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Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the

genus Rollinia A.St.-Hil

Article  in  Annalen des naturhistorischen Museums in Wien, Serie a · January 2007

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 Ann. Naturhist. Mus. Wien 108 B 191 - 205 Wien, Mai 2007

Monographic studies in the genus Annona L. (Annonaceae):

Inclusion of the genus Rollinia A.ST.-HIL.
H. Rainer*

Abstract
Recent molecular studies in the genus Annona, which included two species of Rollinia and one, formerly
placed in Raimondia, revealed the very close relationship of these genera. If Rollinia would be kept as a
separate entity, Annona would become polyphyletic. No satisfactory possibility was found to split up
Annona for maintaining Rollinia as a distinct genus. On the basis of these findings the necessary nomen-
clatural changes are presented here.
Key words: Annonaceae, Annona, Rollinia, Flora Neotropica, new combinations, new names, nomencla-
ture, systematics.

Zusammenfassung
Neueste molekulare Untersuchungen in der Gattung Annona, unter Einbeziehung von zwei Arten der
Gattung Rollinia und einer Art, die früher zu Raimondia gestellt wurde, haben die sehr engen Beziehungen
der Gattungen untereinander verdeutlicht. Wenn man Rollinia als eigenständige Gattung beibehalten würde,
wäre das Ergebnis eine polyphyletische Gattung Annona. Es wurde keine befriedigende Möglichkeit gefun-
den, Annona so abzugrenzen, dass Rollinia als eigenständige Gattung erhalten bleiben könnte. Aufgrund
dieser Ergebnisse werden die dadurch notwendig gewordenen nomenklatorischen Änderungen in dieser
Arbeit präsentiert.

Introduction

The genus Rollinia was established by Antoine de SAINT-HILAIRE in the “Flora Brasiliae
meridionalis” (1825: 28 - 29, tab. 5) as a result of his collecting activities in the early
19th century in south eastern Brazil. The genus was subsequently expanded by SCHLECH-
TENDAL (1834), MARTIUS (1841), TRIANA & PLANCHON (1862), and SAFFORD (1916b).
SAFFORD (1916a) segregated the new genus Rolliniopsis because some species show a
peculiar fruit morphological pattern, namely almost free but sessile carpidia in ripe fruit-
ing stage. FRIES (1934) gave a first comprehensive overview in the series “Monogra-
phien der Arten einiger Annonaceen-Gattungen”, keeping Rolliniopsis as a separate
genus, even until his treatment of the family for ENGLER & PRANTL’s “Natürliche
Pflanzenfamilien” (FRIES 1959). In MAAS & WESTRA (1989) Rolliniopsis was sunk into
synonymy of Rollinia. The monographic treatment by MAAS et al. (1992), in which 42
species were listed, with two more taxa mentioned as new but not formally described,
represents the up to date state of knowledge. In this monograph MAAS though did not
discuss taxa placed in Annona, but mentioned by FRIES (1934: 113) as being closely

* Heimo Rainer, Department of Systematic and Evolutionary Botany, Faculty Center for Botany,
University of Vienna, Rennweg 14, A-1030 Vienna, Austria. – heimo.rainer@univie.ac.at
192 Annalen des Naturhistorischen Museums in Wien 107 B

related or even representing the group within Annona from where Rollinia might have
derived (FRIES 1943: 8). Some very closely related taxa described in Annona, e.g.,
Annona amazonica R.E.FR. A. duckei DIELS, A. neglecta R.E.FR., and A. symphyocarpa
SANDWITH, all of which show Rollinia-characters (baccate outer petals, though not dor-
sally winged, inner petals connate and adnate to outer ones, fruits with a comparatively
long and thin pedicel, pollen in monads) have been omitted from the discussion on the
systematic position of Rollinia by KOEK-NOORMAN & MAAS (1992).

Results and discussion

On the basis of recent molecular studies in the genus Annona (RAINER et al. in prep.), in
which two taxa of Rollinia (R. cuspidata and R. mucosa) and one of Raimondia (R.
deceptrix) were included, and a phylogenetic analysis (RAINER et al. in prep.) revealed
that Rollinia and Raimondia are nested within Annona. These findings are in full accor-
dance with recently published results on biogeographic aspects in the family Anno-
naceae (RICHARDSON et al. 2004).
As a result of morphological, karyological, and palynological studies, and evidence
from numerous additional publications on seed anatomy (VAN DER HEIJDEN & BOUMAN
1988), palynology (WALKER 1971), and chapters on wood anatomy (TER WELLE 1992),
palynology (LE THOMAS et al. 1992) and karyology (MORAWETZ & HEMMER 1992)
included in the revision of the genus Rollinia by MAAS et al. (1992), it becomes clear
that Rollinia and Raimondia cannot be maintained as separate genera. The genus
Raimondia was already placed in synonymy of Annona in an earlier paper (RAINER
2001) and the necessary changes in Rollinia are presented in this paper to provide the
basis for the monographic treatment of Annona (RAINER in prep.).
In the following new combinations, new names, and reestablishment of names originally
published under Annona are provided. Actually accepted names are given in bold face.
Citation of the type material is added where known, the “!” sign is added after citation
of herbarium acronyms for specimens seen by the author. Herbarium acronyms follow
the Index Herbariorum (HOLMGREN & HOLMGREN 1998-2007).

