M. (1917). Neue arten der gattungen Piestus, Leptochirus und Conosoma aus Südamerika. Neue Beiträge zur Systematischen Insektenkunde 1, , 45–53. Bernhauer, M. (1920). Results of Dr. E. Mjöberg’s Swedish scientific expeditions to Australia 1910–1913. 22. Staphylinidae. Arkiv für Zoologi 13(8), 1–27. Bernhauer, M. (1921). Zur Staphylinidenfauna von Südamerika. Deutsche Entomologische Zeitschrift 1921, 65–77. Bernhauer, M. (1928). 33ster Beitrag zur südamerikanischen Staphylinidenfauna. Tijdschrift voor Entomologie 71, 286–288. Bernhauer, M., and Schubert, K. (1910). Staphylinidae I. In ‘Coleopterorum Catalogus, Vol. 5’. (Ed. S. Schenkling.) pp.1–86. (Junk: Berlin.) Blackwelder, R. E. (1942). Notes on the classification of the staphylinid beetles of the groups Lispini and Osoriinae. Proceedings of the United States National Museum 92, 75–90. Blackwelder, R. E. (1943). Monograph of the West Indian beetles of the family Staphylinidae. United States National Museum Bulletin 182, viii + 1–658. Blackwelder, R. E. (1944). Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 1. United States National Museum Bulletin 185, xii + 1–188. Blackwelder, R. E. (1952). The generic names of the beetle family Staphylinidae, with an essay on genotypy. United States National Museum Bulletin 200, iv + 1–483. Bremer, K. (1994). Branch support and tree stability. Cladistics 10, 295–304. doi:10.1111/j.1096-0031.1994.tb00179.x Invertebrate Systematics 565 Cameron, M. (1927). Description of a new species of Siagonium from New Zealand. Entomologist’s Monthly Magazine 63, 222. Caron, E., Ribeiro-Costa, C. S., and Newton, A. F. (2008). New position of an abdominal defensive gland complex in Staphylinidae (Coleoptera) with redescription of Piestus heterocephalus Fauvel, 1902 (Piestinae). Zootaxa 1895, 1–9. Crotch, G. R. (1870). The genera of Coleoptera studied chronologically (1802–21). The Transactions of the Entomological Society of London 1870, 213–241. Dalman, J. W. (1821). Nya genera och species af Insecter. Kongelige Vetenskaps Academiens Handlingar 1821, 372–387. Erichson, W. F. (1839). Genera et species Staphylinorum insectorum coleopterorum familiae. F. H. Morin, Berlin, pp. 1–400. Erichson, W. F. (1840). Genera et species Staphylinorum insectorum coleopterorum familiae. F. H. Morin, Berlin, pp. 401–954. Fabricius, J. C. (1801). ‘Systema Eleutheratorum Secundum Ordines, Genera, Species: Adiectis Synonymis, Locis, Observationibus, Descriptionibus. Vol. 2.’ (Bibliopolii Academici Novi: Kiliae.) pp. 687. Fauvel, A. (1864). Études sur les Staphylinides de l’Amérique centrale, principalement du Mexique. Notices Entomologiques 2, 3–62. Fauvel, A. (1865). Études sur les Staphylinides de l’Amérique centrale, principalement du Mexique. Bulletin de la Société Linnéenne de Normandie 9, 8–66. Fauvel, A. (1895). Staphylinides nouveaux de l’Inde et de la Malaisie. Revue d’Entomologie 14, 180–286. Fauvel, A. (1901). Voyage de M. le Dr. Ed. Bugnion au Venezuela, en Colombie et aux Antilles. Revue d’Entomologie 20, 69–91. Fauvel, A. (1902). Staphylinides exotiques nouveaux. Revue d’Entomologie 21, 8–37. Fitch, W. N. (1971). Toward defining the course of evolution: minimum change for a specific tree topology. Systematic Zoology 20, 406–416. doi:10.2307/2412116 Fleming, J. (1821). Insecta. In ‘Supplement to the Fourth, Fifth, and Sixth Editions of the Encyclopedia Britannica, with Preliminary Dissertations on the History of the Sciences. Vol. 5’. pp. 41–56. (Archibald Constable and Company: Edinburgh.) Fleutiaux, E., and Sallé, A. (1889). Liste des Coléoptères de la Guadeloupe et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France 9, 351–484. Goloboff, P. A. (1993). Estimating character weights during tree search. Cladistics 9, 83–91. doi:10.1111/j.1096-0031.1993.tb00209.x Goloboff, P. A., Farris, J. S., and Nixon, K. C. (2008). TNT, a free program for phylogenetic analysis. Cladistics 24, 774–786. doi:10.1111/j.1096-0031. 2008.00217.x Gravenhorst, J. L. C. (1806). ‘Monographia Coleopterorum Micropterorum.’ (Henricus Dieterich: Gottingae [Göttingen, Germany].) pp. 248. Gray, G. (1832). [Notices of new genera and species]. In ‘The Class Insecta Arranged by the Baron Cuvier, with Supplementary Additions to Each Order’. (Eds E. Griffith and E. Pidgeon.) Insecta, 1: viii + 570. In: E. Griffith, The Animal kingdon arranged in conformity with its organization, by the Baron Cuvier . . . with supplementary additions to each order. Vol. 14. (Whittaker, Treacher, and Co.: London.) Grebennikov, V., and Newton, A. F. (2009). Good-bye Scydmaenidae, or why the ant-like stone beetle should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal of Entomology 106, 275–301. Guérin-Méneville, F. E. (1829). ‘Iconographie de Règne Animal de G. Cuvier, ou Représentation d’Après Nature de l’Une des Espèces les Plus Remarquables et Souvent non Encore Figurées, de Chaque Genre d’Animaux. Avec un Texte Descriptif Mis au Courant de la Science. Ouvrage Pouvant Servir d’Atlas a Tous les Traites de Zoologie. II. Planches des Animaux Invertébrés. Insectes, pls. 3–12.’ (J. B. Baillière: Paris.) 566 Invertebrate Systematics Gusarov, V. I. (2002). A revision of Nearctic species of the genus Geostiba Thomson, 1858 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 81, 1–88. Herman, L. H., Jr (1970). Phylogeny and reclassification of the genera of the rove-beetle subfamily Oxytelinae of the world (Coleoptera, Staphylinidae). Bulletin of the American Museum of Natural History 142(5), 343–454. Herman, L. H., Jr (1977). Revision and phylogeny of Zalobius, Asemobius, and Nanobius, new genus (Coleoptera, Staphylinidae, Piestinae). Bulletin of the American Museum of Natural History 159(2), 45–86. Herman, L. H. (2001a). Nomenclatural changes in the Staphylinidae (Insecta: Coleoptera). Bulletin of the American Museum of Natural History 264, 1–82. doi:10.1206/0003-0090(2001)264<0003:NCITSI> 2.0.CO;2 Herman, L. H. (2001b). Catalog of the Staphylinidae (Insecta: Coleoptera) 1758 to the end of the second millennium, parts I–VII. Bulletin of the American Museum of Natural History 167, 1–4218. doi:10.1206/00030090(2001)264<0003:NCITSI>2.0.CO;2 ICZN (1999). International Code of Zoological Nomenclature, fourth edition [online]. The International Trust for Zoological Nomenclature 1999 c/o The Natural History Museum, Cromwell Road, London, UK (last updated 01 January 2000). Available from http://www.iczn.org/iczn/ index.jsp [Verified 26 January 2009.] Khachikov, E. A. (2007). A new species of the genus Siagonium Kirby et Spence, 1815 (Coleoptera, Staphylinidae) from China. Kavkazskii Entomologicheskii Byulleten; Caucasian Entomological Bulletin 3, 103–104. Kirby, W., and Spence, W. (1815). ‘An Introduction to Entomology: or Elements of the Natural History of Insects: with Plates. 1.’ (Longman, Hurst, Rees, Orme, and Brown: London.) Kraatz, G. (1859). Die Staphylinen-Fauna von Ostindien, insbesondere der Insel Ceylan. Archiv für Naturgeschichte 25(1), 1–196. Lacordaire, J. T. (1833). Essai sur les coléoptères de la Guyane Française. Nouvelles Annales du Muséum d’Histoire Naturelle 2, 35–94. Lacordaire, J. T. (1854). ‘Histoire des Insectes. Genera des Coléoptères ou Exposé Méthodique et Critique de Tous les Genres Proposés Jusqu’ici Dans Cet Ordre d’Insectes. 2.’ (Librairie Encyclopédique de Roret: Paris.) Laporte, F. L. (Castelnau) (1835). ‘Études Entomologiques, ou Description d’Insectes Nouveaux, et Observations sur Leur Synonymie.’ (Méquignon-Marvis: Paris.) pp. 159. Latreille, P. A. (1807). ‘Genera Crustaceorum et Insectorum, Secundem Ordinem Naturalem in Familias Disposita, Iconibus Exemplisque Plurimus Explicata. Vol. 2.’ (Amand Koenig: Paris.) pp. 280. Latreille, P. A. (1829). Crustacés, arachnides et partie des insectes. In ‘Le Règne Animal Distribué d’Après son Organisation, Pour Servir de Base à l’Histoire Naturelle des Animaux et d’Introduction à l’Anatomie Comparée. par M. le Baron Cuvier. . . . Crustacés, Arachnides et Partie des Insectes. 4.’ (Ed. G. Cuvier.) pp. xxvii + 1–584. (Déterville: Paris.) Lawrence, J. F., and Newton, A. F., Jr (1982). Evolution and classification of beetles. Annual Review of Ecology and Systematics 13, 261–290. doi:10.1146/annurev.es.13.110182.001401 LeConte, J. L. (1866). Additions to the coleopterous fauna of the United States. No. 1. Proceedings. Academy of Natural Sciences of Philadelphia 19, 361–394. Leschen, R. A. B., and Löbl, I. (1995). Phylogeny of Scaphidiinae with redefinition of tribal and generic limits (Coleoptera: Staphylinidae). Revue Suisse de Zoologie 102, 425–474. Leschen, R. A. B., and Löbl, I. (2005). Phylogeny and classification of Scaphisomatini (Staphylinidae: Scaphidiinae) with notes on mycophagy, termitophily, and functional morphology. Coleopterists Society Monographs, Patricia Vaurie Series 3, 1–63. E. Caron et al. Naomi, S.-I. (1987–1990). Comparative morphology of the Staphylinidae and the allied groups (Coleoptera, Staphylinoidea), parts I–XI. Kontyû [1987–88]. Japanese Journal of Entomology [1989–90], 55, 450–458, 666–675; 56, 67–77, 241–250, 506–513, 727–738; 57, 82–90, 269–277, 517–526, 720–733; 58, 16–23. Naomi, S.-I. (1995). Revision of the genus Piestoneus Sharp (Coleoptera: Staphylinidae) from Japan. Japanese Journal of Entomology 63(4), 763–780. Naomi, S.-I. (2006). Description of a new species of the genus Siagonium Kirby et Spence from Japan (Coleoptera, Staphylinidae, Piestinae). Natural History Research 9(1), 41–44. Navarrete-Heredia, J. L., Newton, A. F., Thayer, M. K., Ashe, J. S., and Chandler, D. S. (2002). ‘Guía Ilustrada de los Géneros de Staphylinidae (Coleoptera) de México – Illustrated Guide to the Genera of Staphylinidae (Coleoptera) of Mexico.’ (Universidad de Guadalajara y CONABIO: México.) Newton, A. F., Jr (1982a). Redefinition, revised phylogeny, and relationships of Pseudopsinae (Coleoptera, Staphylinidae). American Museum Novitates 2743, 1–13. Newton, A. F., Jr (1982b). A new genus and species of Oxytelinae from Australia, with a description of its larva, systematic position, and phylogenetic relationships (Coleoptera, Staphylinidae). American Museum Novitates 2744, 1–24. Newton, A. F., and Peck, S. B. (2006). Family Staphylinidae, the rove beetles. In ‘The Beetles of the Galápagos Islands, Ecuador: Evolution, Ecology and Diversity (Insecta: Coleoptera)’. (Ed. S. B. Peck.) pp. 131–158. (NRC Research Press: Ottawa, Ontario, Canada.) Newton, A. F., Thayer, M. K., Ashe, J. S., and Chandler, D. S. (2000). 22. Staphylinidae. In: American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. vol. 1. (Eds R. H. Arnett and M. C. Thomas.) pp. 272–418. (CRC Press: Boca Raton, London, New York, Washington.) Newton, A. F., Chacón, C. G., and Chandler, D. S. (2005). Checklist of the Staphylinidae (Coleoptera) of Colombia. Biota Colombiana 6, 1–72. Nixon, K. C. (1999). Winclada (BETA), ver. 0.9.9. (Published by the author, Ithaca, New York.) Nixon, K. C., and Carpenter, J. M. (1993). On outgroups. Cladistics 9, 413–426. doi:10.1111/j.1096-0031.1993.tb00234.x Olivier, A. G. (1811). Insectes. In ‘Encyclopédie Méthodique. Histoire Naturelle. Vol. 8(2).’ pp. 361–722. (H. Agasse: Paris.) Page, R. (2001). NDE (NEXUS Data Editor For Windows) version 0.5.0. (Published by the author, Glasgow, United Kingdom.) Perty, J. A. M. (1830). Insecta Brasiliensia. In ‘Delectus Animalium Articulatorum, Quae in Itinere per Brasiliam Annis MDCCCXVII–MDCCCXX Jussu et Auspiciis Maximiliani Josephi I. Bavariae Regis Augustissimi Peracto. . . . Frid.’ (Eds J. Spix and C. Martius.) pp. 1–60. (Fleischer: Monachii.) Reiche, L. (1865). Étude des espèces de Mylabrides de la collection de L. Reiche, suivie d’une note sur le genre Trigonurus Mulsant et description d’une espèce nouvelle. Annales de la Société Entomologique de France 5, 627–642. Scheerpeltz, O. (1933). Staphylinidae VII. In ‘Coleopterorum Catalogus, 6 (129).’ (Ed. S. Schenkling.) pp. 989–1500. (Junk: Berlin.) Scheerpeltz, O. (1951). Los Staphylinidae (Coleoptera) encontrados en Bolivia por el Sr. Rodolfo Zischka. Folia Universitaria 5(5), 1–16. Scheerpeltz, O. (1952). Revision der Gattung Piestus Gravh. (Coleoptera Staphylinidae). Revista Chilena de Entomologia 2, 281–305. Scheerpeltz, O. (1960). Zur Kenntnis neotropischer Staphyliniden (Col.). Beiträge zur Neotropischen Fauna 2(2), 65–138. doi:10.1080/ 01650526009380624 Sharp, D. (1876). Contribution to an insect fauna of the Amazon Valley. Coleoptera-Staphylinidae. Transactions of the Entomological Society of London 1876, 27–424. Revision of Piestus with remarks on related genera Sharp, D. (1887). Staphylinidae. In ‘Biologia Centrali-Americana. Insecta. Coleoptera. 1(2).’ (Ed) pp. 673–824. (Taylor & Francis: London.) Sharp, D. (1889). The Staphylinidae of Japan. Annals & Magazine of Natural History 3, 28–44, 108–121, 249–267, 319–334, 406–419, 463–476. Sharp, D., and Muir, F. A. G. (1912). The comparative anatomy of the male genital tube in Coleoptera. Transactions of the Entomological Society of London 3, 477–642. Solsky, S.M. (1872). Enumération et description des coléoptères de la famille des Staphylinides recueillis par Mrs. C. Jelsky et le Baron de Nolken pendant leurs voyages dans l’Amérique du Sud en 1870 et 1871. Horae Societatis Entomologicae Rossicae 8, 289–314. Steel, W. O. (1950). Notes on Staphylinidae, chiefly from New Zealand. Transactions of the Royal Society of New Zealand 78, 203–212, 213–235. Thayer, M. K. (2005). 11. Staphylinoidea. 11.7. Staphylinidae Latreille, 1802. Coleoptera, Vol. I. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). In ‘Handbook of Zoology Vol. IV, Arthropoda: Insecta.’ (Coleoptera Eds R.G. Beutel and R. A. B. Leschen; Insecta Eds N. P. Kristensen and R. G. Beutel.) pp. 296–344. (De Gruyter: Berlin, New York.) Invertebrate Systematics 567 Tikhomirova, A. L. (1968). Zhuki-stafilinidy Iury Karatau. In ‘Iurskie Nasekomye Karatau. Akademiia Nauk SSSR, Otdelenie obshei biologii. (Ed. B. B. Rodendorf.) pp. 139–154. (Izdatel’stvo Nauka, Moskva [Moscow].) Wendeler, H. (1955). Neue Staphyliniden aus Brasilien (1. Teil). Dusenia 6 (5), 187–198. Wu, J., and Zhou, H.-Z. (2007). Phylogenetic analysis and reclassification of the genus Priochirus Sharp (Coleoptera: Staphylinidae: Osoriinae). Invertebrate Systematics 21, 73–107. doi:10.1071/IS06010 Zheng, Fa-ke (2004). Coleoptera: Staphylinidae. In ‘Insects of the Great Yarlung Zangbo Canyon of Xizang, China.’ (Ed. Yang, Xing-ke.) pp. 50–60. (China Science and Technology Press: Beijing.) 568 Invertebrate Systematics E. Caron et al. Appendix 1. List of characters Seventy parsimony-informative characters were selected, 62 binary and 10 multistate Head: 0. Front: (0) without pair of frontal processes (Fig. 73); (1) with pair of frontal processes (Fig. 72). 1. Length of the frontal processes in male: (0) equal or longer than the antennal scape (Fig. 65); (1) shorter than the antennal scape (Fig. 64). 2. Basal distance between each frontal process: (0) 2 times or more the basal width of each one (Fig. 66); (1) the same or narrower than the basal width of each one (Fig. 67). 3. Front, without frontal process: (0) slightly ventrally deflected (Fig. 86); (1) suddenly ventrally deflected (Fig. 82). 4. Front in frontal view: (0) without a slight concavity at middle; (1) with a slight concavity at middle. 5. V-shaped frontal sulcus: (0) incomplete (Fig. 86); (1) complete (Fig. 82). 6. Vertex: (0) without transverse carina on base (Fig. 86); (1) with transverse carina on base (Fig. 82). 7. Punctures: (0) fine and inconspicuous (Fig. 82); (1) fine and conspicuous (Fig. 86); (2) large (Fig. 89). 8. Moderate punctures with seta on basal half of the dorsal margin of the eyes: (0) one; (1) two; (2) three (in line); (3) five or more (in line). 9. Antennal length of male: (0) equal or slightly longer than female; (1) conspicuously longer than female (Figs 48, 49, 106, 107). 10. Antennal length of male: (0) short, reaching or exceeding a little the apex of elytra (Fig. 13); (1) long, almost reaching or reaching the apex of abdomen (Fig. 17). 11. Long setae on basal half of exposed dorsal face of the antennal scape: (0) absent; (1) present only in male (Figs 106, 107); (2) present in male and female. 12. Enlarged area on basal half of exposed dorsal face of the antennal scape: (0) absent; (1) present only in male (Figs 106, 107); (2) present in male and female. 13. Antennomere 4: (0) with dispersed microsetae, not densely covered; (1) densely covered with microsetae (Fig. 106). 14. Antenna (antennomere 4 to 11): (0) the same width from the base to the apex (Fig. 18); (1) gradually narrowing to the apex (Fig. 35). 15. Epipharynx as visible dorsally: (0) short, about the same or up to 2 times the median length of labrum (Fig. 108); (1) long, ~2.5 times or so the median length of labrum (Fig. 110). 16. Epipharynx as visible dorsally: (0) with one lobe at each side (Fig. 108); (1) with two lobes at each side (Fig. 111). 17. Mandibles: (0) weakly projected anteriorly, shorter than or equal to the antennal scape (Fig. 26); (1) strongly projected anteriorly, longer than the antennal scape (Fig. 25). 18. Dorsal tooth of the mandibles: (0) absent (Fig. 126); (1) present (Fig. 125). 19. Mandibles with dorsal tooth: (0) weakly developed, or developed but not forming bifurcate apex with first ventral tooth (Figs 113, 115, 123); (1) developed, forming bifurcate apex with first ventral tooth (Figs 121, 122). 20. Mandibles with bifurcate apex: (0) dorsal tooth shorter than the ventral tooth (Fig. 121); (1) dorsal tooth longer than the ventral tooth (Fig. 120). 21. Teeth on internal borders of the mandibles (left–right): (0) 0–0 (Fig. 126); (1) 1–1 or 1–2, right mandible with an area that resembles a tooth (Figs 125, 129); (2) 1–2, right mandible with the apical tooth acute and not close to the other (Fig. 121); (3) 2–2, basal teeth projected anteriorly (Fig. 127); (4) 1–0. 22. Openings of glands at the external margin of mandibles: (0) absent; (1) present (Fig. 114). 23. Small basal tooth on external margin of the mandibles: (0) absent (Fig. 128); (1) present (Fig. 113). 24. Small basal tooth on external margin of the mandibles: (0) weakly acute (Fig. 121); (1) strongly acute (Fig. 113). 25. Maxillary palpus: (0) palpomere 4 equal to or a little longer than 2 and 3 combined (Fig. 131); (1) palpomere 4 shorter than 2 and 3 combined (Fig. 130). 26. Maxillary palpomere 3: (0) wider than long (Fig. 132); (1) as wide as long (Fig. 131). 27. Mentum: (0) rectangular, 2 times as wide as long (Fig. 136); (1) subquadrate, 1.5 times as wide as long or as wide as long (Fig. 135). 28. Anterior angles of the mentum: (0) not emarginate (Fig. 135); (1) emarginate (Fig. 136). 29. Pair of conspicuous setae at the apex of the median sclerotised plate of the ligula: (0) absent (Fig. 133); (1) present (Fig. 137). 30. Pair of conspicuous setae on apical margin of the ligula, near the median sclerotised plate: (0) absent (Fig. 133); (1) present (Fig. 137). 31. Gular sutures: (0) joined; (1) narrowly separated (Fig. 8); (2) broadly separated. Thorax: 32. Anterior angles of the pronotum: (0) not or weakly anteriorly projected (Fig. 61); (1) anteriorly projected (Fig. 65). 33. Sides of the pronotum, anterior to constriction: (0) curved (Fig. 87); (1) subparallel to parallel (Fig. 83); (2) sinuous, with short teeth (Fig. 98). 34. Punctures on pronotum: (0) fine and inconspicuous, sometimes with a few scattered moderate punctures (Fig. 83); (1) fine and conspicuous, sometimes with a few large punctures (Fig. 86); (2) only large punctures (Fig. 91). 35. Median longitudinal sulcus on pronotum: (0) inconspicuous (Fig. 92); (1) conspicuous (Fig. 82). 36. Basal constriction of the pronotum: (0) present on basal half or third (Figs 7, 85); (1) present on basal quarter (Fig. 82). 37. Slight emargination in each side, anterior to basal constriction of the pronotum (in lateral view): (0) absent; (1) present (Fig. 65). 