Nomenclatural issues in the orchid bees - CiteSeerX

Zootaxa 3006: 1–42 (2011) www.mapress.com / zootaxa/ Copyright © 2011 · Magnolia Press

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ZOOTAXA ISSN 1175-5334 (online edition)

Nomenclatural issues in the orchid bees (Hymenoptera: Apidae: Euglossina) and an updated catalogue ANDRÉ NEMÉSIO1 & CLAUS RASMUSSEN2 1

Instituto de Biologia, Universidade Federal de Uberlândia. Rua Ceará, S/N, Campus Umuarama, Uberlândia, MG. 38.400-902. Brazil. E-mail: [email protected] 2 Department of Biological Sciences, Aarhus University, Ny Munkegade 114, Bldg. 1540, DK-8000 Aarhus C, Denmark. E-mail: [email protected]

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Taxonomic notes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 1. Eulaema meriana (Olivier, 1789) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 On the identity of Apis meriana Olivier, 1789 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Neotype designations and repositories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Apis meriana Olivier, 1789 and Apis dimidiata Fabricius, 1793 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 2. Euglossa piliventris Guérin-Méneville, 1844 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 3. Exaerete appendiculata (Romand, 1849), Exaerete subcornuta (Romand, 1849), and Euglossa dentata var. maxima Romand, 1849, nom. nud. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 On the true identity of Chrysantheda subcornuta Romand, 1849: natural history of orchid bees and the first record ever of their association with orchids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Orchid bees and orchids before Darwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 4. Eufriesea danielis (Schrottky, 1907) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 5. Euglossa fimbriata Moure, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 6. Euplusia yepezi Moure, 2000, nom. nud. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 7. Updated catalogue of orchid bees . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Genus Aglae Lepeletier de Saint Fargeau & Audinet-Serville, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Genus Eufriesea Cockerell, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Genus Euglossa Latreille, 1802 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Genus Eulaema Lepeletier de Saint Fargeau, 1841 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Genus Exaerete Hoffmannsegg, 1817. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Appendix 1. Valid species-group taxa of orchid bees (Euglossa and Eulaema) placed in subgenera . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Index of Latin names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

Abstract The taxonomic status (or the taxonomic status of the onomatophores) of seven nomina are revised: Eulaema meriana (Olivier, 1789), Euglossa piliventris Guérin-Méneville, 1844, Exaerete appendiculata (Romand, 1849), Exaerete subcornuta (Romand, 1849), Eufriesea danielis (Schrottky, 1907), Euglossa fimbriata Moure, 1968, and Eufriesea yepezi (Moure, 2000). Lectotype is designated for Euglossa piliventris Guérin-Méneville, 1844. Neotypes are designated for Apis meriana Olivier, 1789 and Apis dimidiata Fabricius, 1793 based on the same specimen and A. dimidiata Fabricius, 1793 became an objective synonym of A. meriana Olivier, 1789. Chrysantheda subcornuta Romand, 1849 is shown to be a senior objective synonym of Chrysantheda appendiculata Romand, 1849 and their synonymy under Exaerete dentata (Linnaeus, 1758) is questioned and discussed. Eumorpha combinata danielis Schrottky, 1907 is confirmed as a junior subjective synonym of Eufriesea auriceps (Friese, 1899), but the status of its onomatophore is reconsidered. Moreover, a long overlooked Euglossa dentata var. maxima Romand, 1849, nom. nud. and Euplusia yepezi Moure, 2000, nom. nud. are discussed for the first time. Both nomina Euglossa fimbriata Moure, 1968 and Euglossa fimbriata Rebêlo & Moure, 1996

Accepted by A.S. Lelej: 12 Jul. 2011; published: 29 Aug. 2011

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are available, but the latter is permamently invalid as a junior primary homonym of the former. An updated catalogue of all orchid bee nomina is also provided. Key words: euglossine bees, lectotype, neotype, nomenclature, systematics, taxonomy, Neotropical region