Annona L., Sp. pl. 536 - 537 (1753)

Type species: Annona muricata L., Sp. pl. 536 (1753).
= Raimondia SAFF., Contr. U. S. Natl. Herb. 16 (5): 217 - 219, pl. 52, 53 (1913).
Type species: Raimondia monoica SAFF. (= Annona cherimolioides TRIANA & PLANCH.)
= Rollinia A.ST.-HIL., Fl. Bras. merid. 1 (1): 28, tab. 5 (1825)
Lectotype species (designated by HUTCHINSON 1923: 260): Rollinia longifolia A.ST.-HIL.
(= Annona dolabripetala RADDI)
Contrary to the citation of Annona dolabripetala RADDI [≡ Rollinia dolabripetala
(RADDI) R.E.FR.] as the lectotype for the genus (MAAS 1992: 65; GREUTER et al.
1993: 1000), which follows the citation of Annona dolabripetala RADDI in PFEIFFER
(1874: 984) the latter did neither make the new combination for “Rollinia
dolabripetala” nor selected this taxon as the type species for the genus Rollinia but
merely listed “Annona dolabripetala RADDI” in brackets without any comments.
RAINER: Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the genus Rollinia 193

= Rolliniopsis SAFF., J. Wash. Acad. Sci. 6: 198 - 204, fig. 1 - 2 (1916)

Type species (SAFFORD 1916a): Rolliniopsis discreta SAFF. (= Annona leptopetala
(R.E.FR.) H.RAINER)

New combinations

Annona andicola (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia andicola MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol.
Med. Sci. 92 (3): 297, fig. 1, 2 (1989)
Type: J.A. Steyermark 52903 [holotype: F]

Annona bahiensis (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia bahiensis MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol.
Med. Sci. 92 (3): 299, fig. 3, 4 (1989)
Type: S.A. Mori & J.A. Kallunki 9888 [holotype: CEPEC; isotype: U!]

Annona boliviana (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia boliviana R.E.FR., Acta Horti Berg. 13 (3): 115 (1941)
Type: B.A. Krukoff 11258 [holotype: S!; isotypes: A!, F!, G!, MO!, NY!, S!, U!]

Annona calcarata (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia calcarata R.E.FR., Acta Horti Berg. 12 (2): 279 - 280 (1937)
Type: B.A. Krukoff 5315 [holotype: S!; isotypes: A!, B!, BM!, F!, G!, M!, MICH,
MO!, NY!, U!, UC]

Annona centrantha (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia centrantha R. E. FR., Acta Horti Berg. 12 (1): 181 - 182, fig. 13c - f (1934)
Type: E.P. Killip & A.C. Smith 27073 [holotype: US!; isotypes: B!, F!, NY!, S!]

Annona cuspidata (MART.) H.RAINER, comb.n.

≡ Rollinia cuspidata MART., Fl. bras. 13 (1): 20, tab. 6: fig. 2 (1841)
Type: C.F.P. von Martius 2725 [lectotype (selected by MAAS 1992: 80): M!; isotypes:
B (fragm.)!, M!]
= Rollinia cardiantha DIELS in PILGER, Verh. Bot. Vereins Prov. Brandenburg 47: 134
(1905)
Type: E. Ule 5002 [holotype: B!; isotypes: G!, MG, S!]
= Rollinia pachyptera DIELS in MILDBRAED, Notizbl. bot. Gart. Berlin-Dahlem 10: 176
(1927)
Type: G. Tessmann 4704 [holotype: B!; isotypes: F!, G!, S!]
= Rollinia uniflora R.E.FR., Acta Horti Berg. 12 (1): 182 - 184, fig. 12c - d (1934)
Type: R. Spruce s.n., Nov. 1856 [holotype: K!; isotypes: K!, S!]
= Rollinia sphaerantha R.E.FR., Acta Horti Berg. 12 (1): 184, fig. 13a - b (1934)
Type: E.P. Killip & A.C. Smith 27650 [holotype: US!; isotypes: B!, F, NY!, S!]
194 Annalen des Naturhistorischen Museums in Wien 107 B

Annona danforthii (STANDL.) H.RAINER, comb.n.

≡ Rollinia danforthii STANDL., Publ. Field Mus. Nat. Hist., Bot. Ser. 18 (2): 444 (1937)
Type: R.E. Danforth 30 [holotype: F; isotypes: BM!, GH!, S!]
= Rollinia chocoensis R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl., ser. 3, 24 (10):
18 (1948)
Type: E.P. Killip 35501 [holotype: S!; isotypes: NY!, UC, US!]

Annona dolichopetala (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia dolichopetala R.E.FR., Ark. Bot., n. s. 4 (2): 26, tab. 3 (1959)
≡ Rollinia dolichopetala var. suberecta R.E.FR., Ark. Bot., n. s. 4 (2): 27, tab. 3 p.p. (1959)
Type: E. Asplund 19768 [holotype: S!; isotypes: C, G, K!, NY!, P!, S!]
= Rollinia dolichopetala var. divergens R.E.FR., Ark. Bot., n. s. 4 (2): 27, tab. 3 p.p.
(1959)
Type: E. Asplund 18467 [holotype: S!; isotypes: C, G, K!, NY!, P!, S!]

Annona edulis (TRIANA & PLANCH.) H.RAINER, comb.n.

≡ Rollinia edulis TRIANA & PLANCH., Ann. Sci. Nat., Bot., sér. 4, 17: 30 (1862)
Type: J.J. Triana s.n., s.d. [holotype: P!; isotypes: BM!, G, K!]
= Rollinia edulis var. acuta R.E.FR., Ark. Bot., n. s. 1 (6): 346 (1950)
Type: J. Cuatrecasas 4665-A [holotype: US]
= Rollinia edulis var. macropus R.E.FR., Ark. Bot., n. s. 1 (6): 346 (1950)
Type: J. Cuatrecasas 11095 [syntypes: COL, US]; J. Cuatrecasas 11140 [syntypes:
COL, US]

Annona emarginata (SCHLTDL.) H.RAINER, comb.n.