38. Pronotum with lateral tooth anterior to the constriction: (0) absent (Fig. 61); (1) present (Fig. 7). 39. Anterior margin of the prosternum: (0) truncate; (1) anteriorly projected at the middle (Fig. 8). 40. Conspicuous ovate set of fine punctures in median region of prosternum of the male: (0) absent; (1) present (Fig. 8). 41. Mesoventrite process: (0) not carinate; (1) carinate (Fig. 9). 42. Moderate punctures on median region of metaventrite: (0) absent; (1) present. 43. Groove on ventral margin of the anterior femur: (0) absent; (1) present (Fig. 139). 44. Stout setae at the groove on ventral margin of the anterior femur: (0) short (Fig. 139); (1) long. 45. Small acute tooth on each basolateral angle of elytra: (0) absent (Fig. 56); (1) present (Fig. 52). 46. Complete longitudinal striae on each elytron: (0) one (Fig. 56); (1) three; (2) five (Fig. 16); (3) six (Fig. 5). 47. Stria 6 at the basal half on each elytron: (0) absent; (1) present (Fig. 5). 48. Epipleural carina of elytra: (0) absent; (1) present (Fig. 6). 49. Punctures on striae of elytra: (0) absent or inconspicuous (Fig. 16); (1) fine (Fig. 19); (2) large (Fig. 55). 50. Integument of striae of elytra: (0) not microgranulate; (1) microgranulate (Fig. 55). (continued next page ) Revision of Piestus with remarks on related genera Invertebrate Systematics 569 Appendix 1. (continued ) 51. Integument of interstriae of elytra: (0) without undulate microstriae; (1) with undulate microstriae. 52. Interstriae of elytra: (0) not carinate (Fig. 86); (1) carinate (Fig. 55). 53. Apical margin of elytra: (0) truncate (Fig. 54); (1) projected (Fig. 53). Abdomen: 54. Sternite 3: (0) without transverse carina (Fig. 143); (1) with transverse carina (Figs 10, 144). 55. Basolateral ridge on tergites 3–7: (0) absent; (1) present. 56. Paratergites on segments 3–6: (0) one pair; (1) two pairs. 57. Punctures on tergites: (0) uniformly distributed (Fig. 46); (1) only in anterolateral regions (Fig. 15). 58. Very small emargination on each posterolateral area on sternite 7, external openings of abdominal defensive gland complex (Caron et al. 2008): (0) absent; (1) present (Fig. 11). 59. Lateral margin on sternite 7: (0) each without longitudinal set of long setae; (1) each with longitudinal set of long setae. 60. Apical margin of sternite 8 of female: (0) without short setae (Fig. 147); (1) with short setae (Fig. 148). 61. Apical margin of tergite 10: (0) without fringe (Fig. 149); (1) with fringe (Fig. 151). 62. Apex of tergite 10: (0) with 1 or 2 pairs of setae (Fig. 149); (1) with more than two pairs of setae (Fig. 151). Genitalia: 63. Base of the median lobe of the aedeagus in ventral view: (0) not bulbous (Fig. 181); (1) bulbous (Fig. 214). 64. Apex of the median lobe of the aedeagus, in lateral view: (0) not curved (Fig. 188); (1) curved (Fig. 186). 65. Apex of the median lobe of the aedeagus, in ventral view: (0) not emarginated medially (Fig. 214); (1) emarginate medially (Fig. 181). 66. Region of median lobe for supporting the base of lateral lobe: (0) not extended anteriorly (Fig. 212); (1) extended anteriorly (Fig. 179). 67. Setae on apex of ovipositor of female: (0) many (Fig. 155); (1) few, one or two (Fig. 154). 68. External margin of ovipositor of female: (0) convexly curved (Fig. 153); (1) emarginate (Fig. 154). 69. Basal region of the spermathecal duct: (0) not sclerotised (Fig. 270); (1) sclerotised (Fig. 262). 570 Invertebrate Systematics E. Caron et al. Appendix 2. Additional material examined Piestus lacordairei Laporte, 1835 NICARAGUA: Río San Juan: 1 specimen, 60 km SE San Carlos Refugio Bartola, 10
58.400 N 84
20.300 W, 100 m, on bark downed logs, 25.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 1 specimen, Hamburgfarm, 30.I.[19]26, Nevermann coll. (FMNH); 1 specimen, the same locality and collector, 4.II.1936 (FMNH); 1 specimen, no locality and collector, VII.[19]38 (FMNH); 3 specimens, Vara Blanca 2000 m, VIII.[19]38, no collector (FMNH); 2 specimens, no locality, 1.V.1940, ilegible coll. (FMNH); 1 specimen, ilegible locality, 3.VIII.39, no collector (FMNH); 1 specimen, ilegible locality, 23–28.I.1941, no collector (FMNH); 1 specimen, ilegible locality, 25.I.1941, no collector (FMNH); Guanacaste: 5 specimens, Cação Biological Station 10
550 38’N 85
270 7’W, 1050 m, under bark, 10.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 2 specimens, 8 km NE Sta. Elena Santa Elena Forest Res. 10
20.701’N 84
47.899’W, 1640 m, 11–17.VI.2001, S. and J. Peck coll. (SEMC); Alajuela: 1 specimen, Sarchí, 1000 m, 17.X.[19]34, Nevermann coll. (FMNH); 1 specimen, E.B. San Ramon R.B. San Ramon 27 km N and 8 km W San Ramon, 10
130 3000 N 84
350 3000 W, 850–950 m, wet premontane forest, 20.VI-6.VII.1999, R. Anderson coll. (SEMC); 1 specimen, Est. Biologica Alberto M. Brenes, 10
13.020 N 84
350 5100 W, 900–1000 m, 29.VI.–6.VII.1999, Johnson coll. (FMNH); Heredia: 4 specimens, La Selva Biol. Station nr Puerto Viejo de Sararapiqui, 18.II.1985, L. Herman coll. (AMNH); San José: 1 specimen, Naranjo, VII.1939, Uloya coll. (AMNH); 4 specimens, Naranjo, VII.[19]39, no collector (1 AMNH, 3 FMNH); 2 specimens, St. Maria Dota, 2.12.1994, H. Forster coll. (NMW); Cartago: 1 specimen, 13–16.II.1939, no collector (FMNH); Limón: 1 specimen, Lasmercedes Santa Clara, 200–300 m, 12.VI.[19]28, F. Nevermann coll. (FMNH); 1 specimen, Hamburg Farm, loose bark in cracks of wood, 22.11.1935, F. Nevermann coll. (FMNH); Puntarenas: 3 specimens, La Reserva de Monteverde, ~4 km E Monteverde, 5000–5500 ft, 4.III.1985, L. Herman coll. (AMNH); 1 specimen, Wilson Botanical Garden Las Cruces Biol. Stn, 1200 m, 27. V.1993, J. S. and A. K. Ashe coll. (SEMC); 2 specimens, UVITA, 3–22.11.1994, H. Forster coll. (NMW); 1 specimen, Corcovado National Park Sirena Stn. upper Ollas Trail, 140 m, 8
290 700 N 83
340 3900 W, flight intercept trap, 24–28.VI.2000, Z.H. Falin coll. (SEMC); 1 specimen, Corcovado National Park Sirena Stn upper Rio Claro Trail, 8
280 2900 N 83
350 800 W, 100 m, flight intercept trap, 28.VI-1.VII.2000, Z. Falin coll. (SEMC); 1 specimen, Rincon de Osa 50 m, 8
41.141’N 83
31.117’W, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 1 specimen, Las Cruces Biol. Sta., 08
47.140 N 82
57.580 W, 1330 m, flight intercept trap, 28–31.V.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC). PANAMA: 4 specimens, Canal Zone Barro Colorado I., under bark, 14.I.1959, H. S. Dybas coll. (FMNH); 6 specimens, the same locality and collector, fermented fibrous log and at light, 16.I.1959 (FMNH); 4 specimens, the same locality and collector, Fairchild Trail, bark and under bark debris from fallen tree, 28.I.1959 (FMNH); 2 specimens, Canal Zone 4 mi. NW Gamboa, under bark, 23.II.1975, J. F. Lawrence coll. (FMNH); 11 specimens, Canal Zone Barro Colorado Is., under bark fermenting, 16–22.II.1976, A. Newton coll. (3 DZUP, 8 FMNH); 2 specimens, the same locality and collector, under bark fermenting logs, 25.II.1976 (FMNH); Bocas del Toro: 1 specimen, Almirante dam on Nigua Creek, berlese, 23.III.1959, H. S. Dybas coll. (FMNH); 3 specimens, the same locality and collector, berlese, 25.III.1959 (FMNH); 1 specimen, the same locality and collector, berlese, 27.III.1959 (FMNH); Chiriquí: 1 specimen, La Fortuna 1200 m, Continental Divide Trail, 8
460 0’N 82
120 0’W, berlese forest litter, 9.VI.1995, R. Anderson coll. (SEMC); 1 specimen, La Fortuna Cont. Divide Trail, 08
460 N 82
120 W, 1150 m, under bark, 9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, La Fortuna Río Hornito Trail, 08
420 N 82
140 W, 1000 m, under bark, 11.VI.1995, the same collector (SEMC); Veraguas: 2 specimens, Cerro Santa Rita, chips from cut end of log, 9.II.1959, H. S. Dybas coll. (FMNH); Colón: 1 specimen, Parque. Nac. Soberania Pipeline Rd km 61, 09
070 N 79
450 W, 40 m, flight intercept trap, 7–21.VI.1995, J. Ashe, R. Brooks coll. (SEMC). COLOMBIA: Cundinamarca: 4 specimens, Finca San Pablo 3 km N Alban, 1800 m, 1–12.VIII.1967, no collector (AMNH); Huila: 1 specimen, La Plata, no date and collector (NMW). FRENCH GUIANA: Saint-Laurent-du-Maroni: 5 specimens, Saül, under bark, III.-IV.1999, A. Berkov coll. (AMNH); Cayenne: 3 specimens, no locality, date and collector (1 IRSNB, 2 ZMHB); 1 specimen, Roura 12.3 km SSE, 4
380 55’N 52
180 4’W, 260 m, under bark, 29.V.1997, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, Roura 55.4 km SSE Kaw marshes, 4
290 58’N 52
30 0’W, 40 m, under fermenting bark, 11.VI.1997, the same collector (SEMC). ECUADOR: 1 specimen, no locality, date and collector (FMNH); Esmeraldas: 1 specimen, Cachabé low c., XI.[19]96, Rosenberg coll. (FMNH); 4 specimens, Cachabé low c., I.[19]97, Rosenberg coll. (2 FMNH, 2 IRSNB); 1 specimen, Cachabé low c., XII.?, Rosenberg coll. (IRSNB); Pichincha: 1 specimen, 45 km NNW Quito Macquipucuna Station, 1600–1650 m, flight intercept trap, 18.IV–5. V.1996, P. Hibbs coll. (SEMC); 1 specimen, Nanegalito 7 km S on Nono Road, 0
00 23’N 78
400 36’W, 1540 m, pyrethrun fogging allen tree trunk, 30.X.1999, Z. H. Falin coll. (SEMC); Napo: 1 specimen, Sierra Azul Hacienda Aragon 0
400 0’S 77
550 0’W, 2300 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); Cotopaxi: 1 specimen, Naranjito nr San Francisco de la Pampas 1500ft., cane-sugar, 18.V.1993, J.J. Morrone, E. Tapia coll. (AMNH); 1 specimen, N of San Francisco de las Pampas vic. Rio Esmeraldas, 4400–5000 ft., litter and debris, 14–15.V.[19]/93, L. Herman coll. (AMNH). PERU: Ucayali: 1 specimen, Boqueron del Padre Abad, 470 m, 8.VIII.1946, no collector (AMNH); Cusco: 1 specimen, Consuelo Manu rd. km 165, rotten palm and leaf litter, 1.X.1982, L.E. Watrous, G. Mazurek coll. (FMNH); 1 specimen, Marcapata, no date and collector (IRSNB). BRAZIL: Pará: 1 specimen, no locality, date and collector (IRSNB); Rondônia: 2 specimens, Cacaulândia, II.1997, Vaz Mello coll. (AMBC). Undetermined country: 1 specimen, no locality, date and collector (IRSNB). Piestus capricornis Laporte, 1835 PERU: Loreto: 2 specimens, Napo River 80 mi NE of Iquitos, 7.XII.1980, T. King coll. (FMNH). BRAZIL: Amazonas: 1 specimen, Fonte Boa Estr. Mamopina 023227S 660408W, 21–24.IX.2005, J.A. Rafael, F.F. Xavier Fo coll. (INPA); Pará: 3 specimens, Benevides, no date and collector (IRSNB); Rondônia: 2 specimens, Porto Velho Campus UNIR, 08
500 04’S 63
560 35’W, 19.IV.2006, J.A. Rafael, F.F. Xavier Fo coll. (1 DZUP, 1 INPA); 5 specimens, Vilhena, 12
460 55’S 60
220 18’W, 25.IV.2006, J.A. Rafael, F.F. Xavier Fo coll. (1 DZUP, 4 INPA). Undetermined country: 1 specimen, Cayenne, Guadalupe, no date and collector (IRSNB); 1 specimen, no locality, date and collector (FMNH). Piestus planatus (Sharp, 1887) MEXICO: 1 specimen, no locality, date and collector (FMNH); 1 specimen, Sr. Durango, V.[19]05, Speyer coll. (ZMHB); Chiapas: 1 specimen, Palenque, 18.VII.1992, O. Hillert coll. (ZMHB); Veracruz: 1 specimen, Córdova, no date, J. Flohr coll. (ZMHB); Guerrero: 2 specimens, Atovac, no date, J. Flohr coll. (ZMHB); Oaxaca: 2 specimens, no locality, date and collector (IRSNB); 2 specimens, 5.7 mi S Valle Nacional, 2000’, under bark hardwood, 11–18.VIII.1973, A. Newton coll. (FMNH). GUATEMALA: 1 specimen, no locality, date and collector (IRSNB); Alta Verapaz: 1 specimen, Panzos, no date and collector (ZMHB). NICARAGUA: Río San Juan: 11 specimens, 60 km SE San Carlos Refugio Bartola, 10
58.400 N 84
20.300 W, 100 m, on bark downned logs, 25.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (1 DZUP, 10 SEMC); 1 specimen, the same locality and collector, under bark, 27.V.2002 (SEMC). COSTA RICA: 1 specimen, no locality, date and collector (ZMHB); 1 specimen, Escocia, 7.IV.[19]24., Nevermann coll. (FMNH); Alajuela: 7 specimens, E.B. San Ramon Revision of Piestus with remarks on related genera Invertebrate Systematics 571 R.B. San Ramon 27 km N and 8 km W San Ramon, 10
130 30’N 84
350 30’W, 1120 m, rotting palm trunk, 29.VI–6.VII.1999, R. Anderson coll. (SEMC); Limón: 1 specimen, Hamburgfarm Reventazón, 14.X.[19]24, F. Nevermann coll. (FMNH); 1 specimen, the same locality and collector, no date (FMNH); 1 specimen, the same locality and collector, 12.XI.1925 (FMNH); 1 specimen, the same locality and collector, 15.XI.1925 (AMNH); 1 specimen, the same locality and collector, 4.X.[19]28 (FMNH); 2 specimens, the same locality and collector, 6.VI.[19]30 (1 FMNH, 2 AMNH); 2 specimens, the same locality and collector, 3.XI.[19]33 (FMNH). PANAMA: Darién: 3 specimens, Cana ANCON Station, 07
45.323’N 77
41.069’W, 500 m, 3–9.VI.1996, S. Lingafelter coll. (1 DZUP, 2 SEMC); 4 specimens, Cana Biological Stn., Serranía de Pirre 7
450 18’N 77
410 6’W, 975 m, 4.VI.1996, A. Gillogly coll. (SEMC); 3 specimens, Cana ANCON Station, 07
450 N 77
410 W, 900–1100 m, 5.VI.1996, S. Lingafelter coll. (SEMC); 6 specimens, Cana Biological Station, 7
450 18’N 77
410 6’W, 530 m, 09.VI.1996, J. Ashe, R. Brooks coll. (SEMC). ECUADOR: Esmeraldas: 1 specimen, Cachabé low c., no date, Rosenberg coll. (IRSNB); 1 specimen, Cachabé low c., XI.[19] 96, Rosenberg coll. (IRSNB). Undetermined country: 4 specimens, no locality, date and collector (ZMHB). Piestus spinosus (Fabricius, 1801) VENEZUELA: Bolívar: 1 specimen, El Pauchi bei Santa Elena, 01.III.1995, O. Hillert coll. (ZMHB). FRENCH GUIANA: Saint-Laurent-du-Maroni: 1 specimen, Juin, no date and collector (FMNH); 1 specimen, no locality, date and collector (AMNH); 1 specimen, Maripasoula Lawa River, under bark, 7.XII.1963, B. Malkin coll. (FMNH); 1 specimen, Saül under bark, III–IV.1999, A. Berkov coll. (AMNH); Cayenne: 1 specimen, Pariacabo, no date and collector (IRSNB); 2 specimens, no locality, date and collector (ZMHB); 7 specimens, Roura 39.4 km SSE, 4
320 43’N 52
80 26’W, 270 m, under fermenting bark, 10.VI.1997, J. Ashe, R. Brooks coll. (SEMC). ECUADOR: Napo: 1 specimen, no locality, date and collector (ZMHB); 2 specimens, 18 km S Tena, 28.IV.1978, C. W. and L. B. O’Brien and Marshall coll. (FMNH); Pastaza: 3 specimens, Cusuimi on Rio Cusuimi 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH). PERU: Loreto: 1 specimen, no locality, date and collector (ZMHB); Ucayali: 1 specimen, Boqueron del Padre Abad, 470 m, 24.VIII.1946, F. Woytkowski coll. (AMNH); Huánuco: 1 specimen, TingoMaria, 13.VII.1968, C. W. and L. B. O’Brien coll. (AMNH); Lima: 1 specimen, Salinas, III.1939, no collector (NMW); Junín: 1 specimen, Rio Toro, no date and collector (ZMHB); Huánuco: 7 specimens, Panguana Sira, 12.IX.1988, Listabarth coll. (NMW); Cusco: 1 specimen, Consuelo Manu rd. km 165, rotten palm and leaf litter, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the same locality and collector, litter under crown of felled tree, 2.X.1982 (FMNH); 2 specimens, the same locality and collector, rotten palm, 4.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 5.X.1982 (FMNH); 2 specimens, the same locality and collector, rotten palm, 6.X.1982 (FMNH); 1 specimen, the same locality and collector, flight intercept trap, 7.X.1982 (FMNH); 1 specimen, the same locality and collector, beating palm branches, 8.X.1982 (FMNH); 2 specimens, the same locality and collector, under bark, 11.X.1982 (FMNH). BOLIVIA: 1 specimen, Yuracaris, no date and collector (IRSNB); 2 specimens, Yuracaris, 6.XI.1900, no collector (1 FMNH, 1 NMW); Cochabamba: 2 specimens, El Chapa?, X.1949, L. Peña coll. (FMNH); 1 specimen, Chapar, no date and collector (DZUP); Santa Cruz: 1 specimen, Loc. Parque Amboró, 17
39’S 63
43’W, bajo corteza de Ficus sp., Rio Saguayo, no date and collector (FMNH); 1 specimen, Prov. Ichilo Loc. Parque Amboró, bajo corteza de Ficus sp., Rio Saguayo, 700 m, 26.XII.1988, no collector (FMNH). BRAZIL: 1 specimen, Marco de Legua, no date and collector (IRSNB); Amazonas: 1 specimen, Manaus, no date and collector (NMW); 3 specimens, no locality, date and collector (1 NMW, 2 FMNH); 1 specimen, Ega, no date and collector (IRSNB); Pará: 1 specimen, no locality and date, E. Sefer coll. (MNRJ); 2 specimens, no locality, date and collector (1 MZSP, 1 FMNH); 10 specimens, Aldeia Aracu Igarapé Gurupu-Umu Maranhão 50 km E of Caninde under bark, V.1963, B. Malkin coll. (FMNH); Rondônia: 4 specimens, Guaporé, 121605S 604230W, 327 m, 23.IV.2006, J.A. Rafael, F.F. Xavier Fo coll. (INPA); 2 specimens, Vilhena, 12
460 55’S 60
220 18’W, 25.IV.2006, J.A. Rafael, F.F. Xavier Focoll. (INPA); Mato Grosso: 7 specimens, Vila Vera, 55
300 12
460 , X.1973, M. Alvarenga coll. (AMNH); 4 specimens, Sinop, 12
310 55
370 , X.1974, M. Alvarenga coll. (AMNH). Undetermined country: 4 specimens, no locality, date and collector (1 NMW, 1 FMNH, 1 ZMHB, 1 ZMUC). Piestus longipennis Fauvel, 1864 COLOMBIA: 1 specimen, no locality, date and collector (IRSNB); Valle del Cauca: 6 specimens, Cali, no date, Fass coll. (1 DZUP, 5 NMW); 1 specimen, Carmen, 1400 m, 17.VI.1908, Fass coll. (FMNH); 1 specimen, Villa Elvira, 1800 m, 2.VII.1908, Fass coll. (FMNH); 1 specimen, Rio Fivaco, 2000 m, IX.1908, Fass coll. (FMNH); 1 specimen, S. Antonio, 2000 m, XI.1908, Fass coll. (FMNH). ECUADOR: Carchi: 1 specimen, 46 km W Tufino west slope, 2600 m, under dead wood bark, 19.XI.1987, C. Young and R. Davidson coll. (FMNH); Pichincha: 1 specimen, near Calicali, 1800 m, 19.XII.87, M. Huybensz coll. (FMNH); Napo: 2 specimens, 52 km N. Tena, 6500ft., 24.V.1993, L. Herman coll. (AMNH); 1 specimen, Cosanga, 4.2 km S on Baeza-Tena Road then 2.9 km W on pipeline access road, 2150 m, 0
370 19’W 77
500 1’W, on under bark downed logs, 7.XI.1999, Z. H. Falin coll. (SEMC); Cotopaxi: 1 specimen, Canton Sigchos Las Pampas Otonga Natural Reserve, 25–28.VII.2005, W. Rossi coll. (DZUP); 3 specimens, Canton Sigchos Las Pampas Bosque Integral Otonga, 7–10.VII.06, W. Rossi coll. (FMNH); 1 specimen, Bosque Integral Otonga, 1975 m, 79
000 204’ 00
250 166’, 02.06.[20]07, A.C. Proaño C. and A. Barragán coll. (DZUP); Tungurahua: 1 specimen, Baños, no date and collector (ZMHB). Piestus zisckai Scheerpeltz, 1951 ECUADOR: Tungurahua: 1 specimen, Santa Jnéz, no date and collector (ZMHB); Zamora-Chinchipe: 1 specimen, Rio Bombuscaro, 1100 m, 4
70 0’S 78
370 48’W, malaise trap, 26.VI–4.VII.1996, P. Hibbs coll. (SEMC). PERU: 1 specimen, no locality, date and collector (ZMHB). BOLIVIA: 1 specimen, Yuracaris, no date and collector (IRSNB). Piestus bicornis (Olivier, 1811) COSTA RICA: Cartago: 4 specimens, Turialba, 800 m, no date and collector (ZMHB); Limón: 2 specimens, Rio Barbilla, no date and collector (NMW); 1 specimen, Hamburgfarm Reventazon, 27.IX.[19]24, F. Nevermann coll. (FMNH); 1 specimen, Las Mercedes 200–300m Sta. Clara, 20.III.1922, the same collector (FMNH). PANAMA: 7 specimens, Canal Zone, Madden Dam, under bark of hardwood ferment. stage, 12.VI.1976, A. Newton coll. (FMNH); Panamá: 1 specimen, Ciricito Canal Zone, 16.I.[19]31, no collector (AMNH). COLOMBIA: 3 specimens, no locality, date and collector (1 ZMUC, 1 ZMHB, 1 IRSNB); Cundinamarca: 3 specimens, Bogota, no date and collector (ZMHB); Meta: 1 specimen, Villavicencio, 11.VII.[19]38, H. Dybas coll. (FMNH); 5 specimens, Villavicencio Meta, 25.VII.[19]38, H. S. Dybas coll. (FMNH); Putumayo: 19 specimens, Santa Rosa headwaters of Rio San Miguel, under bark of log, 16–20.X.1970, B. Malkin and P. Burchard coll. (FMNH). VENEZUELA: 1 specimen, Cafetal, no date and collector (FMNH); 3 specimens, Cafetal, V.1922, no collector (FMNH); Carabobo: 15 specimens, Las Trincheiras, VI.1922, L. R. Reynold coll. (FMNH). FRENCH GUIANA: 572 Invertebrate Systematics E. Caron et al. Saint-Laurent-du-Maroni: 1 specimen, 3.5mi N Saül, Les Eaux Claires, Treefall, 16.IV.[19]96 C. Chaboo coll. (AMNH); Cayenne: 2 specimens, no locality, date and collector (NMW). ECUADOR: Sucumbios: 1 specimen, Sacha Lodge, 0.5
S 76.5