Introduction The discovery of the dependence of male orchid bees (Hymenoptera: Apidae: Euglossina; alternatively ranked as tribe, i.e., Euglossini, by some authors, e.g. Michener 2007) to chemical fragrances naturally found in flowers of many plant species, especially orchids (Vogel 1966), facilitated large collections of these insects from the 1960’s onward, when these fragrances were artificially synthesized and used as lures to attract the bees (Dodson et al. 1969). Many unknown species at that time were revealed to the scientific community (e.g. Moure 1967a, b, 1968, 1969, 1970; Dressler 1978a, b, 1982a, b, c). Much of the taxonomic knowledge on these bees prior to the utilization of fragrances to collect them was based on females. The ease of collecting males with the chemical fragrances completely changed our understanding of the diversity of orchid bees and it was soon realized that there were many more species, especially of Euglossa Latreille, 1802, than previously thought through the study of females. As females of many species are strikingly similar and often morphological near indistinguishable, the taxonomy of orchid bees was based on males from the 1970’s on, particularly because males display many external structures useful in taxonomic studies. Females were, thus, practically ignored in most taxonomic studies, especially those involving species of Euglossa (e.g. Moure 1968, 1970, Dressler 1978a, 1982a, b, c). Recently, however, it has been recognized that a few species had been described twice, i.e., once (before the extensive use of chemical attractants) based on female specimens and again (after the use of chemical attractants) based on male specimens (e.g. Bembé 2007, Nemésio 2009a: 92). The solution for these and many other problematic taxonomic complications has more recently concerned researchers and some recent attempts to deal with them have been carried out (e.g. Nemésio 2009a, b). Nevertheless, due to the complexity of some situations and the intricacies of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature 1999, hereafter referred to as the Code), and also due to a reinterpretation of the old literature (and the designations and/or indications there made), we concluded that the status of some species must be reassigned, as well as the status of some specimens listed as onomatophores (e.g. Nemésio 2009a). Below we discuss six particularly relevant cases, in addition to providing an updated catalogue of all available orchid bee nomina. The type repositories in the previously published checklists and catalogues (e.g. Moure 1967b; Kimsey & Dressler 1986; Ramírez et al. 2002; Roubik & Hanson 2004; Moure et al. 2007; Nemésio 2009a) have often relied on information from the original primary description, but we made an attempt to verify the whereabouts of all type specimens through personal visits or contact with curators of institutions as listed in Table 1. TABLE 1. Repositories and acronyms are cited following the Biodiversity Collections Index (including their permanent links). In parentheses follows the number of primary types of orchid bees in each institution, or in descending order for institutions with more than ten types: USNM, ZMBH, HNHM, DZUP, UFMG, UCDC, SEMC, and BMNH: AMNH USA, New York, New York, American Museum of Natural History (10) urn:lsid:biocol.org:col:32972; BMNH United Kingdom, London, Natural History Museum, Department of Entomology (10) urn:lsid:biocol.org:col:1009 CIBC Trinidad and Tobago, Curepe, International Institute of Biological Control (1) urn:lsid:biocol.org:col:33107 DZUP Brazil, Paraná, Curitiba, Universidade Federal do Paraná, Museu de Entomologia Pe. Jesus Santiago Moure (23) urn:lsid:biocol.org:col:33266 HNHM Hungary, Budapest, Hungarian Natural History Museum (28) urn:lsid:biocol.org:col:33453 IAVH Colombia, Boyacá, Villa de Leyva, Instituto Alexander von Humboldt (1) urn:lsid:biocol.org:col:1022 ICN Colombia, Bogotá, Universidad Nacional de Colombia, Insituto de Ciencias Naturales de la Universidad Nacional (2) urn:lsid:biocol.org:col:33478 INPA Brazil, Amazonas, Manaus, Instituto Nacional de Pesquisas da Amazoonia, Colecão Sistemática da Entomologia (1) urn:lsid:biocol.org:col:33531