≡ Rollinia emarginata SCHLTDL., Linnaea 9 (3): 318 (1834)
Type: F. Sellow s.n., s.d. [lectotype (selected by MAAS 1992: 95): B!]; F. Sellow 2137
[syntype: B (photos: F, GH!, MO!, NY!)]; F. Sellow 4899 [syntypes: G, LE!]; F.
Sellow 5866 [syntypes: G, HAL!, L!, LE!]
= Rollinia glaucescens SOND., Linnaea 22: 557 (1849)
≡ Rollinia sonderiana WALP., Ann. bot. syst. 3 (5): 813 (1852), nom. illeg.
Type: A.F. Regnell II 5 b [syntypes: S (6 sheets)!]
= Rollinia longifolia var. paraguariensis CHODAT, Bull. Herb. Boissier 6, App. 1: 11
(1898)
Type: E. Hassler 1720 [holotype: G; isotypes: K!, NY!]
= Rollinia hassleriana R.E.FR. in CHODAT & HASSLER, Bull. Herb. Boissier, sér. 2, 4:
1172 (1904)
Type: E. Hassler 7862 [holotype: G]
= Rollinia intermedia R.E.FR. in CHODAT & HASSLER, Bull. Herb. Boissier, sér. 2, 4:
1172 (1904)
Type: E. Hassler 3475 [lectotype (selected by FRIES 1934: 172): G; isolectotypes: A!,
B!, K!, MICH, NY!]; E. Hassler 4797 [syntypes: F, G, K!]; E. Hassler 5202 [syn-
types: G, GH!, K!, NY!, UC]
RAINER: Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the genus Rollinia 195

= Rollinia longipetala R.E.FR., in CHODAT & HASSLER: Bull. Herb. Boissier, sér. 2, 4:
1173 (1904)
≡ Rollinia emarginata SCHLTDL. var. longipetala (R.E.FR.) R.E.FR., Acta Horti
Berg. 12 (1): 169 - 170, fig. 6 (1934)
Type: E. Hassler 7407 [holotype: G; isotypes: A!, B!, F, G, K!, MO!, NY!, S!, U!, UC]
= Rollinia odoriflora ROJAS ACOSTA, Bull. Acad. Int. Géogr. Bot. 26: 163 (1918)
Type: Anonymous collector s.n., s.d. [type not located]
= Rollinia occidentalis R.E.FR., Acta Horti Berg. 12 (1): 165 - 166, tab. 18 (1934)
Type: R.W. Pearce s.n., Nov. 1866 [holotype: K!]
= Rollinia glaziovii R.E.FR., Acta Horti Berg. 12 (1): 166, 168, fig. 10a - c (1934)
Type: A.F.M. Glaziou 12407 [holotype: B!; isotypes: A!, C, K!, LE!, MG, P!, S!]
= Rollinia hassleriana var. vestita R.E.FR., Acta Horti Berg. 12 (1): 171 (1934)
Type: T.C.J. Herzog 1283 [holotype: L!; isotype: S (fragm.)!]

Annona fendleri (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia fendleri R.E.FR., Acta Horti Berg. 12 (1): 152, tab. 15 (1934)
Type: A. Fendler 196 [holotype: K!; isotypes: G!, GH!, GOET, MO!, OXF!]
= Rollinia subracemosa PITTIER, Bol. Soc. Venez. Ci. Nat. 4: 349 (1938)
Type: Ll. Williams 10011 [holotype: VEN!; isotypes: A!, F, MICH, S (fragm.)!]

Annona ferruginea (R.E.FR.) H.RAINER, comb.n.

≡ Rolliniopsis ferruginea R.E.FR., Acta Horti Berg. 12 (1): 196 (1934)
≡ Rollinia ferruginea (R.E.FR.) MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch.,
Ser. C, Biol. Med. Sci. 92 (3): 301 (1989)
Type: C. Gaudichaud 801 [holotype: B (destr.), lectotype (selected by MAAS &
WESTRA 1989: 301): P!; isotypes: G, P!]

Annona fosteri (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia fosteri MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol.
Med. Sci. 92 (3): 302, fig. 6 (1989)
Type: R.B. Foster 9657 B [holotype: U!; isotypes: F, MO!]

Annona glomerulifera (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia glomerulifera MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C,
Biol. Med. Sci. 92 (3): 303, fig. 7 (1989)
Type: R. Vásquez & N. Jaramillo 790 [holotype: MO!]

Annona helosioides (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia helosioides MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C,
Biol. Med. Sci. 92 (3): 305, fig. 8 (1989)
Type: W. Palacios 1295 [holotype: MO!; isotype: QCNE!]

Annona herzogii (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia herzogii R.E.FR., Acta Horti Berg. 12 (1): 177 - 178 (1934)
Type: T.C.J. Herzog 1368 [holotype: L!; isotypes: G, S!]
196 Annalen des Naturhistorischen Museums in Wien 107 B

Annona hispida (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia hispida MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol.
Med. Sci. 92 (3): 307, fig. 9 (1989)
Type: J. Schunke V. 2657 [holotype: F; isotypes: COL, G, GH!, NY!, K!, P!, S!]

Annona jucunda (DIELS) H.RAINER, comb.n.

≡ Rollinia jucunda DIELS in MILDBRAED, Notizbl. bot. Gart. Berlin-Dahlem 10: 177
(1927)
Type: G. Tessmann 5341 [holotype: B!; isotypes: F, G, NY!, S!]
= Rollinia peruviana DIELS in PILGER, Verh. Bot. Vereins Prov. Brandenburg 47: 135
(1905)
Type: E. Ule 6426 [holotype: B!; isotypes: F, G, K!, L!, MG, S!]
non: Annona peruviana HUMB. & BONPL. ex DUNAL, Monogr. Anonac. 67 (1817),
(= Annona glabra L.)
= Rollinia rigidiflora R.E.FR., Acta Horti Berg. 12 (1): 154, fig. 9 (1934)
Type: Ll. Williams 3886 [holotype: F; isotype: S (fragm.)!]

Annona leptopetala (R.E.FR.) H.RAINER, comb.n.

≡ Rollinia leptopetala R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl. 34 (5): 50 - 51,
tab. 7: fig. 3 - 4 (1900)
≡ Rolliniopsis leptopetala (R.E.FR.) SAFF., J. Wash. Acad. Sci. 6: 191 (1916)
Type: G. Gardner 2033 [lectotype (selected by FRIES 1934: 191): B!; isolectotypes:
BM!, G, GH!, K!, MG, NY!, OXF!, P!]; J. Blanchet 3101 [syntypes: BM!, BR, C, F,
G, GH!, K!, MG, NY!, OXF!, P!]
= Rolliniopsis discreta SAFF, J. Wash. Acad. Sci. 6: 199, fig. 1 - 2 (1916)
Type: P.H. Dorsett 371 b [holotype: US!]
= Rolliniopsis simiarum SAFF., J. Wash. Acad. Sci. 6: 201 (1916)
Type: J.N. Rose & P.G. Russell 19963 [holotype: US!; isotype: GH!]
= Rollinia leptopetala var. angustifolia R.E.FR., Kongl. Svenska Vetensk.-Akad.
Handl. 34 (5): 51, tab. 7: fig. 5 (1900)
Type: A.F.M. Glaziou 13508 [holotype: B!; isotypes: C, K!, LE!, P!]