W, 270 m, malaise, 27.VIII–10.IX.1994, Hibbs coll. (SEMC); Napo: 1 specimen, Tena, 14.II.1923, F.X. Williams coll. (FMNH); 1 specimen, Tena, 2100’, 3.XI.[19]88, L. Herman coll. (AMNH); Pastaza: 9 specimens, Cusuimi on Rio Cusuimi 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH); 8 specimens, Puyo nr. Santa Clara, 950 m, 18–21.VII.2008, W. Rossi and I. Tapia coll. (FMNH); Morona-Santiago: 1 specimen, Macas, no date and collector (FMNH). PERU: 5 specimens, no locality, date and collector (1 ZMHB, 2 AMNH, 2 FMNH); Loreto: 1 specimen, 20 km from Ucayali on R.Calleria Colonia Calleria, 5.X–10.X.1961, B. Malkin coll. (FMNH); 10 specimens, Yagua Indian Village headwaters of Loreto-Yacu, 2.V.1970, B. Malkin coll. (FMNH); San Martín: 7 specimens, Hera (Jera) Moyobamba San Martin, 860 m, 27.VI.1947, F. Woytkowski coll. (AMNH); Huánuco: 1 specimen, no locality and collector, 15.V.[19]38 (NMW); 3 specimens, Panguana Sira, 12.IX.1988, Listabarth coll. (NMW); 2 specimens, Panguana Rio Pachitea, Rio Yuyapichis, 9
370 S 74
560 W, 260 m, 1–14. IX.1968, Listabarth coll. (NMW); 3 specimens, Bella Durmiente near Tingo Maria, 14.VII.1968, C. W. and L. B. O’Brien coll. (AMNH); Junín: 2 specimens, no locality, date and collector (FMNH); 2 specimens, Chanchamajo, no date and collector (FMNH); 2 specimens, the same locality, 1000 m, no date and collector (ZMHB); 2 specimens, Rio Toro, no date and collector (ZMHB); 4 specimens, San Ramón de Pangoa 40 km SE Satipo, 750 m, 24. III.1972, R.T. and J.C. Schuh coll. (AMNH); Cusco: 1 specimen, Marcapata Perou Yungas Bolivie, 1000 m, no date and collector (IRSNB); 1 specimen, Marcapata, no date and collector (IRSNB); 5 specimens, Callanga, no date and collector (2 NMW, 3 ZMHB); 2 specimens, Quincemil, 2400ft., 15.IV.1947, J. C. Pallister coll. (AMNH); 3 specimens, Consuelo Manu rd. km 165, rotten palm, 3.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the same locality and collector, rotten palm, 4.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 5.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 5–15.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 6.X.1982 (FMNH); 2 specimens, the same locality and collector, beating palm branches, 8.X.1982 (FMNH); Madre de Dios: 2 specimens, Tambopata 15 km NE Puerto Maldonado Reserva Cuzco Amazónico, 12
330 S 69
030 W, 200 m, under bark, 9.VII.1989, J. S. Ashe, R. A. Leschen (MUSM). BOLIVIA: 1 specimen, Yuracaris, no date and collector (IRSNB); 1 specimen, VIII.1953, L. Peña coll. (FMNH); La Paz: 9 specimens, Yungas de la Paz, no date and collector (8 ZMHB, 1 FMNH); Beni: 1 specimen, Rurrenabaque, 23.X.1956, L. Pena coll. (FMNH); Cochabamba: 1 specimen, no locality, date and collector (DZUP); 11 specimens, Cristalmayu, 28.VIII.1949, L. Peña coll. (NMW); 3 specimens, Chapare, 400 m, no date, R. Zischka coll. (1 NMW, 2 ZMHB); 7 specimens, 20 mi SW Villa Tunari, 2.IV.1978, L. and C. W. O’Brien and G. B. Marshall coll. (FMNH); 2 specimens, 16.7 km W. Villan Tunari Avispas, 17
10 13’S 65
320 46’W, 500 m, 11.II.1999, R. Anderson coll. (SEMC); 5 specimens, 16.7 km W. Villa Tunari Parque Machias, 16
580 20’S 65
240 42’W, 300 m, 12.II.1999, R. Anderson coll. (SEMC); Santa Cruz: 1 specimen, Prov. Ichilo Loc. Parque Amboró, 17
390 S 63
430 W, Rio Saguayo bajo corteza de Ficus sp., no date and collector (AMNH); 1 specimen, Ichilo, 17
390 S 63
430 W, bajo corteza de Ficus sp., 31.VII.1988, no collector (FMNH); 3 specimens, Ichilo Cafetal, under bark of stump, 6.VIII.1990, no collector (FMNH); 1 specimen, Potrerillo de Guenda, 2.IX.[19]98, no collector (FMNH). BRAZIL: 4 specimens, no locality, date and collector (3 NMW, 1 IRSNB); 1 specimen, Pebas Ega, no date and collector (IRSNB); 1 specimen, Santa Rita, no date and collector (IRSNB); Acre: 1 specimen, Iquiri, VIII.[19]51, no collector (MZSP); Amazonas: 1 specimen, Manaus, Reserva Adolpho Ducke, 02
550 5100 S 59
580 5900 W, 13.III.2006, V. Rodrigues coll. (INPA); Pará: 16 specimens, Caninde Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH); Goiás: 1 specimen, Jatahy Rio Verde, no date and collector (IRSNB); 1 specimen, Jatahy, no date and collector (FMNH); 1 specimen, Pires do Rio, 2.XI.1956, J. R. Pacheco coll. (DZUP); Minas Gerais: 1 specimen, no date and collector (ZMHB); Espirito Santo: 1 specimen, no date and collector (IRSNB); 3 specimens, Corrego Itá, XI.1956, W. Zikan coll. (MNRJ); Rio de Janeiro: 1 specimen, Neue Friburg., no date and collector (ZMHB); São Paulo: 1 specimen, no locality, date and collector (MZSP); 2 specimens, Umgbg.v.Ribeirão Preto, III–VII.1899 (FMNH); Paraná: 1 specimen, Salto Mauá, V.[19]43, no collector (DZUP); 1 specimen, Caviuna, XII.1945, A. Maller coll. (AMNH); 1 specimen, Rolandia, IV.1948, A. Maller coll. (AMNH); Santa Catarina: 2 specimens, Blumenau, no date and collector (NMW); 1 specimen, N. Teutonia, no date, Plaumann coll. (FMNH); 1 specimen, no locality, date and collector (FMNH); 1 specimen, Corupá, I.1938, A. Maller coll. (MNRJ); 1 specimen, no locality and collector, III.1940 (NMW); 25 specimens, Nova Teutonia, VII.41, Dirings coll. (3 DZUP, 22 MZSP); 1 specimen, Corupa, XI.1948, A. Maller coll. (AMNH). PARAGUAY: Guairá: 2 specimens, 300 m, 25.VII.1951, C. Pfannel coll. (FMNH). Undetermined country: 10 specimens, no locality, date and collector (5 ZMHB, 2 NMW, 2 IRSNB, 1 FMNH). Piestus fronticornis (Dalman, 1821) NICARAGUA: 1 specimen, no locality, date and collector (IRSNB); Granada: 1 specimen, Res. Nat. Volcan Mombacho, 11
50.050 N 85
58.830 W, 1060 m, flight intercept trap, 1–5.VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC); Río San Juan: 1 specimen, 60 km SE San Carlos Refugio Bartola, 10
58.400 N 84
20.300 W, 100 m, on bark downed logs, 25.V.2002, R.Brooks, Z.Falin, and S.Chatzimanolis coll. (SEMC). GUATEMALA: Guatemala: 2 specimens, no locality, 27.VI.1969, G. Ekis coll. (AMNH). COSTA RICA: 1 specimen, no locality, date and collector (ZMHB); Guanacaste: 1 specimen, Puerto Humo, 17.VIII.[19]66, F. Fisk coll. (FMNH); Alajuela: 2 specimens, San Carlos, no date and collector (1 DZUP, 1 ZMHB); Heredia: 1 specimen, La Selva Biol. Statíon nr. Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); San José: 2 specimens, La Uruca, 1100 m, no date and collector (IRSNB); 29 specimens, no locality and date, E. Schmidt coll. (AMNH); 15 specimens, no locality, date and collector (AMNH); 1 specimen, La Caja 8 Kil.W., 1931, Schmidt coll. (FMNH); 2 specimens, La Caja, 1934, no collector (1 AMNH, 1 FMNH); 2 specimens, no locality and collector, 1936 (FMNH); 5 specimens, La Caja, II.1940, no collector (ZMHB); 3 specimens, no locality, VIII.[19]64, F. Fisk coll. (FMNH); 1 specimen, San Antonio de Escazu, malaise trap, 25.III–9.IV.1984, S.A. Cameron coll. (AMNH); 11 specimens, St. Maria Dota, 2.XII.1994, H. Forster coll. (NMW); Limón: 1 specimen, 1000–1200 m, X.[19]28, F. Nevermann coll. (FMNH); 1 specimen, 4.X.[19]28, F. Nevermann coll. (FMNH); 1 specimen, willted banana leaves, 25.XI.1928, F. Nevermann coll. (FMNH); 1 specimen, 1.V.1931, Nevermann coll. (FMNH); 1 specimen, 26.XI.[19]33, Nevermann coll. (FMNH); 1 specimen, decaying banana blossoms leaves, 26.I.1935, F. Nevermann coll. (FMNH); Puntarenas: 21 specimens, UVITA, 3.XI.–22. XI.1994, H. Forster coll. (NMW); 1 specimen, Corcovado National Park Sirena Stn. lower Ollas Trail, 5 m, fogging rotting sap flow, 24.VI.2000, Z. H. Falin coll. (SEMC); 4 specimens, Corcovado National Park Sirena Stn. lower Ollas Trail 5m 8
240 4800 N 83
350 2200 W under bark, 25.VI.2000, the same collector (SEMC). PANAMA: 1 specimen, no locality, date and collector (IRSNB); 1 specimen, Canal Zone Barro Colorado I., 13.I.1959, H.S. Dybas coll. (DZUP); 4 specimens, the same locality and collector, 14.I.1959 (1 DZUP, 3 FMNH); 4 specimens, the same locality and collector, fermented fibrous log at light, 16. I.1959 (FMNH); 1 specimen, the same locality and collector, 21.I.1959 (FMNH); 2 specimens, Canal Zone, Madden Dam, under bark of hardwood ferment stage, 12.VI.1976, A. Newton coll. (FMNH); 5 specimens, Canal Zone Barro Colorado Is., under bark fermenting, 16–22.II.1976, the same collector (FMNH); 1 specimen, Canal Z. Barro Colo. Is., sift fermenting bark, 20.VIII.1978, Q. D. Wheeler (FMNH); 2 specimens, the same locality and collector, 26. VIII.1978 (FMNH); Bocas del Toro: 1 specimen, Almirante trail to dam on Nigua Crk., decayed palm log, 29.III.1959, H. S. Dybas coll. (FMNH); Chiriquí: 2 specimens, V. de Chiriqui 25–4000ft., no date, Champion coll. (FMNH); 3 specimens, Bugaba, no date, the same collector (1 NMW, 1 AMNH, 1 ZMHB); 1 specimen, La Fortuna Cont. Divide Trail, 08
460 N 82
120 W, 1150 m, under bark, 9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); Veraguas: 1 specimen, Revision of Piestus with remarks on related genera Invertebrate Systematics 573 no locality and collector, IX–X.1938 (FMNH); Colón: 1 specimen, no locality and collector, 28.I.1925 (FMNH); 1 specimen, Canal Zone 4mi. NW Gamboa, under bark, 23.II.1975, J. F. Lawrence coll. (FMNH); 3 specimens, Black Tank Rd. NW Gatun Locks, under bark, 2.VI.1995, B. Ratcliffe and M. Jameson coll. (SEMC); 1 specimen, 10–15 km N jct. Escobal and Piña Rds, ~30 m, under bark, 2.VI.1996, J. Ashe, R. Brooks coll. (SEMC); Panamá: 2 specimens, Ciricito Canal Zone, 13.III.[19]30, no collector (AMNH); 5 specimens, the same locality, 14.III.[19]30, no collector (AMNH); 2 specimens, the same locality, 19.III.[19]30, no collector (AMNH); 1 specimen, the same locality, 28.II.[19]31, no collector (AMNH); Darién: 2 specimens, Cana Biological Station, 530 m, 7
450 N 77
410 6’W, 09.VI.1996, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: 1 specimen, no locality, date and collector (ZMHB); Cundinamarca: 1 specimen, Bogota, no date and collector (ZMHB); Valle del Cauca: 2 specimens, R. Dagua, IV.[18]97, W. Rosenberg coll. (IRSNB); 1 specimen, PNN Farallones de Cali Anchicaya, 3
260 N 76
480 W, 650 m, malaise, 19.XII.2000–02.I.2001, S. Sarria coll. (SEMC). FRENCH GUIANA: Saint-Laurent-du-Maroni: 1 specimen, no locality, date and collector (FMNH); 1 specimen, Les Eaux Claires, 3. XI.1995, A. Berkov coll. (AMNH); 2 specimens, 3.5mi N Saül Les Eaux Claires, IV.[19]96, A. Berkov coll. (AMNH); 4 specimens, Saül 7 km N Las Eaux Claires 3
390 4600 N 53
130 1900 W, 220 m, under bark, 1.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, Saül, Lecythis zabucajo fallen branch, III–IV.1999, A. Berkov coll. (AMNH); Cayenne: 3 specimens, no locality, date and collector (IRSNB). ECUADOR: Esmeraldas: 3 specimens, San Marco, 14.IX.1956 (FMNH); 2 specimens, Zapallo Grande, 25–30.X.1987, M. Huybensz coll. (FMNH); Pastaza: 5 specimens, Puyo nr. Santa Clara, 950 m, 18–21.VII.2008, W. Rossi and I. Tapia coll. (FMNH). BRAZIL: Amazonas: 1 specimen, Manaus, XI.1961, R. Arlé coll. (INPA); 1 specimen, Est. Br 17 km 34, 7.VIII.[19]69, E.V. Silva coll. (INPA); 1 specimen, 35 km NE Manaus Res. Flor. Ducke, 25.VII–08.VIII.[19]95, Arndt and Gröger coll. (ZMHB); Amapá: 1 specimen, Serra do Navio, IX.1957, K. Lenko coll. (MZSP); Pará: 1 specimen, no locality, date and collector (IRSNB); 2 specimens, Taperinha Santarem, 1–10.VII.[19]27, Zerny coll. (NMW); 2 specimens, Tiriós Alto Parú d’Oeste, I–II.1963, Machado, Pereira coll. (MZSP); 6 specimens, Caninde Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH). Undetermined country: 1 specimen, no locality, date and collector (NMW). Piestus validus Sharp, 1876 COLOMBIA: Meta: 1 specimen, Villavicencio, 25.VII.[19]88, H. S. Dybas coll. (FMNH). VENEZUELA: Carabobo: 2 specimens, San Esteban Ven. nr. Puerto Cabello, 1–20.XII.1939, P. J. Anduze coll. (FMNH); Aragua: 3 specimens, Rancho Grande Biol. Stn., 10
210 000 N 67
410 000 W, 1300 m, flight intercept trap, 12–14.V.1998, J. Ashe, R. Brooks and R. Hanley coll. (SEMC); 1 specimen, the same locality and collector, flight intercept trap, 14.V–2.VI.1998 (SEMC). ECUADOR: 1 specimen, Macas, no date and collector (FMNH); 2 specimens, no locality, date and collector (FMNH); 2 specimens, Libertad, 12.V.[19]63, L. Peña coll. (FMNH); 2 specimens, 8 km NE Puyo, 28.IV.1978, C. W. and L. B. O’Brien and Marshall coll. (FMNH); 1 specimen, Mera, 25.I.[19]23, F.X. Williams coll.; Napo: 4 specimens, Coca, V.1965, L. Pena coll. (FMNH); Pastaza: 1 specimen, Pomona on Rio Pastaza, 2000ft., 9.VIII.1960, T. Sabine coll. (FMNH); 1 specimen, Cusuimi on Rio Cusuimi 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH); Loja: 1 specimen, Pompeya, 13–25.V.1965, L. Pena coll. (FMNH). PERU: Junín: 1 specimen, Chanchamayo, no date and collector (FMNH); Huánuco: 1 specimen, Chinchao 25 km. Below Carpish, 2500 m, 6.IX.1946, F. Woykowski coll. (AMNH); 1 specimen, Panguana Sira, 12.IX.1988, Listabarth coll. (NMW); 1 specimen, Panguana Río Pachitea Rio Yuyapichis, 9
370 S 74
560 W, 260 m, VI.1986, Listabarth coll. (NMW); Cusco: 1 specimen, Marcapata, no date and collector (IRSNB); 9 specimens, Callanga, no date and collector (NMW); 3 specimens, Consuelo Manu rd. km 165, rotten palm, 3.X.1982, L.E. Watrous and G. Mazurek coll. (1 DZUP, 2 FMNH); 7 specimens, the same locality and collector, rotten palm, 4.X.1982 (1 DZUP, 6 FMNH); 8 specimens, the same locality and collector, 5–15.X.1982 (1 DZUP, 7 FMNH); 5 specimens, the same locality and collector, rotten palm, 6.X.1982 (FMNH); 7 specimens, the same locality and collector, beating dead branches, 8.X.1982 (FMNH); 4 specimens, the same locality and collector, palm litter, 11.X.1982 (FMNH); 4 specimens, the same locality and collector, rotten palm, 12.X.1982 (FMNH); Madre de Dios: 2 specimens, Tambopata, 25.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH). BOLIVIA: 3 specimens, the same locality and collector (1 ZMHB, 2 FMNH); 2 specimens, Yuracaris, 6. XI.1900, no collector (1 NMW, 1 FMNH); 5 specimens, Yuracaris, no date and collector (1 IRSNB, 4 NMW); 1 specimen, Yuracaris, S. Antonio, no date and collector (IRSNB); La Paz: 1 specimen, Yungas de la Paz, no date and collector (ZMHB); 1 specimen, Yungas de La Paz 1000 m, no date, H. Rolle coll. (ZMHB); 1 specimen, Yungas de la Paz, no date, V. Heyne coll. (ZMHB); Cochabamba: 4 specimens, Cochabamba 109 km E Yungas, CochabambaVila Tunari Rd., 17
80 5000 S 65
420 2900 W, 1480 m, flight intercept trap, 1–6.II.1999, F. Genier coll. (SEMC); 1 specimen, Cochabamba 109 km E Yungas, Cochabamba-Vila Tunari Rd., 17
80 5200 S 65
420 5400 W, 1400 m, flight intercept trap, 8–12.II.1999, F. Genier (SEMC). BRAZIL: 4 specimens, Pebas, no date and collector (1 FMNH, 3 IRSNB). Undetermined country: 5 specimens, no locality, date and collector (1 NMW, 4 ZMHB). Piestus pennicornis Fauvel, 1864 NICARAGUA: Matagalpa: 1 specimen, 6 km N Matagalpa, Selva Negra Hotel, 12
59.990 N 85
54.530 W, 1350 m, under bark, 21.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC); Granada: 1 specimen, Res. Nat. Volcan Mombacho, 11
50.050 N 85
58.830 W, 800–1000 m, under bark and upright tree, 2.VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). PANAMA: Colón: 1 specimen, Fort Sherman, 15.I.1980, Stockwell coll. (FMNH); Veraguas: 9 specimens, Cerro Tule, 4 km W of Santa Fe, 850 m, 30.VII.1995, A. Gillogly coll. (SEMC). COLOMBIA: 2 specimens, Bogota, no date and collector (1 ZMHB, 1 FMNH). VENEZUELA: 1 specimen, Caracas, no date and collector (IRSNB); 3 specimens, Las Trincheras, VI.1922, L. R. Reynold coll. (FMNH). ECUADOR: Sucumbios: 2 specimens, Sacha Lodge, 0
280 1400 S 76
270 3500 W, 270 m, fruit fall, 23.III.1999, R. Brooks coll. (SEMC); Napo: 1 specimen, Cosanga, 4.2 km S on Baeza-Tena Road then 2.9 km W on pipeline access road, 0
370 1900 S 77
500 100 W, 2150 m, under bark, 6. XI.1999, Z. H. Falin coll. (SEMC); 1 specimen, the same locality and collector, 7.XI.1999 (SEMC). PERU: Cusco: 1 specimen, Marcapata, no date and collector (IRSNB); 10 specimens, Consuelo, Manu rd km 165, rotten palm, 3.X.1982, L. E. Watrous and G. Mazurek coll. (1 DZUP, 9 FMNH); 1 specimen, the same locality and collector, beating palm branches, 8.X.1982 (FMNH); 1 specimen, the same locality and collector, rotten palm, 11.X.1982 (DZUP). BOLIVIA: 3 specimens, Yuracaris, no date and collector (IRSNB); Cochabamba: 1 specimen, 17 mi N Villa Tunari, 1.IV.1978, L. and C.W. O’Brien and G.B. Marshall coll. (FMNH). BRAZIL: 3 specimens, Pebas, no date and collector (IRSNB); 1 specimen, no locality and date, A. Böttcher coll. (ZMHB); 2 specimens, no locality, date and collector (1 MNRJ, 1 FMNH); Rondônia: 1 specimen, Guaporé, 12
160 0500 S 60
420 3000 W, coleta manual em tronco, 23. IV.2006, J.A. Rafael and F.F. Xavier Fo, coll. (INPA); Pará: 2 specimens, no locality, date and collector, ‘Ex-Typis’ (IRSNB); 1 specimen, no locality, date and collector (FMNH); 4 specimens, Caninde, Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH); Rio de Janeiro: 4 specimens, Nova Friburgo, no date, L.W. Schaufuß coll. (ZMHB); 1 specimen, no locality and date, F. Sahlhberg coll. (FMNH); 3 specimens, no locality and collector, Fry coll. (2 FMNH, 1 ZMHB); 1 specimen, no locality, date and collector (FMNH); 2 specimens, Pétropolis, no date, Lfévrier coll. (IRSNB); 1 specimen, the same locality, no date and collector (IRSNB); São Paulo: 6 specimens, no locality and date, M. Ráz coll. (5 FMNH, 1 NMW); 1 specimen, São Bernardo, IX.1932, J. Guerin coll. (FMNH); Paraná: 1 specimen, Rolândia, XII.1947, A. Maller coll. (AMNH); 1 specimen, the same locality and collector, IX.1948 (AMNH); Santa 574 Invertebrate Systematics E. Caron et al. Catarina: 1 specimen, no locality and date, Klimsch coll. (FMNH); 2 specimens, Blumenau, no date and collector (NMW); 1 specimen, no locality, date and collector (ZMHB); 1 specimen, Rio Vermelho, I.1946, A. Maller coll (AMNH); 1 specimen, Corupá, XI.1948, A. Maller coll. (AMNH). PARAGUAY: Caazapá: 4 specimens, Hermosa, prop. Sosa family, San Rafael Reserve, 26
190 1500 S 55
440 5500 W, 90 m, fermenting tree wound, 4.XII.2000, Z.H. Falin coll. (SMEC). Undetermined country: 1 specimen, no locality, date and collector (AMNH); 4 specimens, no locality and date, L.W. Schaufuß coll. (ZMHB). Piestus pygmaeus Laporte, 1835 MEXICO: 8 specimens, no locality, date and collector (3 ZMHB, 3 IRSNB, 1 FMNH, 1 NMW); 1 specimen, Tuxtepec, no date, Flohr coll. (ZMHB); 3 specimens, Almolonga, no date, Flohr coll. (ZMHB); 8 specimens, Jalapa, no date, W. Schaus coll. (AMNH); 5 specimens, no locality and date, J. Flohr coll. (ZMHB); 4 specimens, Jalapa, no date, Hoege coll. (1 FMNH, 1 NMW, 2 ZMHB); 7 specimens, no locality and date, Truqui col. (FMNH); 4 specimens, no locality, 30.VI.1897, C. Hoge coll. (1 NMW, 3 AMNH); San Luís Potosí: 5 specimens, 12 mi S Cd. Mante, 21.VI.1975, L. E. Watrous coll. (FMNH); Jalisco: 7 specimens, 1 Mi.S.W. La Resolana, 20.XI.1950, R. F. Smith coll. (AMNH); 1 specimen, Las Jarillas, 62 km S/P. Vallarta, under bark and logs, 22.VII.1984, Chemsak and Doyen coll. (FMNH); Veracruz: 2 specimens, Las Vigas, no date, Hoege coll. (NMW); 2 specimens, Jalapa, no date and collector (1 ZMHB, 1 AMNH); 1 specimen, Almolonga, no date and collector (ZMHB); 2 specimens, El Fortin, 20.VII.1936, C. H. Seevers coll. (FMNH); 2 specimens, Tejeria, 4.VII.1944, H. Dybas coll. (FMNH); 1 specimen, Tierra Blanca, 28.VII.1944, H. Dybas coll. (FMNH); 1 specimen, 6 mi NE Catemaco, rain for., 1500ft., under bark, 30.VII–8.VIII.1970, A. Newton coll. (FMNH); México: 2 specimens, Real de Arriba, Temescaltepec, 19.VI.[19]33, H. E. Hinton and R. L. Usinger coll. (AMNH); Oaxaca: 4 specimens, Oaxaca, 4.5 mi S Valle Nacional, under bark hardwood, 13–16.VIII.1973, A. Newton coll. (FMNH); 3 specimens, Oaxaca, 5.7 mi S Valle Nacional, under bark hardwood, 11–18.VIII.1973, A. Newton coll. (FMNH); 2 specimens, Tuxtepec, no date and collector (ZMHB); Chiapas: 3 specimens, Cahabon, Vera Paz, no date, Champion coll. (1 AMNH, 1 NMW, 1 ZMHB); Nayarit: 1 specimen, El Cora, Tepic, no date and collector (ZMHB). BELIZE: Orange Walk: 1 specimen, Rio Bravo Conservation Area, La Milpa Ruins, 17
500 N 89
020 W, FIT, 15–25.V.1997, C. Carlton coll. (SEMC); Belize: 8 specimens, Altun Ha, 11.VIII.1977, L. and C. W. O’Brien and G. B. Marshall coll. (FMNH); Toledo: 1 specimen, BARC San Pedro Columbia, 16
160 4300 N 88