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A. NEMÉSIO & C. RASMUSSEN

MACN MCZ

Argentina, Buenos Aires, Museo Argentina de Ciencias Naturales (1) urn:lsid:biocol.org:col:33736 USA, Massachusetts, Cambridge, Harvard University, Museum of Comparative Zoology (1) urn:lsid:biocol.org:col:33791 MHNAN France, Nantes, Muséum d’Histoire Naturelle de Nantes (4) urn:lsid:biocol.org:col:35082 MHNG Switzerland, Geneva, Muséum d'Histoire Naturelle (1) urn:lsid:biocol.org:col:33823 MIZA Venezuela, Aragua, Maracay, Universidad Central de Venezuela, Museo del Instituto de Zoología Agrícola (3) urn:lsid:biocol.org:col:33834 MNCE Brazil, Paraná, Curitiba, Museu de Historia Natural Capão da Embuia (1) urn:lsid:biocol.org:col:33854 MNHN France, Paris, Muséum National d'Histoire Naturelle (4) urn:lsid:biocol.org:col:33864 MNRJ Brazil, Rio de Janeiro, São Cristovão, Universidade do Rio Janeiro, Museu Nacional (3) urn:lsid:biocol.org:col:33870 MPEG Brazil, Pará, Belem, Museu Paraense Emilio Goeldi (5) urn:lsid:biocol.org:col:33880 MPSP Brazil, São Paulo, Universidade de São Paulo, Museu Paulista (3) urn:lsid:biocol.org:col:33888 MRSN Italy, Torino, Museo Regionale di Scienze Naturali (2) urn:lsid:biocol.org:col:33892 MSNG Italy, Genova, Museo Civico di Storia Naturale "Giacomo Doria" (6) urn:lsid:biocol.org:col:33900 MUCR Costa Rica, Ciudad Universitaria, Universidad de Costa Rica, Museo de Insectos (0) urn:lsid:biocol.org:col:33917 MUSM Peru, Lima, Universidad Nacional Mayor de San Marcos, Museo de Historia Natural (1) urn:lsid:biocol.org:col:33924 MZCR Costa Rica, Ciudad Universitaria, Universidad de Costa Rica, Museo de Zoologia (0) urn:lsid:biocol.org:col:33936 NHRS Sweden, Stockholm, Naturhistoriska riksmuseet (2) urn:lsid:biocol.org:col:33998 NMW Austria, Wien, Naturhistorisches Museum Wien (4) urn:lsid:biocol.org:col:34043 OUMNH United Kingdom, Oxford, University Museum of Natural History (2) urn:lsid:biocol.org:col:34102 QCAZ Ecuador, Quito, Pontificia Universidad Catolica del Ecuador, Catholic Zoology Museum (4) urn:lsid:biocol.org:col:34156 RPSP Brazil, São Paulo, Ribeirão Preto, Universidade de São Paulo (3) urn:lsid:biocol.org:col:35083 SEMC USA, Kansas, Lawrence, University of Kansas, Snow Entomological Museum (11) urn:lsid:biocol.org:col:34264 UCDC USA, California, Davis, University of California, R.M. Bohart Museum of Entomology (11) urn:lsid:biocol.org:col:34411 UF USA, Florida, Gainesville, University of Florida, Florida Museum of Natural History (2) urn:lsid:biocol.org:col:34858 UFMG Brazil, Minas Gerais, Belo Horizonte, Universidade Federal de Minas Gerais, Taxonomic Collections (21) urn:lsid:biocol.org:col:35084 UNAM Mexico, Mexico D.F., Universidad Nacional Autonoma de Mexico, Instituto de Biologia (1) urn:lsid:biocol.org:col:34485 USNM USA, Washington D.C., National Museum of Natural History (67) urn:lsid:biocol.org:col:1019 UUZM Sweden, Uppsala, Uppsala University, Evolutionsmuseet (3) urn:lsid:biocol.org:col:34534 WACA USA, Oregon, Ashland, Work amber collection (1) urn:lsid:biocol.org:col:34575 ZMHB Germany, Berlin, Museum für Naturkunde der Humboldt-Universität (33) urn:lsid:biocol.org:col:34628 ZMUC Denmark, Copenhagen, Natural History Museum of Denmark (2) urn:lsid:biocol.org:col:34644 ZSM Germany, Munich, Zoologische Staatssammlung (5) urn:lsid:biocol.org:col:34660 “Ettore Morone amber collection”, Turin, Italy (1)