Annona mammifera (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia mammifera MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C,
Biol. Med. Sci. 92 (3): 308, fig. 10 (1989)
Type: P.J.M. Maas et al. 6025 [holotype: U!; isotypes: COL, K!, MO!, NY!, USM!]

Annona maritima (ZÁCHIA) H.RAINER, comb.n.

≡ Rollinia maritima ZÁCHIA, Bradea 6 (28): 242 - 247, fig. (1993)
Type: B. Rambo s.n. (= PACA 54826) [holotype: B!; isotypes: HBR!, PACA]

Annona pachyantha (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia pachyantha MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C,
Biol. Med. Sci. 92 (3): 310, fig. 11 (1989)
Type: A.H. Gentry et al. 53389 [holotype: MO!; isotype: U!]
RAINER: Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the genus Rollinia 197

Annona papilionella (DIELS) H.RAINER, comb.n.

≡ Rollinia papilionella DIELS, in MILDBRAED, Notizbl. bot. Gart. Berlin-Dahlem 10:
176 (1927)
Type: G. Tessmann 4664 [holotype: B!; isotypes: G, F (fragm.), GH (fragm.)!, NY
(fragm.)!]
= Rollinia pittieri SAFF., J. Wash. Acad. Sci. 6: 376 (1916)
Type: H.F. Pittier 4358 [holotype: US!]
non: Annona pittieri DONN.SM., Bot. Gaz. (Crawfordsville) 24 (6): 389 (1897)
= Rollinia microsepala STANDL., Publ. Field Columbian Mus. Bot. Ser. 4 (8): 205
(1929)
Type: G.P. Cooper & G.M. Slater 102 [holotype: F; isotypes: G (fragm.), S (fragm.)!]

Annona parviflora (A.ST.-HIL.) H.RAINER, comb.n.

≡ Rollinia parviflora A.ST.-HIL., Fl. Bras. merid. 1 (1): 30 (1825)
≡ Rolliniopsis parviflora (A.ST.-HIL.) SAFF., J. Wash. Acad. Sci. 6: 202 (1916)
≡ Rollinia parviflora var. angustifolia MART., Fl. bras. 13 (1): 19 (1841)
≡ Rolliniopsis parviflora var. angustifolia (MART.) R.E.FR., Acta Horti Berg. 12 (1):
195 - 196 (1934)
Type: A.F.C.P. de Saint-Hilaire 454 [holotype: P!]
non: Annona parviflora RUIZ & PAV. in LÓPEZ, Anales Inst. Bot. Cavanilles 17 (1):
427 - 428, tab. 485 (1959), nom.inval.; cf. MCNEILL et al. 2006: § 37. 1, (= Annona
mucosa JACQ.)
= Guatteria apodocarpa MART., Fl. bras. 13 (1): 30 (1841)
Type: C.F.P. von Martius s.n., Aug. 1817 [holotype: M!; isotype: B (fragm.)!]
= Rollinia parviflora var. latifolia MART., Fl. bras. 13 (1): 19 (1841)
≡ Rolliniopsis parviflora var. latifolia (MART.) R.E.FR., Acta Horti Berg. 12 (1): 195
(1934)
Type: J. Lhotsky s.n., s.d. [syntypes: B! (photos: F, GH!, MO!, NY!, U!), G, HAL!]

Annona pickelii (DIELS) H.RAINER, comb.n.

≡ Rollinia pickelii DIELS in PILGER, Notizbl. bot. Gart. Berlin-Dahlem 11: 783 (1933)
Type: B.D. Pickel 3004 p.p. [holotype: B!, isotype: SPSF (?)]

Annona rensoniana (STANDL.) H.RAINER, comb.n.

≡ Rollinia rensoniana STANDL., J. Wash. Acad. Sci. 13: 351 (1923)
Type: P.C. Standley 23022 [holotype: US!; isotypes: GH!, NY!]
= Rollinia membranacea TRIANA & PLANCH., Ann. Sci. Nat., Bot., sér. 4, 17: 31 (1862)
Type: J. Goudot s.n., Mar. 1844 [lectotype (selected by MAAS 1992: 124): P!; isolec-
totype: K!]; J.J. Triana s.n., s.d. [syntype: COL]
non: Annona membranacea R.E.FR., Acta Horti Berg. 13 (3): 115, fig. 2a (1941)
= Rollinia standleyi R.E.FR., Acta Horti Berg. 12 (1): 128 - 129 (1934)
Type: P.C. Standley & J. Valerio 45920 [holotype: US!; isotype: S!]
= Rollinia cherimolioides R.E.FR., Acta Horti Berg. 12 (1): 133 - 134 (1934)
Type: H. Karsten s.n., s.d. [holotype: W (destr., photos: A, F, MO!); isotype: S!]
198 Annalen des Naturhistorischen Museums in Wien 107 B

= Rollinia mexicana STANDL., Publ. Field Mus. Nat. Hist., Bot. Ser. 11 (5): 155 (1936)
Type: E. Bourgeau 2448 [holotype: LE!; isotypes: BR, F, G, GH!, L!, P!, S!]
= Rollinia rensoniana var. granatensis R.E.FR., Acta Horti Berg. 12 (1): 130 (1934)
Type: J. Mutis 3842 [holotype: US!; isotypes: MA!, S (fragm.)!]

Annona rufinervis (TRIANA & PLANCH.) H.RAINER, comb.n.