570 4900 W, under bark at night, 25.IX.2004, P. W. Kovarik coll. (FMNH). GUATEMALA: 3 specimens, Zapote, no date, G.C. Champion coll. (1 ZMHB, 2 FMNH); Zacapa: 4 specimens, 24 km ENE Gualán, 280 m, under bark, 26.VI.1993, J. Ashe and R. Brooks coll (SEMC); Escuintla: 1 specimen, Finca El Zapote, 2400ft., under bark, 6.VII.[19]48, R.D. Mitchell coll. (FMNH); 8 specimens, the same locality and collector, 7.VII.[19]48 (FMNH); 1 specimen, the same locality and collector, 9.VII.[19]48 (FMNH); 5 specimens, the same locality and collector, 10.VII.[19]48 (FMNH); 6 specimens, the same locality and collector, 11.VII.[19]48 (FMNH). HONDURAS: Santa Bárbara: 1 specimen, El Mochilo, 13 km SE, 22.VII.1977, C. W. and L. O’Brien and Marshall coll. (FMNH); 1 specimen, Santa Barbara, La Fe, Finca La Roca, 5.3 km, S. Peña Blanca, 14
570 N 88
020 W, 740 m, under bark, 21.VI.1994, Brooks and Ashe coll. (SEMC); Colón: 2 specimens, 3mi SW, Sonaguera, 23.VIII.1956, B. and B. Valentine coll. (FMNH); Francisco Morazán: 1 specimen, Zamorano, 14
N 87
W, 820 m, rotting breadfruit, 6.VI.1994, J. Ashe and R. Brooks coll. (SEMC). NICARAGUA: 1 specimen, no locality, date and collector (NMW); Granada: 20 specimens, Reserva Natural Volcan Mombacho, 11
50.50 N 86
00.740 W, 375 m, under fermenting guanacaste lumber, 1.VI.2002, R. Brooks, Z. Falin, and S. Chatzimanolis coll. (SEMC); 39 specimens, the same locality and collector, 4.VI.2002 (SMEC); 3 specimens, Res. Nat. Volcan Mombacho, 11
50.050 N 85
58.830 W, 1200 m, under bark, 2.VI.2002, R. Brooks, Z. Falin, and S. Chatzimanolis (SEMC); Río San Juan: 1 specimen, 60 km SE San Carlos, Refugio Bartola, 10
58.400 N 84
20.300 W, 100 m, on bark downed logs, 25.V.2002, R.Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 1 specimen, Costa Rica, no date, P. Biolley coll. (FMNH); 1 specimen, Rio Virilla, 26.XII.[19]31, H. Schmidt coll. (SEMC); 1 specimen, San Mateo, Macacona, no date and collector (IRSNB); Guanacaste: 1 specimen, S. Cañas, 9–14.II.1989, F.D. Parker coll. (SEMC); Heredia: 2 specimens, La Selva Biol. Station nr. Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); Cartago: 1 specimen, Turialba, 800 m, no date, A. Heyne coll. (ZMHB); 2 specimens, Vulkan Turialba, 10 km N, no date and collector (ZMHB); Limón: 1 specimen, Lasmercedes, Santa clara, 200–300 m, 17.X.[19]26, F. Nevermann coll. (FMNH); 1 specimen, Ramal Parisimina, Santa Clara, 21.X.[19]26, the same collector (FMNH); 1 specimen, Hamburgfarm Reventazon Ebene Limon, 5.X.[19]28, the same collector (FMNH); 1 specimen, 31.X.[19]31, the same collector (FMNH); 1 specimen, Reventazon, Hamburg Farm, 27.IV.1934, the same collector (FMNH); 1 specimen, the same locality and collector, on dry bark of Castilla, 21.VIII.[19]36 (FMNH); 6 specimens, the same locality and collector, in dry inner bark of Lecythis, 1.X.1936 (FMNH); 1 specimen, no localty and collector, VII.[19]38 (FMNH); Puntarenas: 8 specimens, UVITA, 3.XI–22.XI.1994, H. Forster coll. (NMW); 2 specimens, Corcovado National Park, Sirena, logging recently splintered tree, 26.VI.2000, Z. H. Falin coll. (SMEC); 1 specimen, Corcovado National Park, Sirena Stn., upper Rio Claro Trail, 8
280 2900 N 83
350 800 W, 100 m, flight intercept trap, 28.VI–1.VII.2000, Z. Falin coll. (SEMC); 14 specimens, Corcovado National Park, Sirena Stn., junc. Guanacaste-Sirena Trails, 8
280 5500 N 83
350 4600 W, 5 m, under bark, 30.VI.2000, Z. H. Falin coll. (SEMC). PANAMA: 2 specimens, Almirante, IV.[19]43, no collector (FMNH); 12 specimens, Canal Zone, Barro Colorado I., under bark, 14.I.1959, H.S. Dybas coll. (FMNH); 1 specimen, the same locality and collector, berlese, 25.I.1959 coll. (FMNH); 1 specimen, the same locality and collector, under bark, 27.I.1959 (FMNH); 4 specimens, the same locality and collector, berlese, 28.I.1959 (FMNH); 10 specimens, the same locality and collector, under bark, 4. IV.1959 (FMNH); 1 specimen, the same locality, under bark, 3.II.1976, A. Newton coll. (FMNH); 3 specimens, the same locality and collector, under bark, 8–25.II.1976 (FMNH); 2 specimens, the same locality and collector, 10.II.1976 (FMNH); 5 specimens, the same locality and collector, 16–22.II.1976 (FMNH); 1 specimen, the same locality and collector, 25.II.1976 (FMNH); 3 specimens, no locality, date and collector (FMNH); 1 specimen, the same locality, fermenting bark, 26.VIII.1978, Q. D. Wheeler coll. (FMNH); Bocas del Toro: 1 specimen, Almirante, dam on Nigua Creek, berlese, 23.III.1959, H. S. Dybas coll. (FMNH); 4 specimens, the same locality and collector, 25.III.1959 (FMNH); 1 specimen, the same locality and collector, 27.III.1959 (FMNH); Chiriquí: 1 specimen, no locality, VII.1930, David coll. (FMNH); 3 specimens, Chiriqui, no date and collector (ZMHB); 2 specimens, Pto Armuelles, no date and collector (FMNH); Veraguas: 2 specimens, Gabina, VI.[19]38, no collector (FMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd km 6.1, 09
070 N 79
450 W, 40 m, flight intercept trap, 7–21.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, 10–15 km N jct. Escobal and Piña Rds., 30 m, under bark, 02.VI.1996, J. Ashe and R. Brooks coll. (SEMC); Panamá: 2 specimens, Las Cumbres, 15.II.1959, H. S Dybas coll. (FMNH); 2 specimens, Fort San Lorenzo, 20 m, 09
000 N 79
380 W, under bark, 15.V.1995, J. and A. Ashe coll. (SEMC); 3 specimens, Old Plantation Trail km 4.0, Parque Soberania, 175 m, 1–2.VII.1995, A. Gillogly coll. (SEMC); 1 specimen, Old Plantation Trail km 3.5, Parque Soberania, 150 m, 4.VII.1995, A. Gillogly coll. (SEMC). GUADELOUPE: 3 specimens, no locality, date and collector (1 NMW, 1 IRSNB, 1 AMNH); 2 specimens, Trois Rivieres, no date, Dufau coll. (ZMHB); 1 specimen, no locality and date, Goubeyre coll. (AMNH). SAINT VINCENT AND THE GRENADINES: Saint Vincent: 1 specimen, Leeward side, no date, H. H. Smith (IRSNB). GRENADA: 2 specimens, Balthazar, Windward side, no date, H. H. Smith (1 NMW, 1 IRSNB). TRINIDAD AND TOBAGO: Trinidad: 1 specimen, no locality and date, Thailer coll. (FMNH); 1 specimen, no locality, 17.XI.[19]08, Peetz coll. (FMNH); 7 specimens, Balandra Bay, 19.IV.[19]22, L. R. Reynold coll. (FMNH); 5 specimens, the same locality and date, F. Psota coll. (FMNH); 1 Revision of Piestus with remarks on related genera Invertebrate Systematics 575 specimen, the same locality, V.[19]22, L. R. Reynold coll. (FMNH); 10 specimens, the same locality and date, no collector (FMNH); 5 specimens, the same locality, 23.V.[19]22, L. R. Reynold coll. (FMNH); 7 specimens, the same locality and date, F. Psota coll. (FMNH); 3 specimens, the same locality, 24.V. [19]22, L. R. Reynold coll. (FMNH); 1 specimen, the same locality 13.X[19]34, no collector (FMNH); 1 specimen, Arima Vallley, 800–1200ft., 10–22. II.1964, J. G. Rozen and P. Wygodzinsky coll. (AMNH). COLOMBIA: 7 specimens, no locality, date and collector (ZMHB); 1 specimen, R. Dagua, IX– XII.[18]94, W. Rosenberg coll. (IRSNB); Magdalena: 1 specimen, Sierra Nevada de Santa Marta, Puente de los Claves, 15 km. E. Pueblo Bello, 500 m, under bark, 13.VI.1968, B. Malkin coll. (FMNH); Cesar: 14 specimens, Puento de Los Clavos, 15 km E of Pueblo Bello, Sierra Nevada de Santa Marta, 500 m, under bark, 13.VI.1968, the same collector (FMNH); Boyacá: 1 specimen, Muzo, no date and collector (FMNH); Meta: 1 specimen, Villavicencio, 12. VII.1938, H. Dybas coll. (FMNH); Putumayo: 1 specimen, Santa Rosa, headwaters of Rio San Miguel, under bark, 16–10.X.1970, B. Malkin and P. Burchard coll. (FMNH). VENEZUELA: 1 specimen, no locality, 1858, Moritz coll. (NMW); 2 specimens, St. Theresa, 13.V.[19]22, F. Psota coll. (FMNH); 2 specimens, Las Trincheras, VI.[19]22, F. Psota coll. (FMNH); Bolívar: 3 specimens, between Sta. Elena de Uairen and Icabaró, 30.VII.1987, M. A. Ivie coll. (SEMC). GUYANA: Demerara-Mahaica: 2 specimens, Georgetown, 41 km, SW Soesdyke rd. to Linden, 6
280 2400 N 58
130 4800 W, 15 m, under bark, 10.VI.2001, R. Brooks and Z. Falin coll. (SEMC); Potaro-Siparuni: 1 specimen, Tumatumari, no date and collector (AMNH). FRENCH GUIANA: SaintLaurent-du-Maroni: 2 specimens, Maripassoula, Lawa River, 7.XII.1963, under bark, B. Malkin coll. (FMNH); 2 specimens, Les Eaux Claires, 3.XI.1995, A. Berkov coll. (AMNH); 3 specimens, 3.5mi N Saül, Les Eaux Claires, IV.[19]96, A. Berkov coll. (AMNH); Cayenne: 1 specimen, Chimbo, VIII.[18]97, Rosenberg coll. (IRSNB). ECUADOR: Sucumbios: 1 specimen, Sacha Lodge, 0
280 1400 S 76
270 3500 W, 270 m, fungus covered log, 23.III.1999, R. Brooks coll. (SEMC); Pichincha: 1 specimen, 10 km SE Sto.Domingo de los Colorados, 2.V.1978, O’Brien and Marshall coll. (FMNH); Chimborazo: 3 specimens, Chimbo, VIII.[18]97, Rosemberg coll. (FMNH); Pastaza: 1 specimen, Ashuara indian village on Rio Macuma nr. Rio Morona, beating dry foliage, 11–16. VII.1971, B. Malkin coll. (FMNH); 5 specimens, Cusuimi on Rio Cusuimi, 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH). PERU: 6 specimens, no locality and collector, 23.V.1936 (3 AMNH, 3 FMNH); Loreto: 1 specimen, Estiron, Rio Ampiacu, under corticals, 15–22.V.1966, B. Malkin coll. (FMNH); 1 specimen, 1.5 km N Teniente Lopez, 2
35.660 S 76
06.920 W, 210–240 m, flight intercept trap, 18.VII.1993, R. Leschen coll. (SEMC); Cusco: 1 specimen, Callanga, no date and collector (IRSNB); Junín: 1 specimen, Jauja Prov., Saní Beni 8 km E Satipo, 840 m, 6–9.XI.1935, F. Woytkwski coll. (SEMC); 1 specimen, San Ramón de Pangoa, 40 km SE Satipo, 750 m, 24.II.1972, R. T. and J. C. Schuh coll. (AMNH). BOLIVIA: 1 specimen, Yuracaris, no date and collector (IRSNB); Beni: 1 specimen, Chacobo Indian, Village on Rio Benicito, 66
–12
200 18–27.VII.1960, B. Malkin coll. (FMNH); Cochabamba: 1 specimen, Regian Chaparé, 400 m, 20.XII.1949, R. Zischka coll. (NMW); 1 specimen, 20 mi SW Villa Tunari, 2.IV.1978, L. and C. W. O’Brien and G. B. Marshall coll. (FMNH); Santa Cruz: 11 specimens, Santa Cruz de la Sierra, 24–25.VIII.1960, B. Malkin coll. (FMNH); 24 specimens, the same locality and collector, 23–25.IX.1960 (FMNH); 1 specimen, 14 mi SW Porta Chuelo, uv light trap, 24–III.[19]78, G .B. Marshall coll. (FMNH); 1 specimen, 10 mi W. Puerto Chuelo, uv light trap, 26.III.1978, Marshall coll. (FMNH); 16 specimens, Ichilo, Cafetal by Rio Quebrada Palometilla, 5.VIII.1990, P. Parrillo and P. Bettella coll. (FMNH); 2 specimens, the same locality and collector, 6.VIII.1990 (FMNH); 1 specimen, 3.7 km SSE Buena Vista, Hotel Flora Y Fauna, 400–17
29.950 S 63
33.150 W, 440 m, under bark, 5.XI.2002, R. Leschen (SEMC); 1 specimen, the same locality and collector, at light, 6.XI.2002 (SEMC). BRAZIL: 2 specimens, Pebas, no date and collector (IRSNB); 13 specimens, no locality, date and collector (1 MNRJ, 5 ZMHB, 2 IRSNB, 5 FMNH); 1 specimen, Petropolis, II.1850, no collector (FMNH); 1 specimen, the same locality, no date, H. Schulz coll. (NMW); 1 specimen, Guarujá, II.[19]18, A. Bierig coll. (FMNH); Pará: 1 specimen, no locality, date and collector (IRSNB); 6 specimens, no locality and date, Baker coll. (5 FMNH, 1 NMW); 1 specimen, Caninde, Rio Gurupi, Gurupi Uma Maranhao, 50 km E Caninde, under bark, 6.IV.1963, B. Malkin coll. (FMNH); 1 specimen, Tiriós, Alto Pará d’Oeste, I–II.1963, Machado and Pereira coll. (MZSP); 5 specimens, Caninde, Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH); 1 specimen, the same locality and collector, 7–11.IV.1963 (FMNH); 1 specimen, Aldeia Aracu, Igarape, GurupuUmu, Maranhao 50 km E of Caninde, under bark, V.1963, B. Malkin coll. (FMNH); Ceará: 3 specimens, Serra Ibiapaba, Ubajara, VIII.1953, Ulisses coll. (AMBC); Pernambuco: 1 specimen, Pery-Pery, no date and collector (IRSNB); Bahia: 8 specimens, no locality, date and collector (1 IRSNB, 5 FMNH, 2 NMW); Mato Grosso: 7 specimens, Barra do Tapirape, under bark, 8.XI.1964, B. Malkin coll. (FMNH); Goiás: 1 specimen, S. Isabel do Morro, I. Bananal, VI.1961, M. Alvarenga coll. (MNRJ); Minas Gerais: 4 specimens, Viçosa, IX.1943, Wygodzinsky coll. (MNRJ); Espirito Santo: 1 specimen, Rio Bonito Conde, 21.VI.[19]28, Leopoldina coll. (ZMHB); Rio de Janeiro: 1 specimen, Tijuca, I.1857, J. Gray coll. (IRSNB); 1 specimen, Petrópolis, no date and collector (IRSNB); 12 specimens, no locality and date, Fry coll. (2 ZMHB, 10 FMNH); 4 specimens, no locality, 21.II.1932, D. Mendes coll. (MNRJ); 11 specimens, Trapicheiro, 30.IX.1961, H.Schubart coll. (MNRJ); 2 specimens, the same locality and collector, 10.XI.1961 (INPA); 1 specimen, Pq. da Cidade, VI.1995, A. Bello coll. (AMBC); São Paulo: 2 specimens, no locality and date, Mráz coll. (FMNH); 2 specimens, Santos, no date, J. Metz coll. (1 FMNH, 1 ZMHB); 1 specimen, the same locality, no date, Brauns coll. (NMW); 3 specimens, 27.IX.[19]21, A. Bierig coll. (FMNH); Santa Catarina: 12 specimens, no locality, date and collector (1 FMNH, 1 MZSP, 2 DZUP, 3 NMW, 5 ZMHB); 2 specimens, no locality and date, Lüderwaldt coll. (1 FMNH, 1 NMW); 1 specimen, Blumenau, no date and collector (NMW); 6 specimens, the same locality, Hetschko coll. (NMW); 1 specimen, no locality and date, Fry coll. (FMNH); 17 specimens, N. Teutonia, no date, Plaumann coll. (FMNH); 5 specimens, the same locality and collector, II.1950 (FMNH); 3 specimens, Nova Teutonia, 27
110 B 52
230 L., no date, F. Plaumann coll. (2 FMNH, MNRJ); 2 specimens, the same locality and collector, VIII.1935 (MNRJ); 2 the same locality, no date and collector (FMNH); 2 specimens, blumenau, no date, Hetschko coll. (NMW); 2 specimens, Nova Teutonia, VII.1935, B. Pohl coll. (MZSP); 1 specimen, Corupa, XI.1948, Humbolt coll. (AMNH). PARAGUAY: 7 specimens, no locality, date and collector (1 IRSNB, 6 ZMHB); 1 specimen, no locality, 1885, Drake coll. (FMNH); 3 specimens, no locality and date, Fiebrig coll. (ZMHB); 26 specimens, San Luis, no date, Reimoser coll. (NMW); Asunción: 11 specimens, Asunción, no date, B. Podtiaguin coll. (AMNH); Paraguarí: 1 specimen, Sapucai, 25
400 S 56
550 W, 190 m, 18.VI.1994, U. Drechsel coll. (SEMC); Guairá: 3 specimens, Villa Rica, Sta. Barbara, I.1926, Schade coll. (1 NMW, 2 DZUP); 1 specimen, Calle Florida, 11.IV.1992, U. Dreschel coll. (SEMC); 1 specimen, the same locality and collector, 17.IX.1994 (SEMC); Caazapá: 1 specimen, Hermosa, prop. Sosa family San Rafael Reserve, 26
190 1500 S 55
440 5500 W, 90 m, under bark, 5.XII.2000, Z. H. Falin coll. (SEMC); 2 specimens, the same locality, data e coletor, fermenting tree wound (SEMC); Itapúa: 6 specimens, Hohenau, Alto Parana, no date, H. Jacob coll. (NMW); 1 specimen, the same locality and collector, 23.V.[19]32 (NMW); 1 specimen, the same locality and collector V.1932 (FMNH). ARGENTINA: 1 specimen, no locality, date and collector (IRSNB); 1 specimen, Rio Parana, Territorio das Missiones, no date and collector (FMNH); Salta: 3 specimens, Tabacal, VI.1993, F. Schade coll. (AMNH); Tucumán: 2 specimens, no locality, 17. IX.1928, H. E. Box coll. (1 FMNH, 1 AMNH); 2 specimens, no locality, 23.X.1929, H. E. Box coll. (AMNH); Misiones: 3 specimens, no locality and date, M. Richter coll. (FMNH); 1 specimen, Iguazu, luz, 20.II.1967, R. Foerster coll. (ZMHB); 1 specimen, 20 km SE Puerto Iguazu, 220 m, forest uv light, 6. I.1991, S. and J. Peck coll. (FMNH). Undetermined country: 14 specimens, no locality, date and collector (2 NMW, 3 IRSNB, 1 FMNH, 8 ZMHB). Piestus extimus Sharp, 1887 UNITED STATES OF AMERICA: Arizona: 2 specimens, Santa Rita Mts., 5 to 8000ft., no date, F. H. Snow coll. (SEMC); 2 specimens, St. Catalina Mts., Molino Basin, dead sotol, 14.III.1970, K. Stephan coll. (FMNH); 1 specimen, Santa Rita Mts., Madera Cyn., dead sotol, 15.III.1970, K. Stephan coll. 576 Invertebrate Systematics E. Caron et al. (FMNH); 3 specimens, Dragon Mts. Wood Cyn., 29.IV.1972, K. Stephan coll. (FMNH); 1 specimen, Proctor Ranch, Madera Cyn., Pima Co., rotting barrel cactus, 17.III.1973, D. S. Chandler coll. (FMNH); 4 specimens, the same locality, dead sotol, IV.1973, R. Lenczy coll. (1 DZUP, 3 AMNH); 2 specimens, Pima, IV.1973, R. Leczy coll. (FMNH); 4 specimens, PimaCo., Madera Cn., dead sotol, IV.1973, Lenczy coll. (AMNH); 1 specimen, Santa Cruz co., vic. Pena Blanca Lk, 14.XI.1998, Morris and Warmer coll. (FMNH). MEXICO: 11 specimens, no locality, date and collector (2 FMNH, 4 IRSNB, 2 ZMHB, 3 NMW); 1 specimen, no locality, 30.VI.1897, C. Höge coll. (NMW); Durango: 3 specimens, 2 mi NW Nombre de Dios, 12.VII.1975, L. E. Watrous coll. (FMNH); Nayarit: 1 specimen, Tepic, 2–7.VIII.1947, B. Malkin coll. (AMNH); 1 specimen, San Blas, 4–5.VIII.1947, the same collector (AMNH); 1 specimen, Tepic, no date and collector (FMNH); Jalisco: 16 specimens, 1 Mi.S.W.La Resolana, 20.XI.1950, R. F. Smith coll. (1 DZUP, 15 AMNH); 1 specimen, Las Jarillas, 62 km S, P. Vallarta, under bark, 22.VII.1984, Chemsak and Doyen coll. (FMNH); 1 specimen, Chamela Biol. Stn., flight intercept trap, 14.VII.1989, R. W. Brooks coll. (SEMC); 2 specimens, Atoyac, Isla Chica, 1300 m, 27.X.1997, J. L. Navarrete coll. (CZUG); 2 specimens, Unión de Tula, camino a La MARTINIQUE, 1500 m, 9.I.1998, H. Fierros and J. L. Navarrete-Heredia coll. (CZUC); 2 specimens, Casimiro Castillo, Amoyo Tacubaya, 600 m, 11.VII.2005, H. E. Fierros-López (CZUG); México: 2 specimens, Tejupilco, Temescaltepec, 15.VI.[19]33, H. E. Hinton, R. I. Usinger coll. (AMNH); 1 specimen, Real de Arriba, Temescaltepec, 19.VI.[19]33, H.E. Hinton, R. I. Usinger coll. (AMNH); Guanajuato: 1 specimen, Guanajuato, no date and collector (IRSNB); Oaxaca: 2 specimens, Ocotlan, no date and collector (ZMHB); 3 specimens, Zaachila, 4500ft., 21.VII.[19]37, A. R. Mead coll. (FMNH); 8 specimens, Cuicatlan, 27.VII.[19]37, M. Embary coll. (FMNH); 2 specimens, 2.1 mi NW Totolapam, 3500ft., on rotting cacti columnar Opuntia, 6.IX.1973, A. Newton coll. (FMNH); 3 specimens, 14.0 mi E Chacalpa, Hwy 175, montane tropical, (‘Correction: should be 14.0 mi S Suchixtepec, 16
050 N, 96
280 W), 1700 m, under bark fermenting sap, 30.IV.1977, J. S. Ashe coll. (FMNH). Undetermined country: 2 specimens, no locality, date and collector (NMW). Piestus buquetii Fauvel, 1864 PERU: Cusco: 7 specimens, Torentoy Canyon base of Machu Picchu, 2000 m, under bark, 20.VI.1964, B. Malkin coll. (1 DZUP, 6 FMNH). BOLIVIA: 3 specimens, Yuracaris, no date and collector (IRSNB); 1 specimen, no locality and date, N. Hobngren coll. (NHRS); 1 specimen, Rio Beni, La Paz Reyes, 1891, no collector (FMNH); La Paz: 1 specimen, 9.4 km E Chulumani, Apa Apa Ecoll. Reserve, 16
20.990 S 67
30.300 W, 2100–2400 m, fungus covered log, 19.I.2001, J. S. Ashe and R. S. Hanley coll. (SEMC); 1 specimen, La Paz 6 km W Yanacachi, 16
25.950 S 67
47.070 W, 2150–2350 m, fungus covered log, 24.I.2001, J. S. Ashe and R. S. Hanley coll. (SEMC); 1 specimen, Coroico, 16
11.900 S 67
43.440 W, 1850 m, under bark, 27.I.2001, J. S. Ashe and R. S. Hanley (SEMC); Beni: 1 specimen, Yuracaris, no date and collector (FMNH); 1 specimen, Yuracaris, 6.II.1900, no collector (FMNH). Piestus filicornis Fauvel, 1902 BOLIVIA: Cochabamba: 1 specimen, Cochabamba, 105 km E Yungas, nr. Rio Carmen Mayu, Villa Tunari Rd., 17
80 5100 S 65
430 5000 W, 1750 m, flight intercept trap, 1–6.II.1999, R. Hanley coll. (FMNH); 1 specimen, the same locality, 1–6.II.1999, F. Genier coll. (SEMC); 1 specimen, Cochabamba, 109 km E Yungas, Villa Tunari Rd., 17
80 5000 S 65
420 2900 W, 1480 m, flight intercept trap, 1–6.II.1999, R. Hanley coll. (SEMC); 1 specimen, Cochabamba, 105 km E Yungas, nr. Río Carmen Mayu, Villa Tunari Rd., 17
80 5100 S 65
430 5000 W, 1750 m, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC); 1 specimen, Cochabamba, 109 km E, Yungas, Villa Tunari Rd., 17
80 5200 S 65
420 5400 W, 1400 m, flight intercept trap, 8–12.II.1999, F. Genier coll. (DZUP); 1 specimen, the same locality and collector, 6.II–8.VI.1999 (SEMC). Piestus niger Fauvel, 1864 MEXICO: 1 specimen, no locality and date, Reitter coll. (FMNH); 1 specimen, ilegible locality, no date, J. Flohr coll. (ZMHB); 1 specimen, no locality and date, Koltze coll. (FMNH); Veracruz: 7 specimens, Jalapa, no date, Flohr coll. (2 DZUP, 4 ZMHB, 1 NMW); 1 specimen, Fortin, 26.X.[1]961, Pereira and Halfter coll. (MZSP); Oaxaca: 1 specimen, 1590m Pueblo Nuevo, FIT, 1–12.VIII.1986, H. and A. Howden coll. (FMNH). GUATEMALA: 1 specimen, no locality, date and collector (NMW); Guatemala: 2 specimens, Guatemala City, no date, Champion coll. (FMNH). COSTA RICA: Guanacaste: 3 specimens, Cacao Biological Station, 10