≡ Rollinia rufinervis TRIANA & PLANCH., Ann. Sci. Nat., Bot., sér. 4, 17: 30 (1862)
Type: J. Goudot s.n., Sep. 1844 or Sep. 1845 [holotype: P!; isotypes: G, K!]

Annona rugulosa (SCHLTDL.) H.RAINER, comb.n.

≡ Rollinia rugulosa SCHLTDL., Linnaea 9 (3): 316 (1834)
Type: F. Sellow s.n., s.d. [holotype: B!]
= Rollinia rugulosa subsp. australis R.E.FR., Acta Horti Berg. 12 (1): 164 - 165 (1934)
Type: J. Tweedie s.n., s.d. [holotype: K!; isotypes: E, S (fragm.)!]

Annona schunkei (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia schunkei MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol.
Med. Sci. 92 (3): 311 - 313, fig. 12 - 13 (1989)
Type: J. Schunke V. 14117 [holotype: U!; isotypes: F, K!, MO!, NY!, P!, U!, USM!,
WU!]

Annona ubatubensis (MAAS & WESTRA) H.RAINER, comb.n.

≡ Rollinia ubatubensis MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C,
Biol. Med. Sci. 92 (3): 313, fig. 14 - 15 (1989)
Type: P.E. Gibbs et al. 3501 [holotype: SP!; isotypes: E, NY!]

Annona williamsii (RUSBY ex R.E.FR.) H.RAINER, comb.n.

≡ Rollinia williamsii RUSBY ex R.E.FR., Acta Horti Berg. 12 (1): 186 - 187 (1934)
≡ Rollinia williamsii RUSBY, Proc. Intern. Congr. Pl. Sci. Ithaca 2: 1373 (1929), nom.
nud.
Type: R.S. Williams 370 [lectotype (cited by MAAS 1992: 156): NY!; isotypes: K!, S!]

New names

Annona neoamazonica H.RAINER, nom.n.

≡ Rollinia amazonica R.E.FR. Ark. Bot., n. s. 3 (18): 602 (1957)
Type: R.E. Schultes 6758 [holotype: US!; isotype: S (fragm.)!]
non: Annona amazonica R.E.FR., Acta Horti Berg. 10 (2): 266 (1931)

Annona neochrysocarpa H.RAINER, nom.n.

≡ Rollinia chrysocarpa MAAS & WESTRA, Proc. Kon. Ned. Akad. Wetensch., Ser. C,
Biol. Med. Sci. 92 (3): 300, fig. 5 (1989)
Type: B. Berlin 738 [holotype: MO!; isotype: U!]
non: Annona chrysocarpa LEPR. ex GUILL. & PERR., Fl. Seneg. tent. 1: 6 (1831),
(= Annona glabra L.)
RAINER: Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the genus Rollinia 199

Annona neoecuadorensis H.RAINER, nom.n.

≡ Rollinia ecuadorensis R.E.FR., Acta Horti Berg. 12 (3): 566 (1939)
Type: H. Schultze-Rhonhof 2155 [holotype: B (destr.?); isotype: S!]
non: Annona ecuadorensis R.E.FR., Ark. Bot., n. s. 4 (2): 25 (1959)

Annona neoelliptica H.RAINER & MAAS, nom.n.

≡ Rollinia elliptica R.E.FR., Acta Horti Berg. 12 (1): 160 (1934)
Type: BW (Boschwezen Suriname) 6507 [holotype: U!; isotypes: BM!, BR, COL,
K!, NY!, RB!, S!, U!, VEN!]
non: Annona elliptica R.E.FR., Ark. Bot. 21A (9): 13, fig. 3: 1 - 3 (1927), (= Annona
cascarilloides C.WRIGHT ex GRISEB.)

Annona neoinsignis H.RAINER, nom.n.

≡ Rollinia insignis R.E.FR., Acta Horti Berg. 12 (1): 161 - 162, tab. 17 (1934)
Type: Ll. Williams 2685 [holotype: F; isotypes: B!, S!]
non: Annona insignis R.E.FR., Acta Horti Berg. 12 (1): 214 - 215 (1934)
= Rollinia insignis var. pallida R.E.FR., Acta Horti Berg. 12 (3): 565 (1939)
Type: A. Ducke 141 [holotype: S!; isotypes: A!, F, K!, MO!, NY!, RB!, S!]

Annona neolaurifolia H.RAINER, nom.n.

≡ Rollinia laurifolia SCHLTDL., Linnaea 9 (3): 319 (1834)
≡ Rollinia laurifolia var. erecta R.E.FR., Acta Horti Berg. 12 (1): 150, fig. 6e (1934)
Type: F. Sellow 1190 [lectotype (selected by FRIES 1934: 149): B!]; F. Sellow 809
[syntype: LE!]
non: Annona laurifolia DUNAL, Monogr. Anonac. 65 (1817), (= Annona glabra L.)
= Rollinia laurifolia var. divergens R.E.FR., Acta Horti Berg. 12 (1): 150, fig. 6f - g
(1934)
Type: Y. Mexía 5311 [holotype: S!; isotypes: A!, BM!, F, G, GB, K!, MICH, MO!,
NY!, S!, U!, UC]
= Rollinia laurifolia var. longipes R.E.FR., Ark. Bot. 5 (4): 20, tab. 1: fig. 3 - 4 (1905)
≡ Rollinia laurifolia forma longipes (R.E.FR.) R.E.FR., Acta Horti Berg. 12 (1): 150
(1934)
Type: L. Riedel s.n., Oct. 1823 [holotype: S!; isotypes: A!, K!, LE!]
= Rollinia laurifolia var. reflexa R.E.FR., Acta Horti Berg. 12 (1): 150 - 151, fig. 6h
(1934)
Type: Y. Mexía 5350 [holotype: S!; isotypes: A!, BM!, F, G, GB, MICH, MO!, NY!,
S!, U!]

Annona neosalicifolia H.RAINER, nom.n.