550 3800 N 85
270 700 W, 1050 m, under bark, 10.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 4 specimens, the same locality and collector, 11.VII.2000 (SEMC). PANAMA: 1 specimen, no locality, date and collector (ZMHB). Piestus minutus Erichson, 1840 MEXICO: Nayarit: 1 specimen, El Cora, Tepic, no date and collector (ZMHB); BELIZE: Orange Walk: 2 specimens, Rio Bravo Conserv. Area, La Milpa transect, vic. La Milpa Archeol. Site, under Pseudobombax bark, 7–14.IX.1995, P. Kovarik coll. (FMNH); 1 specimen, Rio Bravo Conservation Area, La Milpa Ruins, 17
500 N 89
020 W, FITs, 15–25.V.1997, C. Carlton coll. (SEMC); 1 specimen, RBCA La Milpa Field Stat., FIT, 15–25.V.1997, the same collector (SEMC). GUATEMALA: 2 specimens, San Juan, Vera Paz, no date, Champion coll. (FMNH); Alta Verapaz: 1 specimen, Lanquin, 1000ft., under bark, 4.VI.1948, R. D. Mitchell coll. (FMNH); Izabal: 2 specimens, Quiriqua Ruins, 8.II.1996, D. Brzoska coll. (SEMC); Escuintla: 1 specimen, Finca El Zapote, 2400ft., under bark, 8.VII.[19]48, R. D. Mitchell coll. (FMNH); 2 specimens, the same locality and collector, 9.VII.[19]48 (FMNH); 1 specimen, the same locality and collector, 10.VII.[19]48 (FMNH); 1 specimen, the same locality and collector, 13.VII.[19]48 (FMNH); 1 the same locality and collector, 2000ft., 20.VII.1948 (FMNH). EL SALVADOR: Usulatán: 1 specimen, 3mi. E. El Trinufo, 1200’, 9.VI.1974, O’Briens and Marshall coll. (FMNH). HONDURAS: Atlántida: 2 specimens, Lancetilla Bot. Grd., Tela, 15
460 N, 87
270 W, 10 m, under bark, 23.VI.1994, J. Ashe and R. Brooks coll. (SEMC). NICARAGUA: Granada: 2 specimens, Res. Nat. Volcan Mombacho, 910m 11
50.050 N 85
58.830 W, flight intercept trap, 1–5.VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC); Río San Juan: 1 specimen, 60 km SE San Carlos, Refugio Bartola, 10
58.400 N 84
20.300 W, 100 m, Heliconia fruits, 27.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC); 1 specimen, the same locality and collector, 28.V.2002 (SEMC). COSTA RICA: 1 specimen, no locality and date, B. Haas coll. (FMNH); Guanacaste: 1 specimen, Lomas Barbudal Biological Reserve, 10
300 2200 N 85
220 1500 W, 17m, on and under bark, 15.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); Heredia: 3 specimens, La Selva Biol. Station nr. Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); 1 specimen, the same locality, 25.III.1988, H. A. Hespenheide coll. (SEMC); 8 specimens, 3 km S. Puerto Viejo, OTSLa Selva, 100 m, X.1992, P. Hanson coll. (SEMC); 1 specimen, La Selva Biol. Sta., 3 km S Pto. Viejo, 10
260 N 84
010 W, malaise trap, 3–8.VIII.1992, G. Wright coll. (SEMC); 3 specimens, the same locality, IX.1993, P. Hanson coll. (SEMC); San José: 1 specimen, no locality and collector, 1936 (FMNH); 1 specimen, La Caja, 1942, no collector (FMNH); Cartago: 2 specimens, Turialba, 800 m, no date and collector (ZMHB); Puntarenas: 1 specimen, Revision of Piestus with remarks on related genera Invertebrate Systematics 577 Surrubres, 300’, no date, A. Heyne, (FMNH); 32 specimens, UVITA, 3.XI.–22.XI.1994, H. Forster coll. (NMW); 5 specimens, Corcovado National Park, Sirena Stn. junc. Guanacaste-Sirena Trails, 8
280 5500 N 83
350 4600 W, 5 m, under bark, 30.VI.2000, Z. H. Falin coll. (SEMC). PANAMA: 20 specimens, Canal Zone, Barro Colorado Island, bark and debris from fallen tree, 14.I.1959, H. S. Dybas coll. (FMNH); 29 specimens, the same locality and collector, fermenting fibrous log at light, 16.I.1959 (FMNH); 21 specimens, the same locality, Fairchild Trail, bark and under bark debris from fallen tree, berlese, 28. I.1959, H. S. Dybas coll. (FMNH); 21 specimens, the same locality and collector, Zetek Trail, fibrous debris in tree trunk, 4.IV.1959 (FMNH); 1 specimen, the same locality, Albrook Forest Site, 100ft., 20.I.1968, R. Hutton coll. (AMNH); 3 specimens, the same locality, 4mi. NW Gamboa, under bark, 23.II.1975, J.F. Lawrence coll. (FMNH); 2 specimens, the same locality, under bark rotting log, 3.II.1976, A. Newton coll. (FMNH); 1 specimen, the same locality and collector, under bark rotting logs, 10.II.1976 (FMNH); 1 specimen, the same locality and collector, under bark, 8–25.II.1976 (FMNH); 3 specimens, the same locality, 28.VII.1978, Q. D. Wheeler coll. (FMNH); 4 specimens, the same locality, under bark fermenting, 16–22.II.1976, A. Newton coll. (FMNH); 3 specimens, the same locality and collector, under bark rotting logs, 25.II.1976 (FMNH); 7 specimens, Canal Zone, Madden Dam, under bark, 12.VI.1976, A. Newton coll. (FMNH); 1 specimen, Ft. Gulick, at lights, 18.XI.1978, H. J. Haslan coll. (FMNH); 1 specimen, Canal Zone, Gamboa-Pipeline Rd KM7, flight intercept trap, 20.V–11.VI.1996, S. Lingafelter coll. (SEMC); Chiriquí: 2 specimens, Rio Armuelles, 1930, no collector (FMNH); 1 specimen, 10.5 km NE Caldera, 08
420 N 82
190 W, 340 m, tree fall litter, 24.V.1995, J. and A. Ashe coll. (SEMC); Veraguas: 6 specimens, Gabina, IX.[19]38, no collector (FMNH); 5 specimens, Cerro Santa Rita, chips from cut end of log, 9.II.1959, H. S. Dybas coll. (FMNH); Colón: 1 specimen, no locality, 28.I.1925, G. Bateson coll. (FMNH); 1 specimen, Parque Nac. Soberania, Pipeline Rd. Km 0.8, 09
070 N 79
450 W, 40 m, collected in Heliconia mariae, 2.VI.1995, Ashe and Brooks coll. (SEMC); 3 specimens, Parque Nac. Soberania, Pipeline Rd. Km 6.1, 09
070 N 79
450 W, 40 m, flight intercept trap, 31.V–2.VI.1995, the same collector (SEMC); 1 specimen, 10–15 km N jct. Escobal and Piña Rds, ~30 m, under bark, 02.VI.1996, the same collector (SEMC); 2 specimens, Santa Rita Ridge, 400m, 17 km E Transisthmus Hwy, 16.VI.1996, A. Gillogly coll. (SEMC); Panamá: 1 specimen, Círicito, Canal Zone, 13.III.[19]30, coll. (AMNH); 24 specimens, Canal Zone, Las Cumbres, compost heap, 15.II.1959, H. S. Dybas coll. (FMNH); 2 specimens, Altos de Majé, sifting under tree bark, 6–15.X.1975, D. S. Chandler coll. (FMNH); 1 specimen, Cerro Azul, ~2000’, wet debris small forest stream, 21.II.1976, A. Newton coll. (FMNH); 2 specimens, Cerro Trinidad 400 m, 6.I.1980, Stockwell coll. (FMNH); 2 specimens, Pipeline Rd., flight intercept trap, 16–21.VI.1993, S. Lingfelter coll. (SEMC); 9 specimens, 09
050 N 79
400 W, Old Plantation Rd. 6.9 km, S. Gamboa, 80 m, fungusy log, 3.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 7 specimens, Old Plantation Trail Km 4.0, Parque Soberania, 175 m, 1–2.VII.1995, A. Gillogly coll. (SEMC); 1 specimen, Barro Colorado Island, FIT, 9
11N 79
51W, 1–5.VII.2000, S. Chatzimanolis coll. (SEMC); 2 specimens, Old Plantation Trail km 3.5, Parque Soberania, 150 m, 4.VII.1995, A. Gillogly coll. (SEMC); Darién: 2 specimens, Rio Tuquesa, 8
270 N 77
440 W, 20 m, 18.II.1981, Stockwell coll. (FMNH); 1 specimen, Cana, 7
480 N 77
390 W, 450 m, 24. IX.[19]82 Stockwell coll. (FMNH). DOMINICA: 1 specimen, Salibiaj, under rotten banana leaves on ground, 28.I.1968, B. Malkin coll. (FMNH). GRENADA: 4 specimens, Balthazar, Windward side, no date, H. H. Smith coll. (3 NMW, 1 IRSNB). TRINIDAD AND TOBAGO: Trinidad: 1 specimen, no locality, date ans collector (FMNH); 1 specimen, no locality, 1904, F. Birch coll. (FMNH); 76 specimens, Balandra Bay, IV.1922, F. Psota coll. (FMNH); 49 specimens, the same locality and collector, 23.V.[19]22 (FMNH); 29 specimens, the same locality and date, L. R. Reynold coll. (FMNH); 22 specimens, the same locality and collector, 24.V.[19]22 (FMNH); 1 specimen, W.I. Trinidad, 6 Mi. NE Arima, L’Oranga Valley, 15.VI.1973, no collector (AMNH); 1 specimen, W.I. Trinidad, Asa Wright N.C., under bark, 8.VI.1991, D. Brzoska coll. (SEMC). COLOMBIA: 1 specimen, the same locality, data e coletor (NMW); 1 specimen, Villavicencio, 11.VII.[19]38, H. S. Dybas coll. (FMNH); 3 specimens, Puento de Los Clavos, 15 km E of Pueblo Bello, Sierra Nevada de Santa Marta, 500 m, under bark of log, 13.VI.1968, B. Malkin coll. (FMNH); Magdalena: 1 specimen, PNN Tayrona Cañaveral, 11
200 N 74
20 W, 30 m, Malaise, 30.VIII.–19.IX.[20]00, R. Henriquez coll. (SEMC); 2 specimens, the same locality and collector, 50 m, Malaise, 29.IX–17.X.2000 (SEMC); 1 specimen, the same locality and collector, 2–22.I.2001, R. Henriquez coll. (SEMC); Amazonas: 1 specimen, PNN Amacayacu Matamata, 3
410 S 70
150 W, 150 m, Malaise, 04–28.VIII.2000, A. Parente coll. (SEMC); 1 specimen, the same locality and collector, 17.XII.2000–02.I.2001 (SEMC). VENEZUELA: 1 specimen, no locality, date and collector (IRSNB); 1 specimen, Carupapo, no date and collector (IRSNB); 2 specimens, no locality, 1858, D’Moritz (1 FMNH, 1 NMW); 1 specimen, Tropical Research Sta., Rancho Grande nr. Maracay, 13.IV.1942, no collector (AMNH); Aragua: 1 specimen, Parque Nac. Henri Pittier 4 km N El Limon, 700 m, on large dead tree, 19.VI.1987, M. A. Ivie coll. (SEMC). GUYANA: 1 specimen, Regio 8, Iwokrama Forest, Turtle Mt. base camp, 4
430 500 N 58
430 500 W, 50 m, under bark, 1.VI.2001, R. Brooks and Z. Falin (SEMC); 14 specimens, the same locality and collector, 4. VI.2001 (SEMC); 2 specimens, Region 8 Iwokrama Forest, Kabocalli Field Stn, 4
170 400 N 58
300 3500 W, 60 m, under bark, 5.VI.2001, R. Brooks and Z.Falin (SEMC). FRENCH GUIANA: 1 specimen, Riv. Lumier, no date and collector (IRSNB); Saint-Laurent-du-Maroni: 1 specimen, Les Eaux Claires, 3. XI.1995, A. Berkov coll. (AMNH); 5 specimens, 3.5mi N Saül, Les Eaux Claires, IV.[19]96, the same collector (AMNH); 11 specimens, Saül, under bark, III.–IV.1999, the same collector (AMNH); Cayenne: 1 specimen, Camopi, no date and collector (IRSNB); 3 specimens, Roura, 55.4 km SSE, Kaw marshes, 4
290 5800 N 52
30 000 W, 40 m, under fermenting bark, 11.VI.1997, J. Ashe and R. Brooks coll. (SEMC). ECUADOR: Sucumbios: 6 specimens, Sacha Lodge, 0.5
S 76.5
W, 270 m, malaise, 12–22.II.1994, Hibbs coll. (SEMC); 2 specimens, the same locality and collector, 4–14.III.1994 (SEMC); 3 specimens, the same locality and collector, 14–24.III.1994 (SEMC); 2 specimens, the same locality and collector, 3–13.IV.1994 (SEMC); 3 specimens, the same locality and collector, 24.V–3.VI.1994 (SEMC); 2 specimens, the same locality and collector, 13–23.VI.1994 (SEMC); 1 specimen, the same locality and collector, 13–25.VII.1994 (SEMC); 1 specimen, the same locality and collector, 25.VII–3.VIII.1994 (SEMC); 1 specimen, the same locality and collector, 10–21.X.1994 (SEMC); 3 specimens, Sacha Lodge, 0
280 1400 S 76
270 3500 W, 270 m, under bark, 21–24.III.1999, R. Brooks coll. (SEMC); 2 specimens, the same locality and collector, 22.III.1999 (SEMC); 1 specimen, the same locality and collector, 24.III.1999 (SEMC); Pichincha: 1 specimen, Rio Palenque Science Center, 0
360 000 S 79
210 000 W, 200 m, malaise trap, 25.III–6.VII.1996, P. Hibbs coll. (SEMC); Napo: 2 specimens, 18 km S. Tena, 28.IV.1978, C. W. and L. B. O’Brien and Marshall coll. (FMNH); 3 specimens, Jatun Sacha Biol. Station, 21 km E. Puerto Napo, 400 m, virgin rain forest, F.I.T, 15.VII.1994, Levy and Génier coll. (SEMC); 1 specimen, the same locality and collector, 18.VII.1994 (SEMC); 1 specimen, Yuturi Lodge, Rio Napo, 0
320 5400 S 76
20 1800 W, 270 m, flight intercept trap, 20–21.III.1999, R. Brooks and D. Brzoska coll. (SEMC); Pastaza: 1 specimen, Pomona, on Rio Pastaza, 2000ft., 9.VIII.1960, T. Sabine coll. (FMNH); 5 specimens, Cusuimi on Rio Cusuimi, 150 km SE Puyo, 300 m, under bark, 19–23.VII.1971, B. Malkin coll. (FMNH). PERU: Cusco: 1 specimen, Callanga, no date and collector (IRSNB); Madre de Dios: 1 specimen, Tambopata, 15 km NE Pto. Maldonado, 200 m, at light, 10.VI.1989. J. S. Ashe and R. A. Leschen coll. (SEMC); 1 specimen, the same locality, Maldonado, Reserva Cuzco Amazónica, 12
330 S 69
030 W, 200 m, 20.VI.1989, R.A. Leschen coll. (SEMC); 1 specimen, the same locality, fight intercept, 24.VI.1989, J. S. Ashe and R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, Liana fruit fall, 25.VI.1989 (SEMC); 1 specimen, the same locality, berlese in fruit fall, 25. VI.1989, R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, 1.VII.1989 (SEMC); 1 specimen, the same locality and collector, 9. VII.1989 (SEMC); 1 specimen, the same locality and collector, Swamp trail, flight intercept trap, 16.VII.1989 (SEMC); 1 specimen, the same locality and collector, 17.VII.1989 (SEMC); 1 specimen, the same locality and collector, 19.VII.1989 (SEMC); 1 specimen, Manu, Parque Nac. Manu, Zona Res., Rio Manu, Cocha Juarez, trail nr. Manu, Lodge, flight intercept trap, 18–24.IX.1991, A. Hartman (FMNH); 1 specimen, Cocha Cashu Bio. Stn., Manu National 578 Invertebrate Systematics E. Caron et al. Park, 11
530 4500 S 71
240 2400 W, 350 m, flight intercept trap, 17–19.X.2000, R. Brooks coll. (SEMC); 1 specimen, the same locality, Cocha Salvador, Reserved Zone, Manu National Park, 12
00 1300 S 71
310 3600 W, 310 m, flight intercept trap, 20–21.X.2000, R. Brooks coll. (SEMC); 3 specimens, Pantiacolla Lodge, Monk Saki Trap, Alto Madre de Dios River, 12
390 2200 S 71
130 5500 W, 400 m, 25.X.2000, R. Brooks coll. (SEMC). BOLIVIA: 1 specimen, Yuracaris, no date and collector (IRSNB); Beni: 3 specimens, Chacobo Indian, Village on Rio Benícito, 66
–12
200 , under bark of fallen log, 28–31. VII.1960, B. Malkin coll. (FMNH); Cochabamba: 2 specimens, Cochabamba, 67.5 km NE, Est. Biol. Valle del Sajita, Univ. de San Simon,, 17
60 3300 S 64
470 5200 W, 300 m, flight intercept trap, 7–9.II.1999, F. Genier coll. (SEMC); 1 specimen, Cochabamba, 117 km E., Yungas Lagunitas, 1000 m, 17
60 2200 S 65
400 5700 W, flight intercept trap, 8–12.II.1999, F. Génier coll. (SEMC); 1 specimen, Cochabamba, 109 km E., Yungas, Cochabamba-Villa Tunari Rd., 1480 m, 17
80 5000 S 65
420 2900 W, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC); Santa Cruz: 1 specimen, Sta. Cruz de la Sierra, 23–25.IX.1960, B. Malkin coll. (FMNH); 2 specimens, Ichilo, Cafetal, forest, under bark of log, 6.VIII.1990, P. Parrilo coll. (FMNH). BRAZIL: 1 specimen, no locality, date and collector (NMW); 3 specimens, Pebas, no date and collector (IRSNB); Amazonas: 2 specimens, Manaus, no date and collector (NMW); 1 specimen, no locality, date and collector (ZMHB); 1 specimen, Benjamin Constant, 18–28.IX.1962, K. Lenko coll. (MZSP); 1 specimen, Igarape Belem, nr. Rio Solimoes, 70 km E of Leticia, under bark, 18–28.V.1970, B. Malkin coll. (FMNH); Pará: 3 specimens, Caninde, Rio Gurupi, under bark, 6.IV.1963, B. Malkin coll. (FMNH); 3 specimens, Aldeia Aracu, Igarape, Gurupu-Umu, Maranhao, 50 km E of Caninde, under bark, V.1963, B. Malkin coll. (FMNH); Rondônia: 1 specimen, Porto Velho, Campus UNIR, 08
500 0400 S 68
560 3500 W, 19.IV.2006, J. A. Rafael and F. F. Xavier Fo coll. (INPA); 4 specimens, Ouro Preto do Oeste, Reserva INPA, Ceplac, 10
430 0000 S 62
140 4500 W, 22.IV.2006, J. A.Rafael and F. F. Xavier Fo coll. (INPA); Goiás: 1 specimen, Jatahy, no date and collector (IRSNB); São Paulo: 3 specimens, no locality and date, B. Pohl coll. (AMNH); Santa Catarina: 16 specimens, Nova Teutonia, no date, Plaumann coll. (10 FMNH, 3 AMNH, 3 NMW); 2 specimens, no locality and date, Klimsch coll. (1 FMNH, 1 NMW); 4 specimens, Nova Teutonia, VII.1935, B. Pohl coll. (DZUP); 2 specimens, the same locality, 27
110 B 52
230 L, VIII.1935, F. Plaumann coll. (MNRJ); 12 specimens, the same locality, II.1950, F. Plaumann coll. (FMNH); Rio Grande do Sul: 6 specimens, no locality, date and collector (5 FMNH, 1 AMNH). PARAGUAY: Guairá: 4 specimens, Villa Rica, no date and collector (DZUP); 10 specimens, the same locality, VII.1922, A. Schade coll. (NMW). ARGENTINA: Tucumán: 1 specimen, no locality, 17. IX.1928, H.E. Box coll. (AMNH); 7 specimens, no locality and the same collector, 23.X.1929 (AMNH). Undetermined country: 8 specimens, no locality, date and collector (5 ZMHB, 2 NMW, 1 IRSNB); 1 specimen, Guatemala, Nicaragua, Grenada, Mexico, no date and collector (IRSNB). Piestus penicillatus (Dalman, 1821) MEXICO: 1 specimen, Sierra de Durango, no date and collector (IRSNB). CUBA: 1 specimen, no locality and date, Gundlach coll. (ZMHB); 1 specimen, no locality and collector, B. Haas coll. (AMNH); 1 specimen, Havana, no date and collector (IRSNB); 12 specimens, the same locality, no date, Baker coll. (2 FMNH, 4 ZMHB, 4 SEMC, 2 IRSNB); 1 specimen, Ant. de los Baños, 9.I.[19]28 (ZMHB); 1 specimen, N. Sierra Maestra, X.[19]28, Rambousek coll. (FMNH); Pina del Río: 18 specimens, Aspiro, I.1933, A. Bierig coll. (FMNH); 24 specimens, the same locality and collector, 3.VI.1934 (2 DZUP, 22 FMNH); 1 specimen, the same locality and collector, XII.[19]34 (FMNH); 5 specimens, Sierra Rangel, XII.[19]34, the same collector (FMNH); 1 specimen, the same locality, X.1938, no collector (FMNH); La Habana: 8 specimens, Las Palmas, IX.[19]28, no collector (FMNH); 1 specimen, Baños, 9.I.[19]28, A. Bierig coll. (ZMHB); 11 specimens, Sierra Bonilla, 27.I.[19]29, the same collector (FMNH); 1 specimen, the same locality, 5.II.1933, A. Bierig coll. (FMNH); 9 specimens, the same locality, Cueva del Indio, 20.X.1932, A. Bierig coll. (FMNH); 5 specimens, Sítio Perdido, Jaruco, 4.IV.1932, A. Bierig coll. (FMNH); 3 specimens, Somorostro, Sra. de Tapaste, 14.VIII.1932, A. Bierig coll. (FMNH); Matanzas: 6 specimens, Catalina, VIII.1937, A. Bierig coll. (FMNH); Santiago de Cuba: 1 specimen, no locality, date and collector (ZMUC); Guantánamo: 1 specimen, 22.III.1914, C.T. Ramsden coll. (AMNH). DOMINICAN REPUBLIC: 3 specimens, St. Domingo, no date and collector (ZMHB, 2 IRSNB). PUERTO RICO: 3 specimens, no locality and date, Moritz coll. (ZMHB); 3 specimens, no locality and collector, S.G. Krug coll. (ZMHB). GUADELOUPE: 2 specimens, no locality and date, B. Haas coll. (1 AMNH, 1 FMNH); 7 specimens, Camp Jacob, no date and collector (IRSNB); 1 specimen, no locality and collector, 16.9.1903 (FMNH); 4 specimens, no locality, date and collector (1 FMNH, 2 NMW, 1 ZMHB). Undetermined country: 3 specimens, no locality, date and collector (NMW); 1 specimen, B. Haas coll. (FMNH); 1 specimen, Cuba, Jamaica, 5.X.[19]28, Rambousek coll. (ZMHB). Piestus formicinus, sp. nov. GUYANA: Upper Demerara-Berbice: 80 specimens, Region 8 Iwokrama Forest, 1 km W Kurupukari, Iwokrama Field Stn., 4
400 1900 N, 58
410 400 W, 60m, ex: Acromyrmex hystrix refuse pile, 21.V.2001, R. Brooks and Z. Falin coll. (SEMC); 1 specimen, the same locality and collector, 2.VI.2001 (9 FMNH, 9 DZUP, 9 AMNH, 4 MZSP, 49 SEMC); 1 specimen, Region 8 Iwokrama Forest, Turtle Mt. base camp, 50m, 4
430 500 N, 58
430 500 W, ex: Acromyrmex hystrix refuse pile, 31.V.2001, the same collector (SEMC); 2 specimens, the same locality and collector, 1.VI.2001 (SEMC); 5 specimens, Region 8 Iwokrama Forest, Kabocalli Field Stn., 60m, 4
170 400 N, 58
300 3500 W, 4.VI.2001, the same collector (SEMC); 2 specimens, the same locality and collector, 5.VI.2001 (SEMC). Piestus sulcatus Gravenhorst, 1806 NICARAGUA: Río San Juan: 1 specimen, 8 km SE El Castillo, Refugio Bartola, 10
58.60 N 84
20.40 W, 30 m, rainforest litter, 24–31.V.2002, R. Anderson coll. (SEMC); 2 specimens, the same locality, 100m, flight intercept trap, 28–30.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis (SEMC). COSTA RICA: 4 specimens, Hamburgfarm, 1.II.[19]24, Neverm coll. (FMNH); 1 specimen, the same locality and collector, 27.XI.1936 (FMNH); 14 specimens, Farm Castilla, VI.[19]38, no collector (FMNH); 3 specimens, Finca Castilla, VI.[19]38, no collector (2 FMNH, 1 AMNH); Guanacaste: 1 specimen, Pitilla Biological Station, 10
580 000 N 85
250 000 W, 600 m, berlese leaf litter, 2.V.1995, R. Anderson coll. (SEMC); Alajuela: 1 specimen, Barroeta, 4–8.III.1942, no collector (FMNH); 1 specimen, Alajuela, Peñas Blancas, 800 m, flight intercept, 19.V.1989, J. Ashe, R. Brooks and R. Leschen (SEMC); Heredia: 1 specimen, F. La Selva, 3 km S. Pto. Viejo, 10
260 N 84