≡ Rollinia salicifolia SCHLTDL., Linnaea 9 (3): 317 (1834)
Type: F. Sellow s.n., s.d. [lectotype (selected by RAINER 2003: 4): HAL!; isolecto-
types: B!, G, HAL!, L!, LE!]; F. Sellow 1498 [syntype: B!]; L.C. von Chamisso s.n.,
s.d. [syntype: B?]
non: Annona salicifolia EKMAN & R.E.FR., Ark. Bot. 22B (2): 2, fig. 1a - i (1929)
200 Annalen des Naturhistorischen Museums in Wien 107 B

Annona neosericea H.RAINER, nom.n.

≡ Rollinia dolabripetala var. sericea R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl.
34 (5): 45 (1900)
≡ Rollinia sericea (R.E.FR.) R.E.FR., Acta Horti Berg. 12 (1): 152 - 154 (1934)
Type: F. Sellow 684 [lectotype (selected by MAAS 1992: 145): B!]; F. Sellow 489
[syntypes: B!, UC]; F. Sellow 490 [syntypes: B!, UC]
non: Annona sericea DUNAL, Monogr. Anonac. 69, tab. 5 (1817)
= Rollinia sericea var. longisepala R.E.FR., Acta Horti Berg. 12 (1): 153 - 154 (1934)
Type: A. von Luederwaldt 8900 [holotype: S!; isotype: SP!]

Annona neoulei H.RAINER, nom.n.

≡ Rollinia ulei DIELS in PILGER, Verh. Bot. Vereins Prov. Brandenburg 47: 136 (1905)
Type: E. Ule 6425 [holotype: B!; isotypes: F (fragm.), G, L!, MG, S (fragm.)!]
non: Annona ulei R.E.FR., Acta Horti Berg. 10 (2): 243, tab. 15 (1931), (= Annona
jahnii SAFF.)
= Annona microcarpa RUIZ & PAV. ex G.DON, Gen. hist. 1: 90 (1831)
≡ Rollinia microcarpa (RUIZ & PAV. ex G.DON) R.E.FR., Acta Horti Berg. 12 (1):
187 - 188, tab. 20 (1934)
Type: H. Ruiz s.n., s.d. [lectotype (selected by MAAS 1992: 153): B! (photo K!); iso-
types: G, MA!]
non: Annona microcarpa JACQ., Fragm. bot. 6: 40, tab. 44: fig. 7 (1809), (= Diospyros
cargillia F. MUELL.)

Annona neovelutina H.RAINER, nom.n.

≡ Rollinia velutina VAN MARLE, Proc. Kon. Ned. Akad. Wetensch., Ser. C, Biol. Med.
Sci. 92 (3): 314, fig. 16 (1989)
Type: J.A. Steyermark & M. Rabe 96554 [holotype: VEN!; isotypes: NY!, S!]
non: Annona velutina A.ST.-HIL. & TUL., Ann. Sci. Nat., Bot., sér. 2, 17: 132 (1842),
(= Annona crotonifolia MART.)

Reestablished names

Annona dolabripetala RADDI, Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. 18
(2): 394 (1820)
≡ Rollinia dolabripetala (RADDI) R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl. 34
(5): 45 (1900)
≡ Annona xestropetala SPRENG., Syst. veg., ed. 16 (Sprengel) 2: 641 (1825), nom. illeg.
Type: neotype (designated by MAAS 1992: 85, 88): tab. 5, in SAINT-HILAIRE, Fl. Bras.
merid., vol. 1(1) (1825)
= Rollinia longifolia A.ST.-HIL., Fl. Bras. merid. 1 (1): 29, tab. 5 (1825)
Type: A.F.C.P. de Saint-Hilaire s.n., s.d. [syntypes: F, G, P!]
= Annona minensis GLAZ., Mém. Soc. Bot. Fr. 3 (a): 14 (1905)
≡ Rollinia minensis (GLAZ.) R.E.FR., Acta Horti Berg. 12 (1): 151 - 152, tab. 14 (1934)
Type: A.F.M. Glaziou 14467 [holotype: P!; isotypes: C, K!, P!, S!]
RAINER: Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the genus Rollinia 201

Annona exsucca DC. in DUNAL, Monogr. Anonac. 77 (1817)

≡ Rollinia exsucca (DC.) A.DC., Mém. Soc. Phys. Genève 5: 199, tab. 2: fig. A (1832)
Type: A. Anderson s.n., s.d. [holotype: G; isotypes: B!, BM!]
= Rollinia puberula A.DC., Mém. Soc. Phys. Genève 5: 200, tab. 1 (1832)
Type: Anonymous collector s.n., s.d. [holotype: G!; isotype: B (fragm.)!]
= Rollinia multiflora SPLITG., Tijdschr. Natuurl. Gesch. Physiol. 9: 97 (1842)
Type: F.L. Splitgerber 595 [holotype: L!; isotypes: K!, L!, S (fragm.)!]
= Rollinia tinifolia KLOTZSCH in SCHOMBURGK, Reis. Br.-Guiana 3: 1163 (1849)
Type: M.R. Schomburgk 1302 [holotype: B!]
= Rollinia glaucescens MIQ., Stirp. surinam. select. 108 (1850)
≡ Rollinia resinosa SPRUCE ex BENTH., Hooker's J. Bot. Kew Gard. Misc. 5: 9
(1853)
≡ Rollinia exsucca subsp. resinosa (SPRUCE ex BENTH.) R.E.FR., Acta Horti Berg.
12 (1): 144 - 145 (1934)
Type: H.C. Focke 642 [holotype: U; isotypes: K, S (fragm.)]
= Rollinia incurva S.MOORE, Trans. Linn. Soc. London, Bot. 4 (3): 303 (1895)
Type: S.L.M. Moore 522 [holotype: BM!; isotypes: B!, K!, NY!, P!, WU!]
= Rollinia broadwayi R.E.FR., Acta Horti Berg. 12 (1): 145 - 147 (1934)
Type: W.E. Broadway 7580 [holotype: S!; isotypes: F, G, K!, MO!, S!, US!]
= Rollinia brevipes R.E.FR., Acta Horti Berg. 12 (1): 147 (1934)
Type: F.C. Hoehne 5129 [holotype: S!]
= Rollinia gardneri R.E.FR., Acta Horti Berg. 12 (1): 157 - 158, tab. 16 (1934)
Type: G. Gardner 1914 [holotype: W (destr.); lectotype (selected here): BM!; iso-
types: F, G, K!, NY!, OXF!, P!]
= Rollinia surinamensis R.E.FR., Acta Horti Berg. 12 (1): 147, 149, fig. 8a - b (1934)
Type: A.A. Pulle 230 [holotype: U!; isotype: S!]
= Rollinia procera R.E.FR., Acta Horti Berg. 12 (3): 565 (1939)
Type: B.A. Krukoff 6881 [holotype: S!; isotypes: A!, BM!, BR, F, G, K!, MICH,
MO!, NY!, U!, US!]
= Rollinia exsucca subsp. elongata R.E.FR., Acta Horti Berg. 12 (1): 145 (1934)
Type: A. Ducke s.n. (= RB 19645) [holotype: S!; isotypes: K!, RB!, US!]
= Rollinia broadwayi var. cuneata R.E.FR., Acta Horti Berg. 12 (1): 146 - 147 (1934)
Type: H.A. Gleason 576 [holotype: US!; isotypes: GH, K!, NY!, S (fragm.)!]

Annona mucosa JACQ., Observ. bot. (Jacquin) 1: 16 (1764)

≡ Annona muscosa JACQ. in AUBLET, Hist. pl. Guiane 1: 618 (1775), orth. err.
≡ Rollinia mucosa (JACQ.) BAILL., Adansonia 8: 268 (1868)
≡ Annona reticulata var. mucosa (JACQ.) WILLD., Sp. pl., ed. 4., 2 (2): 1266 (1799)
Type: N. von Jacquin s.n., s.d. (not located); P. Duss 4620 [neotype (designated by
MAAS 1992: 126): NY!]
202 Annalen des Naturhistorischen Museums in Wien 107 B

= Annona obtusiflora TUSSAC, Fl. Antill. 1: 191, tab. 28 (1813)

≡ Annona obtusifolia DC., Prodr. (DC.) 1: 84 (1824), orth. err.
Type: tab. 28 in TUSSAC, Fl. Antill. (1813)
= Rollinia biflora RUIZ & PAV. ex G.DON, Gen. hist. 1: 91 (1831)
≡ Annona biflora RUIZ & PAV. ex G.DON, Gen. hist. 1: 91 (1831), nom.nud.
Type: J.A. Pavón s.n., s.d. [lectotype (selected here): MA!; isotypes: G, OXF!]
= Rollinia pterocarpa G.DON, Gen. hist. 1: 91 (1831)
≡ Annona pterocarpa RUIZ & PAV. ex G.DON, Gen. hist. 1: 91 (1831), nom.nud.
≡ Annona pteropetala RUIZ & PAV. ex R.E.FR., Acta Horti Berg. 10 (2): 314 (1931),
nom.nud.
≡ Annona pteropetala RUIZ & PAV. in LÓPEZ, Anales Inst. Bot. Cavanilles 17 (1):
426 - 427, tab. 484 (1959), nom.inval.
Type: H. Ruiz s.n., s.d. [lectotype (selected here): MA!; isotypes: B!, G, P!]
= Rollinia micrantha G.DON, Gen. hist. 1: 91 (1831)
≡ Annona parviflora RUIZ & PAV. ex G.DON, Gen. hist. 1: 91 (1831), nom.nud.
≡ Annona parviflora RUIZ & PAV. ex R.E.FR., Acta Horti Berg. 12 (1): 189 (1934),
nom.nud.
≡ Annona parviflora RUIZ & PAV. in LÓPEZ, Anales Inst. Bot. Cavanilles 17 (1): 427
- 428, tab. 485 (1959), nom.inval.
Type: H. Ruiz s.n., s.d. [type: MA!]
= Rollinia orthopetala A.DC., Mém. Soc. Phys. Genève 5: 200 (1832)
Type: C.S. Parker 238 [holotype: G; isotype: E]
= Rollinia sieberi A.DC., Mém. Soc. Phys. Genève 5: 200, tab. 2: fig. B (1832)
Type: Franz W. Sieber, Fl. Trinitatis 96 [holotype: G-DC; isotypes: BP (3 sheets)!,
E, G, L!, LE!, M!, MO!, P!, PRC (4 sheets)!, S!, WU!]
= Rollinia pulchrinervia A.DC., Mém. Soc. Phys. Genève 5: 201 (1832)
Type: Anonymous collector s.n., s.d. [holotype: G-DC]
= Annona biflora SESSÉ & MOÇ., Fl. mexic., ed. 2, 133 (1894)
Type: M. Sessé & J.M. Moçiño 2310 [holotype: MA!; isotypes: F, G, OXF!]
= Rollinia deliciosa SAFF., J. Wash. Acad. Sci. 6: 375, fig. 2 (1916)
Type: J.H. Simmonds s.n., 11 Mar. 1903 [syntype: US 8765973!]; J.H. Simmonds
s.n., 30 Aug. 1912 [syntype: US 8765976!]
= Rollinia jimenezii SAFF., J. Wash. Acad. Sci. 6: 378 (1916)
Type: A.M. Jiménez 427 [syntype: US!]; A.M. Jiménez 543 [syntype: US!]
= Rollinia permensis STANDL., Publ. Field Columbian Mus., Bot. Ser. 4 (8): 208 (1929)
Type: G.P. Cooper 645 [holotype: F; isotypes: G, K]
= Rollinia neglecta R.E.FR., Acta Horti Berg. 12 (1): 124 - 125 (1934)
≡ Rollinia mucosa var. neglecta (R.E.FR.) R.E.FR., Field Mus. Nat. Hist., Bot. Ser.
13 (2): 760 (1938)
Type: G. Klug 333 [holotype: US!; isotypes: F, NY!, S!]
= Rollinia curvipetala R.E.FR., Acta Horti Berg. 12 (1): 178 - 179, fig. 11a - c (1934)
Type: E. Ule 9371 [holotype: B!; isotypes: G, K!, MG, S!]
RAINER: Monographic studies in the genus Annona L. (Annonaceae): Inclusion of the genus Rollinia 203