010 W, 20.III.1980, H. A. Hespenheide coll. (SEMC); 3 specimens, La Selva Biol. Station nr., Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); 2 specimens, La Selva, 3.2 km SE Puerto Viejo, 100 m, flight intercept trap, 19.III.1992, W. Bell coll. (SEMC); 1 specimen, the same locality and collector, 24.III.1992 (SEMC); 2 specimens, the same locality and collector, 27.III.1992 (SEMC); 2 specimens, 3 km S. Puerto Viejo, OTS-La Selva, 100 m, IX.1992, P. Hanson coll. (SEMC); 17 specimens, the same locality and collector, X.1992, P. Hanson coll. (1 CZUG, 16 SEMC); 1 specimen, La Selva, 80 m, flight intercept trap, 19.IV.1993, J. S. and A. K. Ashe coll. (SEMC); 2 specimens, La Selva Biol. Res. Sta., 3 km S. Puerto Viejo, 10
250 000 N 84
00 000 W, 80 m, flight intercept trap, 2–15.VI.1996, R. Hanley coll. (SEMC); 1 specimen, the same locality and collector, 4–10.VI.1996 (SEMC); 1 specimen, the same locality and collector, 4–15.VI.1996 (SEMC); Cartago: 1 specimen, Turrialba, 15. Revision of Piestus with remarks on related genera Invertebrate Systematics 579 VII.1973, 600 m, V. O. Becker coll. (DZUP); Limón: 1 specimen, Finca Castilla, VI.[19]38, C. Rica coll. (FMNH); Puntarenas: 3 specimens, Osa Penn., Fundación Neotrop., 10 km W. Rincon, 8
450 3000 N 83
250 000 W, 20 m, berlese forest litter, 21.VI.1997, R. Anderson coll. (SEMC); 7 specimens, Rincon, 7 km W, Osa Penn., Est. F.N. Aguas Buenas, 50 m, flight intercept trap, 21–25.VI.1997, S. and J. Peck coll. (SEMC); 1 specimen, Osa Penn., Fundación Neotrop., 10 km W. Rincon, 8
420 3000 N 83
310 3000 W, 20 m, berlese forest litter, 22.VI.1997, R. Anderson coll. (SEMC); 2 specimens, Corcovado National Park, Sirena Stn., upper Rio Claro Trail, 8
280 2900 N 83
350 800 W, 100 m, flight intercept trap, 28.VI–1.VII.2000, Z. H. Falin coll. (SEMC); 7 specimens, Rincon de Osa, 8
41.1410 N 83
31.1170 W, 50 m, FIT, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 6 specimens, Rincon de Osa, 8
41.1410 N 83
31.1170 W, 150 m, FIT, 23–26.VI.2001, S. and J. Peck (SEMC); 5 specimens, Corcovado National Park, Sirena Stn., Corcovado Trail, 8
290 700 N 83
340 3900 W, 150 m, flight intercept trap, 28.VI–1.VII.2000, Z.H. Falin coll. (SEMC); 1 specimen, Corcovado National Park, Sirena Stn., lower Rio Claro Trail, 8
240 4800 N 83
350 2200 W, 40 m, UV light, 30.VI.2000, Z. H. Falin coll. (SEMC); 1 specimen, the same locality and collector, FIT, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 1 specimen, Osa Penninsula, Fnd. Neotrop., 10 km W. Rincon, 08
420 3000 N 83
310 3000 W, 200 m, lowland for litter, 24.VI.2001, R. Anderson coll. (SEMC). PANAMA: 1 specimen, no locality and date, 7 day old tapir dung, H. Dybas coll. (FMNH); 1 specimen, Almirante, IV.[19]43, no collector (FMNH); 4 specimens, Canal Zone, Barro Colorado I., fermented fibrous log at light, 16.I.1959, H. Dybas coll. (FMNH); 6 specimens, Barro Colorado I., 1. II.1959, 7 day old tapir dung, H. Dybas coll. (FMNH); 1 specimen, the same locality and collector, Zetek Trail, 4.IV.1959, fibrous debris in tree trunk, H. S. Dybas coll. (FMNH); 1 specimen, the same locality, 6.II.1976, wet leaves and debris forest stream, A. Newton coll. (FMNH); 3 specimens, San Blas, Nusagandi Reserve, 09
210 N 78
590 W, 350 m, rotting palm flrs., 17.V.1995, J. and A. Ashe coll. (SEMC); 3 specimens, Bocas d. Toro, Fortuna, Chiriquí Grande road, 8
470 N 82
110 W, 800 m, premontane rain forest sifting litter, 14–16.VII.1987, D.M. Oison coll. (FMNH); Veraguas: 1 specimen, Gabina, IX. [19]38, no collector (FMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd., 09
070 N 79
450 W, leaf litter, 29.V.1995, J. Jolly and C. Chaboo coll. (SEMC); 1 specimen, the same locality, 40 m, flight intercept trap, 4–7.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and collector, 7–21.VI.1995 (SEMC); 2 specimens, 15 km N jct. Escobal and Piña Rds., ~30 m, flight intercept trap, 02–11.VI.1996, J. Ashe and R. Brooks coll. (SEMC); Panamá: 1 specimen, Las Cumbres, 24.VIII.1975, no collector (FMNH); 4 specimens, the same locality, 24.X.1975, D. S. Chandler coll. (FMNH); 1 specimen, Old Gamboa Rd., flight intercept trap, 13.VII.1993, D. Windsor coll. (SEMC); 2 specimens, the same locality and collector, 09
040 N 79
400 W, 14–19.XI.1994 (SEMC); 2 specimens, road to Cerro Campana, 8
410 0100 N 79
550 2100 W, 760 m, litter on forest floor and near trail, 1.I.[20]02, L. Herman coll. (AMNH); Darién: 2 specimens, Cana Biological Station, 7
450 1800 N 77
410 600 W, 550 m, flight intercept trap, 03–07.VI.1996, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and collector, 06.VI.1996 (SEMC); 1 specimen, the same locality and collector, 07.VI.1996 (SEMC). GUADELOUPE: 6 specimens, Capesterre, III-25–61, Neufchateau coll. (AMNH). DOMINICA: Saint David: 14 specimens, Salibia, under rotten banana leaves, 28–29.I.1968, B. Malkin coll. (FMNH). MARTINIQUE: 1 specimen, Fort de France, 9–12.VI.1960, P. and C. Vaurie coll. (AMNH). SAINT VINCENT AND THE GRENADINES: Saint Vincent: 2 specimens, Leeward side, no date, H. H. Smith coll. (1 NMW, 1 IRSNB). TRINIDAD AND TOBAGO: Trinidad: 1 specimen, Balandra Bay, IV.1922, no collector (FMNH); 12 specimens, the same locality and date, L. R. Reynold coll. (2 AMNH, 10 FMNH); 5 specimens, the same locality and date, F. Psota coll. (FMNH); 1 specimen, Balandra Bay, 23.V.[19]22, F. Psota coll. (FMNH); 4 specimens, the same locality and date, L.R. Reynold coll. (FMNH); 2 specimens, Balandra Bay, 24.IV.[19]22, the same collector (FMNH); 1 specimen, Arima Valley, 800–1.200ft., 10–22.II.1964, Rozen and Wygodzinsky coll. (AMNH); 1 specimen, Asa Wright N.C., 15.I.1981, G.E. Bohart coll. (SEMC); 2 specimens, Simla Res. Sta., 2–15.VI.1981, H. Clemons coll. (SEMC). COLOMBIA: Chocó: 2 specimens, Quebrada Docordo, between Cucurrupi and Noanama, Rio San Juan, beating dry foliage, 1–5.I.1969, B. Malkin coll. (FMNH). VENEZUELA: Aragua: 8 specimens, 18.9 km S Choroni, 2900ft., litter near stream, 2. III.1992, L. Herman coll. (AMNH); 24 specimens, Rancho Grande-Maracay rd, first stream SE of Rancho Grande, 3600ft., litter near stream, 1.IV.1992, the same collector (AMNH); 16 specimens, 20 km NW Maracay, Rancho Grande, sendero a la Toma de Agua, 3800ft., litter near stream, 22.III[19]92, the same collector (AMNH); 17 specimens, the same locality and collector, La Toma, 1150m, litter nr stream, 15.IV.1994 (AMNH); 1 specimen, Rancho Grande Biol. Stn., 10
210 N 67
410 W, 1140m, flight intecept trap, 25–28.II.1995, R.W. Brooks coll. (SEMC); 4 specimens, the same locality and collector, 1–8.III.1995 (SEMC); 1 specimen, the same locality and collector, 1115 m, along stream, 9.III.1995 (SEMC). GUYANA: Upper Demerara-Berbice: 1 specimen, Region 8, Iwokrama Forest, 1 km W Kurupukari, Iwokrama Field Stn., 4
400 1900 N 58
410 400 W, 60 m, flight intercept trap, 26–29.V.2001, R. Brooks and Z. Falin coll. (SEMC). SURINAMEE: Marowijne: 2 specimens, Nassau Mountain, 4
470 5300 N 54
310 1600 W, 500 m, flight intercept trap, 1–2.VI.1999, Z.H. Falin and B. Dedjin coll. (SEMC); 1 specimen, the same locality, 4
480 5600 N 54
360 2000 W, 480 m, flight intercept trap, 3–4.VI.1999, Z.H. Falin and B. Dedijn coll. (SEMC); 2 specimens, Palumeu, 3
200 5600 N 55
260 1800 W, 160 m, flight intercept trap, 5–9.VII.1999, Z. H. Falin and D. Konoe coll. (SEMC); 1 specimen, the same locality, flight intercept trap, 7–8.VII.1999, Z. H. Falin coll. (SEMC). FRENCH GUIANA: 2 specimens, no locality, date and collector (IRSNB); 1 specimen, Cunani, no date and collector (IRSNB); 3 specimens, Boeuf Mort, ~3
380 N 53
130 W, 160 m, 10.X.1995, L. Herman coll. (AMNH); 1 specimen, Wanaboo, near Nason, Marowjine River, 4
430 3500 N 54
260 3600 W, 40 m, flight intercept trap, 31.V–5.VI.1999, Z. H. Falin, B. Dedjin coll. (SEMC); Saint-Laurent-du-Maroni: 8 specimens, nr. Eaux Claires, 3.5 mi N Saul, 3
38–400 N 53
13–140 W, 155–260 m, leaflitter near stream, 5–13.X.[19] 95, L. H. Herman coll. (AMNH); 2 specimens, the same locality, data e coletor, fig. fruit fall (AMNH); 7 specimens, the same locality, data e coletor, leaf litter and Eschweilera flower fall (AMNH); 3 specimens, Saül, 7 km N, 1 km NW, Les Eaux Claires, along Rue de Belizon trail, 3
390 4600 N 53
130 1900 W, 280 m, flight intercept trap, 4–8.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 4 specimens, Saül, Lecythis zabucajo fallen branch, III–IV.1999, A. Berkov coll. (AMNH); 3 specimens, Saül, sifting, IV.1999, A. Berkov coll. (AMNH); 1 specimen, Saül, 7 km N, Les Eaux Claires, 3
390 4600 N 53
130 1900 W, 220 m, flight intercept trap, 30.V–4.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Saül, Mt. Galbao summit, 3
370 1800 N 53
160 4200 W, 740 m, rotting fruit, 6. VI.1997, J. Ashe and R. Brooks coll. (SEMC); Cayenne: 1 specimen, no locality, date and collector (IRSNB); 2 specimens, Roura, 18.4 km SSE, 4
360 3800 N 52
130 2500 W, 240 m, flight intercept trap, 22–24.V.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Roura, 8.4 km SSE, 4
400 4100 N 52
130 2500 W, 200 m, flight intercept trap, 22–24.V.1997, J. Ashe and R. Brooks coll. (SEMC); 4 specimens, Roura, 18.4 km SSE, 4
360 3800 N 52
130 2500 W, 240 m, flight intercept trap, 25–29.V.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Cayenne, 33.5 km S and 8.4 km NW of Hwy N2 on Hwy D5, 4
480 1800 N 52
280 4100 W, 30 m, flight intercept trap, 26–28.V.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and collector, 29.V–9.VI.1997 (SEMC); 1 specimen, Matoury, 41.5 km SSW on Hwy N2, 4
370 2200 N 52
220 3500 W, 50 m, flight intercept trap, 29.V–9.VI.1997, the same collector (SEMC); 1 specimen, Roura, 18.4 km SSE, 4
360 3800 N 52
130 2500 W, 240 m, flight intercept trap, 29.V–10.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Roura, 13.0 km SSE, 4
380 3800 N, 52
170 5600 W, 240 m, rotting fruit, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, Roura, 27.4 km SSE, 4
440 2000 N 52
130 2500 W, 280 m, flight intercept trap, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC). ECUADOR: Esmeraldas: 1 specimen, Bilsa, 0
200 000 S 79
430 000 W, flight intercept trap, 28.IV–10.V.1996, P. Hibbs coll. (SEMC); 2 specimens, the same locality and collector 10.V–5.VI.1996 (SEMC); 5 specimens, the same locality and collector, 5.VI–7.VII.1996 (SEMC); 1 specimen, the same locality and collector, 7–19.VII.1996 (SEMC); 2 specimens, Bilsa Biological Station, 0
200 2400 N 79
420 3600 W, 500 m, flight intercept trap, 7–19.VII.1996, P. Hibbs coll. (SEMC); Sucumbios: 2 specimens, Sacha Lodge, 0.5
S 76.5
W, 270 m, malaise, 22.II–4.III.1994, Hibbs coll. (SEMC); 1 specimen, the same locality and collector, 14–24.III.1994 (SEMC); 1 580 Invertebrate Systematics E. Caron et al. specimen, the same locality and collector, 23.IV–4.V.1994 (SEMC); 2 specimens, the same locality and collector, 24.V–3.VI.1994 (SEMC); 1 specimen, the same locality and collector, 23.IV–3.VII.1994 (SEMC); 1 specimen, the same locality and collector, 3–13.VII.1994 (SEMC); 3 specimens, the same locality and collector, 13–25.VII.1994 (SEMC); 6 specimens, the same locality and collector, 23.VII–3.VII.1994 (SEMC); 2 specimens, the same locality and collector, 25.VII–3.VIII.1994 (SEMC); 2 specimens, the same locality and collector, 3–16.VIII.1994 (SEMC); 1 specimen, the same locality and collector, 16–29.VIII.1994 (SEMC); 2 specimens, the same locality and collector, 27.VIII–10.IX.1994 (SEMC); 1 specimen, the same locality and collector, 20–30. IX.1994 (SEMC); 20 specimens, the same locality and collector, 21–24.III.1999 (SEMC); 2 specimens, Sacha Lodge, 0
280 1400 S 76
270 3500 W, 270 m, Bignoniaceae, large fruits, 22.III.1999, R. Brooks coll.; Pichincha: 1 specimen, 16 km E Sto.Domingo, Tinalandia, 680 m, malaise rainforest, 4.V–25. VII.1985, S. and J. Peck coll. (FMNH); 2 specimens, 17 km SE Sto. Domingo de Colorado, Tinalandia, 3000’, 16–21.X.[19]88, L. Herman coll. (AMNH); 1 specimen, W of Alluriquim, Tinalandia, 2600–2800ft., mercury vapor light 19–20.V.1993, L. Herman coll. (AMNH); 1 specimen, 45 km NNW Quito, Macquipucuna Station, 1600–1650 m, flight intercept trap, 3–18.IV.1996, P. Hibbs coll. (SEMC); 3 specimens, Tinalandia, Santo Domingo, 16 km E, 0
160 5300 S 79
30 3900 W, 750 m, flight intercept trap, 26–27.III.1999, R. Brooks and D. Brzoska coll. (SEMC); Napo: 1 specimen, Jatun Sacha Biol. Station, 21 km E. Puerto Napo, 400 m, 8–11.VII.1994, F. Génier coll. (SEMC); 3 specimens, the same locality and collector, 9.VII.1994 (SEMC); 5 specimens, the same locality and collector, 21.VII.1994 (SEMC); 2 specimens, the same locality and collector, 13.VII.1994 (SEMC); 3 specimens, the same locality, 18.VII.1994, Levy and Génier coll. (SEMC); 1 specimen, km 7.3 Sarayacu Loreto Rd., 1200 m, cloud forest, leces trap, 20.VII.1994, F. Génier coll. (SEMC); 1 specimen, Onkone Gare Camp, 00
390 1000 S 76
260 0000 W, 220 m, under leaf litter, 4–12.X.1995, T.L. Erwin, G.E. Ball and D. Shpeley coll. (SEMC); 2 specimens, Sierra Azul, Hacienda Aragon, 0
400 000 S 77
550 000 W, 2300 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); 1 specimen, Yuturi Lodge, Rio Napo, 0
320 5400 S 76
20 1800 W, 270 m, flight intercept trap, 20–21.III.1999, R. Brooks and D. Brzoska coll.; Cotopaxi: 1 specimen, Guasaganda, 500 m, 1. IV.1988, M. Huybensz coll. (FMNH); Pastaza: 3 specimens, Puyo, nr. Santa Clara, 950 m, 18–21.VII.2008, W. Rossi and I. Tapia coll. (FMNH). PERU: 1 specimen, Tambopata Prov., 15 km NE Pto. Maldonado, 200 m, flight intercept, 3.VI.1989, J. Ashe and R. Leschen coll. (SEMC); 1 specimen, the same locality and collector, at light, 10.VI.1989 (SEMC); 1 specimen, the same locality and collector, flight intercept, 22.VI.1989 (SEMC); 2 specimens, the same locality and collector, 18.VII.1989 (SEMC); Loreto: 1 specimen, Campamento San Jacinto, 2
18.750 S 75
51.770 W, 175–215 m, berlese, 2.VII.1993, R. Leschen coll. (SEMC); 3 specimens, the same locality and collector, flight intercept trap, 3.VII.1993 (SEMC); 1 specimen, the same locality and collector, 7. VII.1993 (SEMC); 4 specimens, 1.5 km N Teniente Lopez, 2
35.660 S 76
06.920 W, 210–240 m, palmfruit berlese, 17.VII.1993, R. Leschen coll. (SEMC); 1 specimen, the same locality and collector, 20.VII.1993 (SEMC); 4 specimens, the same locality and collector, 21.VII.1993 (SEMC); 1 specimen, the same locality and collector, 22.VII.1993 (SEMC); 1 specimen, the same locality and collector, 24.VII.1993 (SEMC); 1 specimen, the same locality and collector, 24.VII.1993 (SEMC); 3 specimens, the same locality and collector, 27.VII.1993 (SEMC); San Martín: 1 specimen, Prov. Huallaga, Tocache, 500 m, 10. XI.1900, G. A. Baer coll. (IRSNB); Huancavelica: 1 specimen, Panguana, Rio Pachitea, Rio Yuyapichia, 9
370 S 74
560 W, 260 m, 1–14.IX.1986, Listabarch coll. (NMW); Cusco: 1 specimen, Consuelo, Manu rd., km 165, 3.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 1 specimen, the localidade and collector, 6.X.1982 (FMNH); 1 specimen, the same locality and collector, 7–8.X.1982 (FMNH); 2 specimens, the same locality and collector, 8.X.1982 (FMNH); 1 specimen, the same locality and collector, 10.X.1982 (FMNH); Madre de Dios: 1 specimen, Tambopata, 25.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 1 specimen, the same locality and collector, 28.X.1982 (FMNH); 1 specimen, Tambopata Prov., 15 km NE Puerto, Maldonado Reserva Cuzco Amazónico, 12
330 S 69
030 W, 200 m, 8.VI.1989, R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, pitfall trap, 22. VI.1989, J. S. Ashe and R. A. Leschen coll. (SEMC); 1 specimen, the same locality and collector, 25.VI.1989 (SEMC); 1 specimen, the same locality and collector, 30.VI.1989 (SEMC); 1 specimen, the same locality and collector, 2.VII.1989 (MUSM); 1 specimen, Manu Prov., Parque Nac. Manu, Zona Res., Rio Manu, Cocha Juarez, trail nr. Manu, Lodge, flight intercept trap, 18–24.IX.1991, A. Hartman coll. (FMNH); 5 specimens, Cocha Cashu Bio. Stn., Manu National Park, 350 m, 11
530 4500 S 71
240 2400 W, flight intercept trap, 17–19.X.2000, R. Brooks coll. (SEMC); 1 specimen, Cocha Salvador, Reserve Zone Manu National Park, 310 m, 12
00 1300 S 71
310 3600 W, 20.X.2000, R. Brooks coll. (SEMC); 3 specimens, Pantiacolla Lodge, 8 km NW El Mirador Trail, Alto Madre de Dios River, 800 m, flight intercept trap, 12
380 3000 S 71
160 4100 W, 23–26.X.2000, R. Brooks coll. (SEMC). BOLIVIA: 3 specimens, Yuracaris (IRSNB); Cochabamba: 8 specimens, Cochabamba, 109 km E, Yungas, Cochabamba – Villa Tunari Rd., 1480 m, 17
80 5000 S 65
420 4900 W, flight intercept trap, 1–6.II.1999, F. Génier coll. (SEMC); 2 specimens, the same locality, 8–12.II.1999, R. Hanley coll. (SEMC); 5 specimens, the same locality, 1400 m, 17
80 5200 S 65
420 5400 W, 6.II–8.VI.1999, F. Génier coll. (SEMC); 2 specimens, the same locality and collector, 1400 m, 17
80 5200 S 65
420 5400 W, 8–12.II.1999 (SEMC); 2 specimens, Cochabamba, 117 km E, Yungas Lagunitas, 1000 m, 17
60 2200 S 65
400 5700 W, flight intercept trap, 1–6.II.1999, F. Génier coll. (SEMC); 4 specimens, Cochabamba, 124 km E, Yungas, Cochabamba – Villa Tunari Rd., 700 m, 17
30 4500 S 65
380 3800 W, carrion trap, 1–6.II.1999, F. Génier coll. (SEMC); 1 specimen, Cochabamba, 67.5 km NE, Est. Biol. Valle del Sajita, Univ. de San Simon, 300 m, 17
60 3300 S 64
470 5200 W, flight intercept trap, 9–13.II.1999, F. Génier coll. (SEMC); Santa Cruz: 1 specimen, 3.7 km SSE Buena Vista, Hotel Flora Y Fauna, 17
29.950 S 63
33.15W, 400–440 m, primary forest, FIT, 4–9.XI.2002, R. Leschen coll. (SEMC); 1 specimen, the same locality and collector, Chiquitano forest, at light, 6.XI.2002, S. Lingafelter coll. (SEMC). BRAZIL: 1 specimen, Chapada, no date and collector (FMNH); 1 specimen, Pebas, no date and collector (IRSNB); 2 specimens, no locality, date and collector (1 NMW, 2 MNRJ); 1 specimen, Vassouras,18.IX.1933, Eldmann coll. (DZUP); Amazonas: 1 specimen, Ega, no date and collector (IRSNB); 1 specimen, Reserva Ducke, 26 km. NE Manaus, 28.II.1977, B. C. Ratcliffe coll. (INPA); 4 specimens, 35 km NE Manaus, Res. Flor. Ducke, 25. VII–08.VIII.[19]95, Arndt and Gröger coll. (ZMHB); Pará: 1 specimen, Aldeia Maracacume, Maranhao 80 km E. of Caninde, 22–23.V.1963, B. Malkin coll. (FMNH); 7 specimens, Belém, Guamá, 22.III.1974, I. S. Gorayeb coll. (INPA); 16 specimens, Belém, Rio Guamá, 22.III.1974, R. T. Schuh coll. (AMNH); Bahia: 1 specimen, San Antonio da Barra, no date and collector (IRSNB); 1 specimen, no locality and date, Bohm. coll. (FMNH); 6 specimens, no locality and date, Fry coll. (FMNH); 1 specimen, no locality, date and collector (FMNH); Goiás: 1 specimen, Jatahy, no date and collector (IRSNB); Distrito Federal: 1 specimen, Brasília, XI-1987, V. de Mello coll. (AMBC); Rio de Janeiro: 8 specimens, no locality and date, Fry coll. (1 ZMHB, 7 FMNH); 1 specimen, no locality and date, Boehm. coll. (FMNH); 4 specimens, Tijuca, XII.1884, E. Gounelle coll. (2 ZMHB, 2 IRSNB); 15 specimens, Deodoro, 3.IV.[1]937, W. Zikán coll. (MNRJ); 1 specimen, Tijuca, 5.III.1959, H. Schubart coll. (MNRJ); 1 specimen, the same locality and collector, 18. V.1959 (MNRJ); 2 specimens, Trapicheiro, 13.XII.1959, the same collector (MNRJ); 1 specimen, Estr. das Canoas, 3.XI.1962, H. Schubart coll. (INPA); 1 specimen, Rio de Janeiro, XII.1967, M. Alvarenga coll. (AMNH); 1 specimen, the same locality and collector, X.1969 (AMNH); 3 specimens, Represa Rio Grande, III.1972, the same collector (AMNH); São Paulo: 2 specimens, São Paulo, Ypiranga, Ihering coll. (FMNH); Paraná: 2 specimens, nr. Porto de Cima, 31.III–1.IV.2006, W. Rossi coll. (DZUP); Santa Catarina: 1 specimen, no locality, date and collector (NMW); 1 specimen, no locality and date Lüderwaldt coll. (FMNH); 3 specimens, no locality and date, Klimsch coll. (2 FMNH, 1 NMW); 2 specimens, Blumenau, no date, Hetschko coll. (NMW); 1 specimen, no locality and date, Fry coll. (FMNH); 1 specimen, Blumenau, no date and collector (DZUP); 1 specimen, no locality, 1930, W. Ehrhardt coll. (ZMHB); 1 specimen, Corupa, XI.1848, H. Humboldt coll. (AMNH); 1 specimen, Corupa, XII.1948, H. Humboldt coll. (AMNH). Undetermined country: 10 specimens, no locality, date and collector (1 NMW, 3 MNRJ, 2 ZMHB, 4 MZSP). Revision of Piestus with remarks on related genera Invertebrate Systematics 581 Piestus gounellei Fauvel, 1902 BRAZIL: Espírito Santo: 2 specimens, Pedra Azul, 1500 m, Domingos Martins, I.2000, A. Bello coll. (AMBC); Rio de Janeiro: 1 specimen, no locality, XII.1899, ilegible coll. (FMNH); 1 specimen, no locality, date and collector (FMNH); 1 specimen, Nova Friburgo, 2–3.IV.1903, E. Gounelle coll. (IRSNB); 1 specimen, Guanabara, Represa Rio Grande, III.1972, M. Alvarenga coll. (AMNH); 1 specimen, Nova Friburgo, Macaé de Cima, 1500 m, 1–15.I.2006, P. Grossi, coll. (DZUP); São Paulo: 1 specimen, no locality and date, B. Pohl coll. (AMNH); 1 specimen, no locality and date, B. Mráz coll. (FMNH); 1 specimen, Casa Grande, IX.1938, J. Guerin coll. (FMNH); 4 specimens, Serra da Bocaina, S. Jose Barreiro, 1650 m, XI.1968, M. Alvarenga coll. (AMNH); Paraná: 1 specimen, Marumbi, Morretes, 500 m, 12.XI.1966, Laroca and O. Mielke coll. (DZUP); 1 specimen, Mananciais da Serra, Piraquara, III.2005, A. J. C. Aguiar coll. (DZUP); 1 specimen, the same locality, 1000 m, XI.2005, P. Grossi, coll. (DZUP); 5 specimens, the same locality and collector, 30.XI.2005 (1 ZMHB, 4 DZUP); 1 specimen, the same locality and collector, 07.XII.2005 (DZUP); 1 specimen, the same locality and collector, XII.2005 (DZUP); 3 specimens, the same locality and collector, 25
180 4000 S 48
580 0400 W, 1000 m, 09.XII.2005 (1 ZMHB, 2 DZUP); 7 specimens, the same locality, 17.X.2007, E. Caron and P. Grossi coll. (DZUP); Santa Catarina: 1 specimen, no locality, date and collector (FMNH); 2 specimens, Blumenau, no date, Hetschko coll. (NMW); 1 specimen, the same locality, no date and collector (FMNH); 1 specimen, Nova Teutonia, II.1933, B. Pohl coll. (MZSP); 1 specimen, the same locality, 27
110 B 52
230 L, VIII.1935, F. Plaumann coll. (MNRJ); 1 specimen, the same locality and collector, 24.IX.1938 (FMNH); 1 specimen, the same locality and collector, XII.1953 (FMNH). PARAGUAY: Caazapá: 1 specimen, Hermosa. Prop. Sosa family, San Rafael Reserve, 26
190 1500 S 55
440 5500 W, 90 m, flight intercept trap, 3–6.XII.2000, Z. H. Falin coll. (SEMC); Itapúa: 1 specimen, Yatai, prop. Hostettler family, San Rafael Reserve, 100 m, 26
380 1700 S 55
390 5000 W, flight intercept trap, 21–25.XI.2000, Z. H. Falin coll. (SEMC). ARGENTINA: Misiones: 1 specimen, P.N. Iguzu, Salto Macuco, 26.XII.1990, S. and J. Peck coll. (FMNH); 1 specimen, 15 km SE Puerto Iguazu, mature forest, 27.XII.1990–6.I.1991, S. and J. Peck (FMNH). Undetermined country: 2 specimens, no locality, date and collector (NMW). Piestus mexicanus Laporte, 1835 MEXICO: 31 specimens, no locality, date and collector (4 ZMHB, 2 IRSNB, 13 NMW, 11 FMNH, 1 AMNH); 1 specimen, no locality and collector, 1871 (NMW); 7 specimens, Jalapa, no date, Flohr coll. (ZMHB); 1 specimen, no locality an date, L. W. Schaufuss coll. (ZMHB); 1 specimen, no locality and date, Hoege coll. (NMW); 1 specimen, no locality and collector, 1890 (NMW); 3 specimens, no locality and date, Truqui col. (FMNH); 3 specimens, Jalapa, no date, Hoge (FMNH); 4 specimens, no locality and date, Reitter coll. (FMNH); Tamaulipas: 4 specimens, Mpo. Gomez Farias, Atlas Cimas, 1000 m, leaf litter, 16.III.1989, R. W. Jones coll. (1 DZUP, 3 FMNH); Durango: 2 specimens, no locality, date and collector (1 ZMHB, 1 NMW); San Luís Potosí: 2 specimens, El Salto de Agua, 20.VI.1975, L. E. Watrous coll. (FMNH); Hidalgo: 2 specimens, 55 km NE Jacala, 1190 m, 4.VI.1987, R. Anderson coll. (SEMC); Veracruz: 1 specimen, Jalapa, no date, F. Schneider coll. (ZMHB); 6 specimens, Sierra de Zongolica, no date and collector (5 ZMHB, 1 SEMC); 2 specimens, Cordoba, no date, A. Fenyes coll. (FMNH); 2 specimens, Fortin de las Flores, 28.VI.1975, Q. W. Wheeler coll. (FMNH); 1 specimen, Balzapote, berlese, rain forest leaf litter, 7.VII.1976, A. Newton coll. (FMNH); 6 specimens, 4.4 mi N Huatusco, cloud forest, 4200ft., sifting litter at edge of stream, 24. IV.1977, J. S. Ashe coll. (FMNH); 1 specimen, the same locality and collector, 25.IV.1977 (FMNH); 1 specimen, 33 km NE Catemaco, Los Tuxtlas Biol. Sta., 160 m, ridge rainforest, FIT, 1.VII.1983–1.VIII.1983, S. and J. Peck coll. (FMNH); 1 specimen, the same locality, 20–27.VII.1984, H. and A Howden coll. (FMNH); 1 specimen, 1.7 km W Huatusco, Hwy. 125, 1250 m, mushrooms, rotting vegetation, 16.VII.1990, I. Yarom coll. (SEMC); 8 specimens, 18.8 km S Huatusco, Hwy. 125 and 13.3 km E on Ixhuatlán Rd., 1130 m, sifting trash, 17.VII.1990, J. S. Ashe, K. J. Ahn and R. Leschen coll. (SEMC); 1 specimen, the same locality and collector, rotten banana trunk, 19.VII.1990 (SEMC); Oaxaca: 6 specimens, 7.3 mi N Chacalapa, 15
490 N 96
280 W, Hwy 175, tropical deciduous, 410 m, sifting litter along stream, 29.IV.1977, J. S. Ashe coll. (FMNH); Chiapas: 1 specimen, 2.5 mi SW El Bosque, 4000’, near nest entrance Atta cephalotes, 25.VIII.1973, A. Newton coll. (FMNH); 1 specimen, 8 mi N Pueblo Nuevo Solistahuacán, 6000’, berlese cloud forest/leaf litter, 26–27.VIII.1973, A. Newton coll. (FMNH); 2 specimens, 13.5 mi s. Mapastepec, under bark log. Lowland, rainforest, 120 m, 30.XII.1975, H. Frania coll. (AMNH); 12 specimens, 4 km N of Escuintla, wet rooting cut vege. and leaves along stream lowland rainforest, 120 m, 4.I.1977, H. Frania coll. (AMNH); 25 specimens, 9.7 mi S Solusuchiapa, Hwy 195, montane tropical, 530 m, sifting litter along stream, 5.V.1977, J. S. Ashe coll. (FMNH); 1 specimen, 4.7 km N Finca Prucia, 24.6 km S Jaltinango de la pas., 1050 m, oak-pine-montane, trop. transition, 3.VII.1979, J. S. Ashe coll. (FMNH); 3 specimens, Bonampak Rd., 100 km SE Palenque, berlese rainforest, 8.VII.1983, S. and J. Peck coll. (FMNH); 16 specimens, the same locality and collector, 230 m, 24.VII.1983 (FMNH); 11 specimens, Parque Laguna Belgica, 19.3 km N Ocozocoautla, 970 m, 8.VI.1991, J. Ashe coll. (SEMC); 36 specimens, 5.9 km E Bochil, 1300 m, riparian mesophytic forest litter, 15.IX.1992, R. S. Anderson coll. (SEMC); 4 specimens, 10 km W El Bosque 1475 m, 15.IX.1992, R. S. Anderson coll. (SEMC). BELIZE: Cayo: 1 specimen, District Chiriqui N.P., 16
290 3500 N 89
020 4900 W, Doyles Delight, nr. Campground, 1100 m, 19–28. VIII.2007, P. W. Kovarik coll. (FMNH). GUATEMALA: 8 specimens, no locality, date and collector (7 ZMHB, 1 IRSNB, 2 FMNH); Zacapa: 1 specimen, 3.5 km SE La Union, 1500 m, 23.VI.1993, J. Ashe and R. Brooks (SEMC); 1 specimen, the same locality and collector, 23–25.VI.1993 (SEMC); 1 specimen, the same locality and collector, 25–27.VI.1993 (SEMC); Escuintla: 1 specimen, Finca San Victor, 600ft., decaying bananas, 10.IX.[19]48, R. D. Mitchell coll. (FMNH). NICARAGUA: Granada: 1 specimen, Res. Nat. Volcan Mombacho, 11
50.050 N 85
58.830 W, 1210 m, flight intercept trap, 1–5. VI.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 3 specimens, San Juan, no date, E. Schimidt coll. (AMNH); 8 specimens, Vara Blanca, 2000 m, H. Schmidt coll. (4 AMNH, 4 FMNH); 3 specimens, no locality, date and collector (FMNH); 1 specimen, Volcan Irazu, no date and collector (IRSNB); 1 specimen, Puente de las Mulas, III.1916, no collector (FMNH); 1 specimen, Rio Virilla, 26.XII.[19]31, H. Schmidt coll. (SEMC); 2 specimens, Río Virillo, II.[19]35, no collector (FMNH); 1 specimen, Vara Blanca, II.1936, no collector (FMNH); 1 specimen, no locality, 12–16.II.[19]40, no collector (FMNH); 3 specimens, El Poró, 800 m, 18.II.1941, no collector (FMNH); 2 specimens, Sardinal, 23.II.1941, no collector (FMNH); Guanacaste: 6 specimens, Lomas Bardubal Biological Reserve, 10
300 2200 N 85
220 1500 W, 17 m, dead log, no date and collector (SEMC); 2 specimens, Maritza Biol. Stn., 550 m, flight intcpt. Trap, 22.V.1993, J. and A. Ashe coll. (SEMC); 13 specimens, 14 km WNW Bagaces, Lomas Barbudal, Berlese litter, 29.VI.1994, S. O’Keefe coll. (FMNH); Alajuela: 1 specimen, Sn. Mateo, 500 m, 28.II.[19]39, no collector (FMNH); 1 specimen, Hiquito, San Mateo, 5.IX.[19]40, Sales coll. (FMNH); 3 specimens, Escobal,4.IV.[19]43, no collector (FMNH); 1 specimen, 155 km N Jet.Trs. 126(9) and 120, 1.2 km E on Rd to Virgen de Socorro, Río Sarapiquí, 2300ft., under bark of log, 28.III.1991, L. Herman coll. (AMNH); Heredia: 69 specimens, 9.7 km N Heredia, Rt. 9, 4200ft., litter near stream, 9.II.1985, L. Herman coll. (AMNH); 2 specimens, 22.8 km N Heredia, Rt.9, 5700ft., litter from stream, 9.II.1985, L. Herman coll. (AMNH); 2 specimens, Sto. Domingo del Heredia, INBio Cafetal, 1100 m, flight intercept trap, 25–28.VI.1997, S. and J. Peck coll. (SEMC); San José: 3 specimens, La Uruca, 1100 m, no date and collector (IRSNB); 1 specimen, La Caja, 8 Kil., 1934, Schmidt coll. (FMNH); 9 specimens, no locality, date and collector (AMNH); 32 specimens, no locality, date and E. Schmidt coll. (AMNH); 3 specimens, La Caja, no date, Nevermann coll. (2 NMW, 1 DZUP); 5 specimens, the same locality, no date, Reimoser coll. (NMW); 2 specimens, the same locality, no date, Schmidt coll. (FMNH); 1 specimen, Finos La Caja, 1930, 582 Invertebrate Systematics E. Caron et al. H. Schmidt coll. (FMNH); 12 specimens, La Caja: 8 kil., 1931, Schmidt coll. (FMNH); 4 specimens, no locality and collector, 1935 (FMNH); 1 specimen, III.[19]37, no collector (FMNH); 5 specimens, Caja, VII-[19]38, no collector (FMNH); 3 specimens, 8.5 km S Orotina, 300 ft., 16.II.1985, L. Herman coll. (AMNH); 1 specimen, Escazu, 25–29.VI.1988, F. D. Parker coll. (SEMC); 1 specimen, the same locality and collector, 28–31.V.1989 (SEMC); Cartago: 2 specimens, Irazu, no date and collector (NMW); Puntarenas: 9 specimens, Las Cruces Botanical Garden near San Vito, 3500 ft., 27–28.II.1985, L. Herman coll. (AMNH); 25 specimens, 65 km, W Rincon, nr. Fundacion Neotropica, litter nr. stream, 25.III.1991, the same collector (AMNH); 1 specimen, Corcovado National Park, Sirena Stn., upper Rio Claro Trail, 8
280 2900 N 83
350 800 W, 100 m, flight intercept trap, 28.VI–1.VII.2000, Z. H. Falin coll. (SEMC). PANAMA: 1 specimen, Canal Zone, Barro Colorado Island, Zetek Trail, fibrous debris in tree trunk, 4.IV.1959, H. S. Dybas coll. (FMNH); 1 specimen, Canal Zone, Albrook Forest Site, 100ft., on ground, 20.I.1968, R. Hutton coll. (AMNH); 1 specimen, the same locality, ground black light trap, 30–31.I.1968, R. S. Hutton coll. (AMNH); Chiriquí: 3 specimens, no locality and collector, 1930 (FMNH); 1 specimen, 9.7 road miles S. of El Volcan, alt. 2750ft., 10.III.1959, G. A. Solem coll. (FMNH); 1 specimen, Cordillera Abajo, 08
400 5600 N 82
360 1000 W, 920 m, litter near stream, 21.V.[20]01, L. Herman and W. Opitz coll. (AMNH); 1 specimen, near Dolega, 380 m, 8
360 1000 N 82
250 2000 W, litter near stream, 23.XII.[20]01, L. Herman coll. (AMNH); 1 specimen, 1.7 km E. Rio Sereno, 8
490 2300 N 82
500 2900 W, 890 m, litter nr stream, 26.XII.[20]01, L. Herman coll. (AMNH); Panamá: 1 specimen, Altos de Majé, berlese litter in young forest, 6–15.X.1975, D. S. Chandler coll. (FMNH); 12 specimens, Las Cumbres, sift forest litter, 24.X.1975, the same collector (FMNH); 3 specimens, Cerro Azul, 2000’, wet debris small forest stream, 21.II.1976, A. Newton coll. (1 DZUP, 2 FMNH); 25 specimens, Parque Nac. Soberamia, Old Gamboa Rd., 4 km E Gamboa, leaf-fruitfall, 11.VI.1993, Jameson coll. (SEMC); 19 specimens, road to Cerro Campana, 8
420 3400 N 79
530 0600 W, 280 m, litter nr stream, 2.I[20]02, L. Herman coll. (AMNH). COLOMBIA: Boyacá: 1 specimen, no locality, date and collector (IRSNB). Undetermined country: 5 specimens, no locality, date and collector (2 ZMHB, 3 NMW). Piestus foveolatus, sp. nov. COSTA RICA: 10 specimens, Farm Castilla, VI.[19]38, no collector (FMNH); 4 specimens, Finca Castilla, VI.[19]38, no collector (FMNH); 1 specimen, ilegible locality,1.V.1939, no collector (FMNH); 1 specimen, ilegible locality, 3.VIII.[19]39, no collector (FMNH); 2 specimens, Tapauté?, 9.IX.[19]39, no collector (FMNH); 1 specimen, Tapauté?, 15.III.[19]40, no collector (FMNH); 2 specimens, Tapauté?, 22–28.I.1941, no collector (FMNH); 1 specimen, Tapauté?, 27.I.1941, no collector (FMNH); 1 specimen, Peralta, 21.II.1943, no collector (FMNH); 1 specimen, Capellades, 1650m, 19–21.III.[19]40, no collector (FMNH); 3 specimens, Cervantes, 8.IV.1940, no collector (FMNH); Alajuela: 1 specimen, 96 km. N Jct Rts. 9 and 120, Rd. to Puerto Viejo, 4600’, litter near stream, 17.III.1991, L. Herman coll. (AMNH); 8 specimens, 7.7 km, N Jet. Rt./126(9) and 120, Rd. to Puerto Viejo, 4600’, litter near stream, 17.III.1991, L. Herman coll. (AMNH); 1 specimen, 14 km S. Volcan Arenal, 10
200 N, 84
430 W, 1000m, sifted leaf litter, 29.IV.1968, J. Longino coll. (SEMC); 1 specimen, Eladio’s River trail, 800m, flight intercept, 19.V.1989, J. Ashe, R. Leschen and R. Brooks coll. (SEMC); 1 specimen, Peñas Blancas, 970m, flight intercept trap, 20.V.1989, the same collector (SEMC); 2 specimens, the same locality and collector, 1190m, flight intercept trap, 20. V.1989, J. Ashe, R. Leschen and R. Brooks coll. (SEMC); Heredia: 4 specimens, La Selva, 3.2 km SE, Puerto Viejo, 100m, flight intercept trap, 14.II.1992, W. Bell coll. (SEMC); 1 specimen, the same locality and collector, 21.II.1992 (SEMC); 1 specimen, the same locality and collector, 17.III.1992 (SEMC); 2 specimens, the same locality and collector, 24.III.1992 (SEMC); 1 specimen, the same locality and collector, 27.III.1992 (SEMC); 2 specimens, La Selva Biol. Res. Sta. 3 km. S. Puerto Viejo, 80m, 10
250 000 N, 84
00 000 W, flight intercept trap, 2–15.VI.1996, R. Hanley coll. (SEMC); 14 specimens, Finca Zurqui, Tajo, 10
020 300 N, 84
010 2600 W, 1575m, wet cloud forest litter, 27.VI.2000, R. Anderson coll. (SEMC); San José: 29 specimens, Cerros de Escazu, 2 km S. San Antonio, 1650m, 9
530 3000 N, 84
90 000 W, berlese forest litter, 13.VI.1997, R. Anderson coll. (SEMC); 13 specimens, Cerro Chompipe 2 km. N. Monte de la Cruz, 2000m, 9
530 3000 N, 84
90 000 W, berlese forest litter, 13.VI.1997, R. Anderson coll. (SEMC); Puntarenas: 3 specimens, 35 km NE San Vito nr. Las Alturas, Rio Bella Vista, Rd to Gravel Pit, 4300’elev., dry leaf litter near river, 22.III.1991, L. Herman coll. (AMNH); 1 specimen, the same locality and collector, wet leaf litter, 23.III.1991 (AMNH); 8 specimens, the same locality, date and collector, moist leaf litter (AMNH); 6 specimens, the same locality, date and collector, dry leaf litter (AMNH); 5 specimens, San Vito, Estac. Biol. Las Alturas, 2 km NE, 8
560 5600 N, 82
500 100 W, 1520m, berlese leaf litter, 20.VI.1998, R. Anderson coll. (SEMC); 1 specimen Las Cruces Biol. Sta., San Vito 5 km SW, 1425m, 8
460 5900 N, 82
590 1800 W, berlese leaf litter, 22.VI.1998, the same collector (SEMC); 15 specimens, Las Alturas Biol. Sta., 1660m, 08
56.170 N, 82
50.010 W, flight intercept trap, 31.V–3.VI.2004, J.S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC); 4 specimens, Altamira Biol. Sta., 1510–1600m, 09
01.760 N, 83
00.490 W, flight intercept trap, 4–7.VI.2004, the same collector (SEMC). PANAMA: Chiriquí: 1 specimen, Finca Palo Santo, nr. Nueva Califórnia, alt. 4200 ft., at light, 9–10.III.1959, H.S. Dybas coll. (FMNH); 1 specimen, nr. Nueva California, W. of Finca Palo Santo, 5000 ft., 10.III.1959, H. Dybas coll. (FMNH); 3 specimens, NW of Volcán, 1410m, 08
490 1600 N, 82
400 2600 W, litter near stream, 17.V.[20]01, L. Herman and W. Opitz coll. (AMNH); 2 specimens, La Fortuna, Cont. Divide Trail, 08
460 N, 82
120 W, 1150m, flight intercept trap, 23.V–9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality and collector, slash, 9.VI.1995; 1 specimen, the same locality and date, berlese forest litter, R. Anderson coll. (SEMC); 1 specimen, 20 km N Gualaca, Finca La Suiza, 1350m, 8
390 000 N, 82
120 000 W, flight intercept trap, 24.V–9.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 3 specimens, the same locality, 1450m, oak forest litter, 12.VI.1995, R. Anderson coll. (SEMC); 1 specimen, 27.7 km W. Volcan Hartmann’s Finca, 08
450 N, 82
480 W, 1450m, flight intercept trap, 14–17.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 1 specimen, the same locality, 1500m, 8
510 4200 N, 82
440 3600 W, oak forest litter, 16.VI.1995, R. Anderson coll. (SEMC); 2 specimens, the same locality, 1600m, 8
510 4200 N, 82
440 4800 W, treefall litter, 17.VI.1996, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: Valle Del Cauca: 1 specimen, PNN Farallones de Cali Anchicaya, 3
260 N, 76
480 W, 730m, Winkler, 21–23.I.2001, S. Sarria coll. (SEMC). Piestus sulcipennis Scheerpeltz, 1952 BRAZIL: 1 specimen, no locality and date, Bang coll. (FMNH). Piestus costatus Sharp, 1887 COSTA RICA: 1 specimen, Coronado, 1400–1500 m, 15.VIII.[19]31, Nevermann coll. (FMNH); 1 specimen, no locality and collector, 5.V.[19]40 (FMNH); 1 specimen, Puntarenas-Guanacaste, border Monte Verde, Cerro Amigos, 1760 m, flight intercept trap, 12.V.1989, J. Ashe, R. Brooks and R. Leschen coll. (SEMC); San José: 7 specimens, km 113 Pan-Am. Hwy, 23 km. N. San Isidro, 2000 m, 9
290 000 N 83
420 2000 W, berlese forest litter, 20. VI.1997, R. Anderson coll. (SEMC); 4 specimens, km 117 Pan-Am. Hwy, 19 km. N. San Isidro, 1800 m, 9
280 000 N 83
420 2000 W, berlese forest litter, 20. VI.1997, R. Anderson coll. (SEMC); 1 specimen, the same locality and collector, 25.VI.1997 (SEMC); 1 specimen, PanAmerican Hwy Km 80.5, 7 km SSW, Cabinas de Quetzal, 9
330 5300 N 83
480 500 W, 2150 m, fogging fungus covered log, 21.VII.2000, J.Ashe, R. Brooks and Z. Falin coll. (SEMC); Cartago: 2 Revision of Piestus with remarks on related genera Invertebrate Systematics 583 specimens, Tapantí, 22–28.I.1944, no collector (FMNH); 3 specimens, P.N. Tapantil, 1150 m, 9
450 4100 N 83
470 500 E, flight intercept trap, 17–20.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); Puntarenas: 7 specimens, OTS Sta. Finca Las Cruces, 4000ft., San Vito, 82
580 W 8
460 N, Berlese 1500cc. leaf litter in stream bed, away from flowing water steep banks, Virgin forest cover, 18.III.1973, J. Wagner and J. Kethley coll. (FMNH); 9 specimens, the same locality and collector, 20.III.1973 (FMNH); 3 specimens, Prov. 35 km. NE San Vito nr. Las Alturas, Rio Bella Vista, Rd. To Gravel Pit, 4300’elev., dry leaf litter near river, 22.III.1991, L. Herman coll. (1 DZUP, 2 AMNH); 15 specimens, the same locality and collector, moist leaf litter, 23.III.1991 (AMNH); 3 specimens, the same locality and collector wet leaf litter, III-23–1991 (AMNH); 19 specimens, Las Cruces Botanical Garden near San Vito, 3500ft., 27–28. II.1985, L. Herman coll. (AMNH); 1 specimen, the same locality and collector, 27.II.1985 (AMNH); 1 specimen, Monte Verde, 1400 m, flight intercept trap, 21.V.1989, J. Ashe, R. Brooks and R. Leschen (SEMC); 7 specimens, Altamira Biol. Sta., 1510–1600 m, 09
01.760 N 83
00.490 W, inside hepialid webs on trees, 6.VI.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC). PANAMA: Bocas del Toro: 1 specimen, 25 km NNE San Felix, 81
500 8
340 , 1500 m, Berlese floor litter and root mat. Quebrada Alicia cloud forest, 11.VI.1980, J. Wagner coll. (FMNH); Chiriquí: 3 specimens, nr. Nueva California, Finca Palo Santo, 4900’, 6.III.1959, H. S. Dybas coll. (FMNH); 2 specimens, Finca Lerida, nr. Boquete, 5650ft., under bark of log on ground, 12.III.[19]59, H. Dybas coll. (FMNH); 5 specimens, 4 km NW Volcán, 1450 m, 8
490 1900 N 82
400 3100 W, litter near stream, 24.XII.[20]01, L. Herman coll. (1 DZUP, 4 AMNH); 3 specimens, Tizingal Rd. NW of Volcán, 8
480 3700 N 82
390 5300 W, 1350 m, litter nr stream, 25.XII.[20]01 L. Herman coll. (AMNH). Piestus chiriquensis Sharp, 1887 NICARAGUA: Río San Juan: 1 specimen, 60 km SE San Carlos, Refúgio Bartola, 100 m, 10
58,400 N 84
20.300 W, 25.V.2002, R. Brooks, Z. Falin and S. Chatzimanolis coll. (SEMC). COSTA RICA: 2 specimens, Farm Castilla, VI.[19]38, no collector (FMNH); 1 specimen, Finca Castilla, VI.[19]38, no collector (FMNH); 1 specimen, Chitaría, 23–25.VII.1941, no collector (FMNH); Guanacaste: 1 specimen, Heliconias Biol. Sta., 10