= Rollinia mucosa subsp. aequatorialis R.E.FR., Acta Horti Berg. 12 (1): 123 - 124
(1934)
Type: R. Spruce 2403 [holotype: B!; isotypes: BM!, E, G, GH!, GOET, K!, M!,
OXF!, P!]
= Rollinia mucosa subsp. portoricensis R.E.FR., Acta Horti Berg. 12 (1): 124 (1934)
Type: P. Sintenis, Pl. portoricenses 4170 [holotype: S!; isotypes: B!, BM!, BP!, E, F,
G, GH!, GOET, HAC!, L!, LE!, M!, MO!, NY!, P!, S!, UC, WU!]
= Rollinia mucosa var. macropoda R.E.FR., Acta Horti Berg. 12 (1): 122 - 123 (1934)
Type: E.L. Ekman H12420 [holotype: S!; isotype: NY!]
= Rollinia jimenezii var. nelsonii R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl., ser.
3, 24 (10): 18 (1948)
Type: E.W. Nelson 406 [holotype: US!]

Annona sylvatica A.ST.-HIL., Pl. usuel. bras. 6: tab. 29 (1825)

≡ Rollinia sylvatica (A.ST.-HIL.) MART., Fl. bras. 13 (1): 18 (1841)
Type: A.F.C.P. de Saint-Hilaire s.n., s.d. [lectotype (selected by MAAS 1992: 148): P!;
isotype: P!]
= Rollinia fagifolia A.ST.-HIL., Fl. Bras. merid. 1 (1): 29 (1825)
Type: A.F.C.P. de Saint-Hilaire s.n., s.d. [holotype: P!; isotype: P!]
= Annona silvestris VELL., Fl. flumin. 239 (1829), Fl. flumin. icon. 5: tab. 128 (1831)
Type: tab. 128, in VELLOZO, Fl. flumin. icon. 5 (1831)
= Annona exalbida VELL., Fl. flumin. 240 (1829), Fl. flumin. icon. 5: tab. 131 (1831)
≡ Rollinia exalbida (VELL.) MART., Fl. bras. 13 (1): 19 (1841)
Type: tab. 131, in VELLOZO, Fl. flumin. icon. 5 (1831)
= Annona fagifolia A.ST.-HIL. & TUL., Ann. Sci. Nat., Bot., sér. 2, 17: 132 (1842)
Type: C. Gaudichaud s.n. (= Herbier Impérial du Brésil 636) [holotype: P!; isotype:
P (fragm.)!]

Annona xylopiifolia A.ST.-HIL. & TUL., Ann. Sci. Nat., Bot., sér. 2, 17: 132 (1842)
≡ Rollinia xylopiifolia (A.ST.-HIL. & TUL.) R.E.FR., Acta Horti Berg. 10 (2): 315
(1931)
Type: A. Richard s.n., s.d. [holotype: P!]
= Rollinia lanceolata R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl. 34 (5): 49 - 50,
tab. 6: fig. 6 (1900)
≡ Rollinia xylopiifolia var. lanceolata (R.E.FR.) R.E.FR., Acta Horti Berg. 12 (1):
176 - 177 (1934)
Type: A.F.M. Glaziou 13509 [lectotype (selected by FRIES 1934: 176): C; isolecto-
types: B!, BR, C, G, K!, LE!, P!]; A.F.M. Glaziou 16691 [syntypes: BR, C, F, LE!, P!]
= Rollinia warmingii R.E.FR., Kongl. Svenska Vetensk.-Akad. Handl. 34 (5): 49, tab.
6: fig. 7 (1900)
Type: A.F.M. Glaziou 6079 [holotype: B!; isotypes: C, K!]
204 Annalen des Naturhistorischen Museums in Wien 107 B

Acknowledgments
Special thanks go to Paul Maas and Lubbert Westra, Utrecht University, for continuous discussions on taxo-
nomic evaluation of generic concepts in Annonaceae, particularly concerning Annona, furthermore
Bernhard von Hagen, Bärbel Hildebrandt, and Martin Röser, Martin-Luther University Halle-Wittenberg,
for help with molecular analyses, thus providing the main trigger for this paper; to Walter Till, University
of Vienna, and to Bruno Wallnöfer, Natural History Museum Vienna, for critical comments on the manu-
script, as well as to the curators of the herbaria A, B, BM, BP, C, E, F, FHO, G, GH, GUAY, HAC, HAL,
HBR, JE, K, L, LE, LOJA, M, MA, MG, MO, NY, OXF, QCA, QCNE, P, PRC, RB, S, SP, U, UPS, US,
USM, W, WU for borrowing herbarium specimens or providing the facilities to study the specimens at their
institutions.

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 Virtual Herbaria Austria
http://herbarium.univie.ac.at/database/search.php

A common initiative of the herbaria

GJO Landesmuseum Joanneum Graz

GZU Karl-Franzes-University of Graz
W Naturhistorisches Museum Wien
WU Universität Wien, Faculty Center Botany (former Institute of Botany)

The Austrian collections are extremely rich of old and rare specimens. These
four institutions together hold probably more than 400.000 types. The databas-
ing was started few years ago and at the moment for c. 63.000 specimens the
data are available, of these c. 4.700 types, most of them with a scanned picture.

The database has standardised fields for taxa (genera / epithets / authors), col-
lectors, countries and provinces. It is possible to search for each of these fields
or combining them, or for continents. For geo-referenced specimens it is possi-
ble to get a map or satellit photo of the area. It is also possible to create a map
for a selected list of specimens.

The data are entered directly online and are stored for each instution separately.

Any instiution which wants to join this initiative, is welcome.

For testing the database you may use Briza media with as reasonable number of geo-
referenced specimens and the Bromeliaceae or Africa with many scanned types.

contact: heimo.rainer@univie.ac.at
ernst.vitek@nhm-wien.ac.at

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