42.920 N 85
02.380 W, 600 m, flight intercept trap, 20–23.XI.2001, R.Brooks coll. (SEMC); Alajuela: 1 specimen, 9.6 km N. Jct Rts. 9 and 120, Rd. to Puerto, Viejo 4600’, litter near stream, 17.III.1991, L. Herman coll. (AMNH); 1 specimen, E.B. Sam Ramon, R.B. San Ramon, 27 km. N. and 8 km. W. San Ramon, 10
130 400 N 84
350 4600 W, flight intercept trap, 8.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); Heredia: 15 specimens, OTS La Selva Field Sta. Puerto Viejo de Sarapiqui, Rio Puerto Viejo, 10
260 N 83
590 W, Ber. 100cc conc. Epiphytic leaf mould w. Peripatus, 5–11.III.1973, J. Wagner and J. Kethley coll. (1 DZUP, 14 FMNH); 1 specimen, La Selva Biol. Station nr. Puerto Viejo de Sarapiqui, 18.II.1985, L. Herman coll. (AMNH); 1 specimen, La Selva, 3.2 km SE Puerto Viejo, 100 m, flight intercept trap, 21.III.1992, W. Bell coll. (SEMC); 1 specimen, 3 km S Puerto Viejo OTS, La Selva, 100 m, X.1992, P. Hanson coll. (SEMC); 2 specimens, Porrosati, 6 km N. San José de la Montana, 1900 m, 10
50 3000 N 84
70 000 W, berlese forest litter, 27.VI.1997, R. Anderson coll. (SEMC); San José: 2 specimens, 117 km. Pan American Highway 19 km N. San Isidro, 9
280 000 N 83
420 2000 W, 1800 m, cloud forest litter, 15.II.1998, R. Anderson coll. (SEMC); 1 specimen, PanAmerican Hwy Km 80.5, 7 km SSW, Cabinas de Quetzal, 9
330 5300 N 83
480 500 W, 2150 m, fogging fungus covered log, 21.VII.2000, J.Ashe, R. Brooks and Z. Falin coll. (SEMC); 1 specimen, PanAmerican Hwy, Km 80.5, 7 km SSW, Cabinas de Quetzal, 9
330 5300 N 83
480 500 W, 2150 m, fogging fungus covered log, 22.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 1 specimen, PanAmerican Hwy. km 80.5, 9.5 km SSW, on San Gerado Rd., Cataraia Trail, 9
320 4700 N 83
480 4000 W, 2020 m, 22.VII.2000, J. Ashe, R. Brooks and Z. Falin coll. (SEMC); 2 specimens, Genesis II Reserve, 09
42.570 N 83
54.640 W, 2360 m, 15.VI.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC); Cartago: 1 specimen, Tapantí, 1100 m, 9.IX.[19]39, no collector coll. (FMNH); 1 specimen, Capellades, 19–21.III.[19]40, no collector (FMNH); 1 specimen, Cañon, 2450 m, 28.XI.1994, H. Forster coll. (NMW); Limón: 1 specimen, Hamburgfarm, 15.XII.[19]24, Nevermann coll. (FMNH); 1 specimen, the same locality and collector, 4.X.[19]28 (FMNH); 3 specimens, the same locality and collector, 1.X.1936 (FMNH); 3 specimens, Finca Castilla, VI.[19]38, no collector (FMNH); Puntarenas: 1 specimen, OTS Sta. Finca Las Cruces, San Vito, 82
580 W 8
460 N, 4000ft., Ber. 1500cc. conc. litter, base of lrg. banana cump, 15.III.1973, J. Wagner and J. Kethley coll. (FMNH); 4 specimens, the same locality and collector, 16.III.1973 (FMNH); 3 specimens, the same locality and collector, Berlese 1500cc. leaf litter in stream bed, away from flowing water steep banks, Virgin forest cover, 18.III.1973 (FMNH); 1 specimen, the same locality and collector, 19.III.1973 (FMNH); 6 specimens, the same locality and collector, Berlese leaf litter in stream bed Rio Jaba, rocky crevices 800cc. conc., 20.III.1973 (FMNH); 2 specimens, Las Cruces Botanical Garden nr. San Vito, 3500ft., 27–28.II.1985, L. Herman coll. (AMNH); 1 specimen, Monte Verde, 1240 m, berlese of upper streambed, in sand, 10.V.1989, J. Ashe, R. Brooks and R. Leschen coll. (SEMC); 2 specimens, 35 km NE San Vito nr. Las Alturas, Río Bella Vista, Rd. to Gravel Pit, 4300., wet leaf litter, 23.III.1991, L. Herman coll. (AMNH); 1 specimen, San Vito, Estac. Biol. Las Alturas, 1500 m, V.1992, P. Hanson coll. (SEMC); 1 specimen, Rincon, Cerro Helado, flight intercept trap, 21–25.VI.1997, S. and J. Peck coll. (SEMC); 1 specimen, Rincon de Osa, 8
41.1410 N 83
31.1170 W, 50 m, 23–26.VI.2001, S. and J. Peck coll. (SEMC); 1 specimen, Las Cruces Biol. Sta., 08
47.140 N 82
57.580 W, 1330 m, rotting banana plant stems, 29.V.2004, J. S. Ashe, Z. Falin and I. Hinojosa coll. (SEMC). PANAMA: Bocas del Toro: 1 specimen, Almirante, trail to dam on Nigua Creek, Berlese: fibrous centre of decayed palm log, 25.III.1959, H. S. Dybas coll. (FMNH); Chiriquí: 1 specimen, La Fortuna, Vivero, 08
420 N 82
140 W, 1150 m, flight intercept trap, 14–18.VI.1994, A. R. Gillogly coll. (SEMC); 3 specimens, 27.7 km W Volcan, Hartmann’s Finca, 8
450 000 N 82
480 000 W, 1800 m, oak forest litter, 16.VI.1995, R. Anderson coll. (SEMC); Panamá: 6 specimens, Cerro Campana, 3200’, wet debris small forest stream, 23.II.[19]76, A. Newton coll. (1 DZUP, 5 FMNH); Darién: 1 specimen, Cana Biological Station, 7
450 1800 N 77
410 600 W, 530 m, 09.VI.1996, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: 1 specimen, Aguatal, no date and collector (FMNH); Cundinamarca: 1 specimen, Finca San Pablo, 8 km N. Alban, 1800 m, 1–12.VIII.1987, P. and B. Wygodzinsky coll. (AMNH); Nariño: 1 specimen, R.N. La Planada Parcela Permanente, 01
150 N 78
150 W, 1885 m, Pitfall, 04–06.XI.2000, G. Oliva coll. (SEMC). VENEZUELA. Lara: 1 specimen, Sanaré, 14.2 km SE Yacambú N.P., 9
410 4500 N 69
360 4800 W, 1650 m, flight intercept trap, 18.V–1.VI1998, J. Ashe, R. Brooks and R. Hanley coll. (SEMC). ECUADOR: Esmeraldas: 16 specimens, Bilsa, 0
200 000 S 79
430 000 W, flight intercept trap, 28.IV–10.V.1996, P. Hibbs coll. (SEMC); 2 specimens, Bilsa Biological Station, 0
200 2400 N 79
420 3600 W, 500 m, Malaise trap, 10.V–4.VI.1996, P. Hibbs coll. (SEMC); 19 specimens, Bilsa, 0
200 000 S 79
430 000 W, flight intercept trap, 10.V–5.VI.1996, P. Hibbs coll. (SEMC); 2 specimens, Bilsa, 0
200 000 S 79
430 000 W, flight intercept trap, 5.VI–7.VII.1996, P. Hibbs coll. (SEMC); 1 specimen, Bilsa Biological Station, 0
200 2400 N 79
420 3600 W, 500 m, flight intercept trap, 7–19.VII.1998, P. Hibbs coll. (SEMC); 1 specimen, the same locality and collector, 19.VII.1996 (SEMC); Sucumbios: 1 specimen, Sacha Lodge, 0
280 1400 S 76
270 3500 W, 270 m, flight intercept trap, 21–24.III.1999, R. Brooks coll. (SEMC); Pichincha: 1 specimen, Prov., 47 km(S). Sto. Domingo, Rio Palenque Sta, Ber. leaf litter with palm fruits, 260 m, 21–24.II.1976, S. Peck coll. (FMNH); 3 specimens, the same locality and collector, Ber. litter under decaying fruit, 250 m, 25.II.1976 (FMNH); 1 specimen, 16 kmE Sto. Domingo Tinalandia, 680 m, malaise rainforest, 25.VII.1985, S. and J. Peck coll. (FMNH); 1 specimen, 17 km SE Sto. Domingo de Colorados, Tinalandia 3000’, light, 16–21.X.[19]88, L. Herman coll. (AMNH); 1 specimen, W of Alluriquin, Tinalandia, 2600–2800ft., litter nr. stream, 19–20.V.1993, L. Herman coll. (AMNH); 3 specimens, 45 km NNW Quito Macquipucuna Station, 1600–1650 m, flight intercept trap, 3–18.IV.1996, P. Hibbs coll. (SEMC); 2 specimens, Rio Palenque Science Center, 0
360 S 79
210 000 W, 200 m, malaise trap, 25.V–6.VII.1996, P. Hibbs coll. (SEMC); 2 specimens, Tinalandia, Santo Domingo, 16 km E, 0
160 5300 S 79
30 3900 W, 750 m, flight intercept trap, 26–27.III.1999, R. Brooks and D. Brzoska coll. (SEMC); Napo: 1 specimen, 33 km N Tena, 59 km E on Loreto Rd, 584 Invertebrate Systematics E. Caron et al. 2,800’, 4.XI.[19]88, blk. Light, L. Herman coll. (AMNH); 1 specimen, Baeza, nr. Rio Papallacta, 5800ft., litter on bark, 24.V.1993, L. Herman coll. (AMNH); 1 specimen, Sierra Azul, Hacienda Aragon, 0
400 000 S 77
550 000 W, 2200 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); 1 specimen, Cosanga, 4.2 km S on Baaza-Tena, Road then 1.5 km W on pipeline access road, 0
370 1900 S 77
500 100 W, 2150 m, flight intercept trap, 5–7. XI.1999, Z. H. Falin coll. (SEMC); Cotopaxi: 2 specimens, N of San Francisco de las Pampas, visc. Rio Esmeraldas, litter and debris, 4400–5000ft., 14–15. V.[19]93, L. Herman coll. (AMNH); 1 specimen, Canton Sigchos Las Pampas, Otonga Natural Reserve, 25–28.VII.2005, W. Roosi coll. (DZUP); Tungurahua: 1 specimen, 12.2 km E Baños, 5000ft., litter near stream, 22.V.[19]93, L. Herman coll. (AMNH). PERU: Huánuco: 1 specimen, Divisoria, 1700 m, IX.1946, F. Woytkowski coll. (AMNH); 5 specimens, Cordillera Azul, 39 km NE Tingo María, 1700 m, montane rainforest, window trap, 11–14. I.1983, A. Newton and M. Thayer coll. (FMNH); Cusco: 1 specimen, pillahuata, Manu rd., km 128, leaf litter, 22.IX.1982, L. E. Watrous and G. Mazurek coll. (FMNH). BOLIVIA: Cochabamba: 1 specimen, 105 km E Yungas, nr. Río Carmen Mayu, Cochabamba-Villa Tunari Rd., 17
80 5100 S 65
430 5000 W, 1750 m, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC). Piestus nevermanni Scheerpeltz, 1952 COSTA RICA: 1 specimen Vara Blanca, VIII.[19]38, no collector (FMNH); Cartago: 1 specimen, P.N. Tapantil, 9
450 4100 N 83
470 500 E, 1150 m, flight intercept trap, 17–20.VII.2000, J. Ashe, R. Brooks, Z. Falin coll. (SEMC). PANAMA: Chiriquí: 2 specimens, La Fortuna, Hydro. Trail, 08
420 N 82
140 W, 1150 m, flight intercept trap, 23.V–9.VI.1995, J. Ashe and R. Brooks coll. (1 DZUP, 1 SEMC); 2 specimens, 5.9 km NE Cerro Punta, 08
220 N 82
340 W, 2400 m, Par. Nac. Volcan Banu, under bark, 14.VI.1995, J. Ashe and R. Brooks coll. (SEMC); 2 specimens, La Fortuna, ‘Hydro Trail’, 08
420 N 82
140 W, 1150 m, treefall litter, 22.V.1995, J. and A. Ashe coll. (SEMC). Piestus paradoxus Bernhauer, 1917 VENEZUELA: Aragua: 15 specimens, 16 km N. Maracay, 10
240 N 67
290 W, 1500 m, under bark, 7.III.1995, R. W. Brooks coll. (1 DZUP, 14 SEMC); 6 specimens, Rancho Grande Biol. Stn., 10
210 000 N 67
410 000 W, 1200–1300 m, fungusy log, 12.V.1998, J. Ashe, R. Brooks and R. Hanley coll. (1 AMNH, 1 DZUP, 4 SEMC). Piestus aper Sharp, 1876 PANAMA: 2 specimens, Barro Colo. Is., 18.VIII.[19]38, E. Williams coll. (FMNH); Bocas del Toro: 1 specimen, Fortuna, Grande road, 8
470 N 82
120 W, 1050 m, premontane rain forest, sifting litter, 12–14.VII.1987, D. M. Olson coll. (FMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd. Km 6.1, 09
070 N 79
450 W, 40 m, flight intercept trap, 7–21.VI.1995, J. Ashe and R. Brooks coll. (SEMC). COLOMBIA: Amazonas: 1 specimen, PNN Amacayacu Matamata, 03
410 S 70
150 W, 150 m, Winkler, 13–15.IX.2000, A. Parente coll. (SEMC). SURINAME: Marowijne: 1 specimen, Perica, 70 km E Paramaribo on East–west Road, 5
400 2800 N 54
360 3100 W, 5 m, flight intercept trap, 31.V–5.VI.1999, Z. H. Falin and B. DeDjin coll. (SEMC); 1 specimen, Palumeu, 3
200 5600 N 55
260 1800 W, 160 m, flight intercept trap, 5–9.VII.1999, Z. H. Falin and D. Konoe coll. (DZUP); 1 specimen, the same locality and collector, 7–8.VII.1999 (SEMC). FRENCH GUIANA: Saint-Laurent-du-Maroni: 2 specimens, Eaux Claires, 3.5 mi N Saul, 3
38–400 N 53
13–140 W, 155–260 m, fig fruit fall, 5–13.X.[19]95, L. Herman coll. (AMNH); 5 specimens, Saül, Sifting, IV.1999, A. Berkov coll. (AMNH); Cayenne: 1 specimen, Roura, 27.4 km SSE, 4
440 2000 N 52
130 2500 W, 280 m, flight intercept trap, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC). PERU: Loreto: 7 specimens, Campamento San Jacinto, 2
18.750 S 75
51.770 W, 175–215 m, flower fall berlese, 2.VII.1993, R. Leschen coll. (SEMC); 4 specimens, the same locality and collector, 10.VII.1993 (SEMC); 2 specimens, the same locality and collector, 11.VII.1993 (SEMC); 9 specimens, 1.5 km N. Teniente Lopez, 2
35.660 S 76
06.920 W, fruit fall, 17.VII.1993, 210–240 m, R. Leschen coll. (1 DZUP, 8 SEMC); Cusco: 2 specimens, Consuelo, Manu rd. km 165, litter under rotten palm, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 1 specimen, the same locality and collector, rotten palm, 3.X.1982 (FMNH); 2 specimens, the same locality and collector, 5.X.1982 (FMNH); 3 specimens, the same locality and collector, 12.X.1982 (FMNH); 6 specimens, the same locality and collector, leaf litter, 13.X.1982 (FMNH); 6 specimens, the same locality and collector, 14.X.1982 (FMNH); Madre de Dios: 1 specimen, Tambopata Lago Sandoval, 12
36.63S 69
2.24W, 936ft., leaf litter winkler, primary rain forest, 18.V.1998, P. Parrillo coll. (FMNH); 1 specimen, Pantiacolla Lodge, 8 km NW, El Mirador Trail, Alto Madre de Dios River, 12
380 3000 S 71
160 4100 W, 800 m, flight intercept trap, 23–26.X.2000, R. Brooks coll. (SEMC). BOLIVIA: Santa Cruz: 3 specimens, 3.7 km SSE Buena Vista, Hotel Flora Y Fauna, 17
29.950 S 63
33.150 W, 400–440 m, Chiquitano for. leaf litter, 5.XI.2002, R. Leschen coll. (SEMC). BRAZIL: Amazonas: 1 specimen, 35 km NE Manaus, Res. Flor. Ducke, 25.VII– 08.VIII.[19]95, Arndt and Gröger coll. (ZMHB); Bahia: 1 specimen, Ilheus, Mata da Esperanca, 14
470 200 S 39
30 4500 W, 50 m, atlantic forest, 29.I.1995, D. Agosti coll. (AMNH); Mato Grosso: 1 specimen, Barra do Tapirape, forest stream near shore, 10.VIII.[19]62, B. Malkin coll. (FMNH). PARAGUAY: Guairá: 2 specimens, Villa Rica, 8.XII.1924, Fr. Schade coll. (NMW); 1 specimen, Melgarejo, Tucuara Creek, flood detritus, 20.X.1994, U. Drechsel coll. (SEMC); Itapúa: 1 specimen, San Pedro Mi, San Rafael Reserve, 26
310 2400 S 55
480 1800 W, 90 m, fruitfall, 27.XI.2000, Z. H. Falin coll. (SEMC). Piestus angularis Fauvel, 1864 MEXICO: Veracruz: 2 specimens, Cordoba, no date, F.A. Fenyes coll. (1 NMW, 1 FMNH); 2 specimens, El Fortin, 8.VIII.1941, H. Dybas coll. (FMNH). BELIZE: Cayo: 1 specimen, Mountain Pine Ridge Area, Rio Frio Cave, riparian litter, 16
580 18.900 N 88
590 46.600 W, 28.V.1997, C. Carlton coll. (SEMC); 1 specimen, Cayo District Chiriqui N.P., Doyle’s Delight nr. Campground, 16
290 3500 N 89
020 4900 W, 1100 m, 19–28.VIII.2007, P. W. Kovarik coll. (FMNH). GUATEMALA: Zacapa: 3 specimens, 3.5 km SE La Union, 1500 m, flight intercept trap, 23–25.VI.1993, R. Brooks and J. Ashe coll. (SEMC). COSTA RICA: 2 specimens, San Pedro Poás, 10.II.1942, no collector (FMNH); 1 specimen, Carrizal, 9–13.IV.[19]44, no collector (FMNH); Alajuela: 1 specimen, Atenas, 13.X.[19]40, no collector (AMNH); 1 specimen, San Ramon, 5 km W, 1220 m, 1–30.XI.1997, P. Hanson coll. (SEMC); Heredia: 1 specimen, OTS, La Selva Field Sta., Puerto Viejo de Sarapiqui, Rio Puerto Viejo, 10
260 N 83
590 W, 5–11.III.1973, J. Wagner and J. Kethley coll. (FMNH); 2 specimens, La Selva, E. River Trail, flight interrcept trap, 11.III.1992, W. Bell coll. (SEMC); San José: 1 specimen, Puriscal, IX.[19]39, ilegible collector (FMNH); Limón: 1 specimen, Reventazon, Hamburg Farm 27.XI.1936, F. Nevermann coll. (FMNH); Puntarenas: 6 specimens, OTS Sta. finca Las Cruces, 4000ft., San Vito, 82
580 W 8
460 N, leaf litter in stream bed, away from flowing water steep banks, virgin forest cover, 19.III.1973, J. Wagner and J. Kethley coll. (FMNH); 2 specimens, the same locality and collector, 20.III.1973 (FMNH); 17 specimens, Las Cruces Botanical, Garden nr. San Vito, 3500ft., 27–28. II.1985, L. Herman coll. (AMNH); 1 specimen, Las Alturas Biol. Sta., 08
56.170 N 82
50.010 W, 1660 m, flight intercept trap, 31.V–3.VI.2004, J. S. Ashe, Revision of Piestus with remarks on related genera Invertebrate Systematics 585 Z. Falin and I. Hinojosa coll. (SEMC); 2 specimens, Altamira Biol. Sta., 09
01.760 N 83
00.490 W, 1510–1600 m, 7.VI.2004, J. S. Ashe, Z. Falin, and I. Hinojosa coll. (SEMC). PANAMA: 1 specimen, Canal Zone, Barro Colorado Is., wet leaves and flood debris forest stream, 6.II.1976, A. Newton coll. (FMNH); 1 specimen, the same locality and collector, 8.II.1976 (FMNH); Chiriquí: 1 specimen, 4 km NW Volcán, 8
490 1900 N 82
400 3100 W, 1450 m, litter near stream, 24.XII.[20]01, L. Herman coll. (AMNH); Colón: 1 specimen, Parque Nac. Soberania, Pipeline Rd., 09
070 N 79
450 W, 17–18.V.1995, J. Joly and C. Chaboo coll. (SEMC). SURINAMEE: Marowijne: 1 specimen, Nassau Mountain, 4
480 3600 N 54
310 1600 W, 500 m, beating treefall, 2.VI.1999, Z. H. Falin coll. (SEMC); Para: 1 specimen, Carolina Creek, 11 km SE, Zanderij Airport, 5
230 3600 N 55
90 2900 W, 30 m, flight intercept trap, 18–20.VI.1999, Z. H. Falin and A. Gangadin coll. (SEMC). FRENCH GUIANA: Saint-Laurent-du-Maroni: 6 specimens, Eaux Claires, N of Saül, 3
390 4400 N 53
130 1700 W, 160 m, 5.X.1995, L. Herman coll. (AMNH); 1 specimen, nr. Eaux Claires, 3.5 mil N Saul, 3
38–400 N 53
13–140 W, 155–260 m, leaf litter near stream, 5–13.X. [19]95, L. H. Herman coll. (AMNH); 1 specimen, Saül, 7 km N, Les Eaux Claires, 3
390 4600 N 53
130 3900 W, 220 m, treefall litter, 4.VI.1997, J. Ashe and R. Brooks coll. (SEMC); Cayenne: 1 specimen, Roura, 39.4 km SSE, 4
320 4300 N 52
80 2600 W, 270 m, treefall litter, 10.VI.1997, J. Ashe and R. Brooks coll. (SEMC). ECUADOR: Esmeraldas: 2 specimens, Bilsa Biological Station, 0
200 2400 N 79
420 3600 W, 500 m, Malaise trap, 28.IV–10.V.1996, P. Hibbs coll. (SEMC); Napo: 1 specimen, Jatun Sacha Biol. Station, 21 km E. Puerto Napo, 400 m, lowland rain for., 7.VII.1994, F. Génier coll. (SEMC); 1 specimen, Sierra Azul, Hacienda Aragon, 0
400 000 S 77
550 000 W, 2300 m, flight intercept trap, 17.II–26.III.1996, P. Hibbs coll. (SEMC); Zamora-Chinchipe: 1 specimen, Río Bombuscaro, 4
70 000 S 78
590 000 W, flight intercept trap, 26.VI–4.VII.1996, P. Hibbs coll. (SEMC). PERU: Cusco: 1 specimen, Consuelo, Manu rd. km 165, litter under rotten palm, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the same localiy and collector, 2.X.1982 (FMNH); 16 specimens, the same locality and collector, 4.X.1982 (FMNH); 26 specimens, the same locality and collector, 5.X.1982 (2 DZUP, 24 FMNH); 11 specimens, the same locality and collector, 6.X.1982 (FMNH); 2 specimens, the same locality and collector, 7.X.1982 (FMNH); 1 specimen, the same locality and collector, 7–8.X.1982 (FMNH); 6 specimens, the same locality and collector, 8.X.1982 (FMNH); 1 specimen, the same locality and collector, 9.X.1982 (FMNH); 7 specimens, the same locality and collector, 12.X.1982 (FMNH); 1 specimen, the same locality and collector, 13–15.X.1982 (FMNH). BOLIVIA: Cochabamba: 3 specimens, Cochabamba, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 17
80 5000 S 65
420 2900 W, 1480 m, flight intercept trap, 1–6.II.1999, F. Génier coll. (SEMC); 26 specimens, Cochabamba, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 17
80 5000 S 65
420 2900 W, 1480 m, flight intercept trap, 6–8.II.1999, R. Hanley coll. (SEMC); 3 specimens, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 17
80 5000 S 65
420 2900 W, 1480 m, flight intercept trap, 8–12.II.1999, R. Hanley coll. (SEMC). BRAZIL: Santa Catarina: 1 specimen, Blumenau, no date and collector (NMW); 1 specimen, no locality, date and collector (FMNH). PARAGUAY: Alto Paraná: 1 specimen Alto Piray, VII-[1]900, Silvestri coll. (FMNH). Piestus rugosus Sharp, 1876 GUYANA: 1 specimen, Kartabo, Bartica, 14.V.1924, no collector (AMNH). ECUADOR: Sucumbios: 3 specimens, Sacha Lodge, 0
280 1400 S 76
270 3500 W, 270 m, under bark, 24.III.1999, R. Brooks coll. (1 DZUP, 2 SEMC). PERU: Cusco: 1 specimen, Marcapata, no date and collector (IRSNB). BOLIVIA: 2 specimens, Yuracaris, no date and collector (IRSNB); Cochabamba: 1 specimen, Cochabamba, 117 km E Yungas, Cochabamba-Villa Tunari Rd., 17
60 3200 S 65
410 1200 W, 1040 m, flight intercept trap, 1–6.II.1999, R. Hanley coll. (SEMC); 2 specimens, 67.5 km NE, Est. Biol. Valle del Sajita, Univ. de Sam Simon, 17
60 3300 S 64
470 5200 W, 300 m, fungus covered log, pyrethrum insecticide fogging, 7.II.1999, R. Hanley coll. (SEMC). BRAZIL: Pará: 1 specimen, no locality and collector, Fry coll. (FMNH); 1 specimen, Aldeia Aracu, Igarape, Gurupu-Umu, Maranhao, 50 km E of Caninde, under bark, V.1963, B. Malkin coll. (FMNH). Hypotelus laevis (Solsky, 1872) PERU: Cusco: 3 specimens, Consuelo, Manu rd. km 165, palm fruit, winkler, 1.X.1982, L. E. Watrous and G. Mazurek coll. (FMNH); 2 specimens, the same locality and collector, 3.X.1982 (FMNH); 2 specimens, the same locality and collector, 4.X.1982 (FMNH); 2 specimens, the same locality and collector, 5.X.1982 (FMNH). Hypotelus andinus (Bernhauer, 1917) COLOMBIA: Boyacá: 1 specimen, SFF Iguague La Planada, 5
250 N 73
270 W 2850 m, Malaise, 11.X–01.XI.2000, P. Reina coll. (SEMC); 1 specimen, the same locality and collector, 01–23.XI.2000 (SEMC). ECUADOR: 1 specimen, Banos, 1500’, 6.V.[19]39, W. C. Macintyre coll. (AMNH). www.publish.csiro.au/journals/is View publication stats