A large number of nomina nuda found among orchid bees are briefly mentioned here, but not included in the checklist. The majority of these nomina nuda are a result of the delay in their description after an initial published reference to the nomen. Many of those nomina have since been made available by formal descriptions, but their original usage shall be regarded as nomina nuda. This includes the following publications and the nomina listed: Illiger (1806) for Centris infernalis (and possibly also the following nomina of orchid bees or Centridini, C. chrysura, C. mediata, C. pellicollis, C. ruficornis, C. scopipes, C. scutellaris, C. ursina, and C. verripes, Euglossa apicia and Eg. dives), Romand (1849) for Euglossa dentata maxima, Friese (1899) for Euglossa xanthogastra, Moure (1967b) for the following nomina of Euglossa: Eg. alleni, Eg. allosticta, Eg. asarophora, Eg. augaspis, Eg. bursig-

NOMENCLATURAL ISSUES IN THE ORCHID BEES

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era, Eg. coruscans, Eg. crassipunctata, Eg. cupreicolor, Eg. cyanaspis, Eg. cyanosoma, Eg. cybelia, Eg. deceptrix, Eg. dressleri, Eg. erythrochlora, Eg. flammea, Eg. heterosticta, Eg. iopyrrha, Eg. jamaicensis, Eg. macropsectra, Eg. maculilabris, Eg. melanotricha, Eg. nigrosignata, Eg. orichalcea, Eg. perfulgens, Eg. platymera, Eg. pleosticta, Eg. rugilabris, Eg. sapphirina, Eg. spinigaleis, Eg. stilbonota, Eg. tridentata, Eg. villosa, Eg. villosiventris; and of Eulaema: El. basicincta, El. bennetti, El. luteola, El. mimetica; Casolari & Casolari (1979) for Euglossa albilabris and Eg. semipurpurata, Aglae coerulea virescens, and Centris splendida and C. similis (Spinola, nec Fabricius), and Moure (2000) for Eufriesea yepezi (see below). Nomina proposed by Pérez in Dominique (1898) were long considered nomina nuda, but as discussed and clarified elsewhere, they are in fact available (Michener 1987; Rasmussen et al. 2007). During the period comprising the XVIII through the XX century, we did not see evidence from the original publication that the description was based on a single specimen, despite the fact that unique holotypes are listed in several of the previous catalogues. We also notice that none of the collections of the authors of these new taxa are completely united in a single museum, and we therefore prefer to treat them as syntypes (Recommendation 73F of the Code) as additional specimens may have existed or still exist in European museums: Fabricius (Apis dimidiata, A. mussitans); Friese (eight species described as Euglossa: Eufriesea chalybaea, Ef. excellens, Eg. igniventris, Eg. lazulina, Ef. longipennis, El. mocsaryi, El. nigrifacies, El. sarapiquiensis); Lepeletier de Saint Fargeau (Euglossa brullei and Eulaema analis, Ef. coerulescens, El. cajennenis, El. elegans); Mocsáry (three species described as Euglossa: Ef. formosa, Ef. opulenta, Ef. theresiae). As indicated, all of these taxa have been inferred as “holotypes” in publications before 2000, and through article 74.6 of the Code, the actual specimens previously regarded as holotypes, were fixed as lectotypes when first cited as such holotypes, in particular by Moure (1967b), and since followed by others [up until the new Code in 1999]. The only taxa not fixed by that action were Apis dimidiata and Eulaema analis, as no specimens were located in any collection, and despite a reference to a holotype, it is not certain which specimen that would have been. Country of origin for the type specimens may only be a reflection of where taxonomists did their field work, but we notice that only Brazil and Panama – out of a total of 19 type countries – are represented with more than 20 new taxa. The countries and the number of new species described from each are: Argentina (1), Bolivia (13), Brazil (94), Colombia (16), Costa Rica (15), Dominican Republic (2 fossils), Ecuador (20), French Guiana (19), Guatemala (1), Guyana (6), Jamaica (2), Mexico (13), Nicaragua (1), Panama (38), Paraguay (3), Peru (20), Suriname (6), Trinidad & Tobago (5), Venezuela (17), and six of unknown origin. Studying material from all of these countries, the most prolific describers of new species-group and genus-group taxa were the following, each with 20 or more proposed nomina: Moure (60), Friese (53), Dressler (42), and Mocsáry (20). As for collectors of specimens that became the onomatophores of new taxa, Dressler caught 48 of those in his extensive orchid and baiting studies. Only others with significant collections were Dodson, Ducke, and Roubik with six each. Although descriptions have been continuous since 1758, the few peaks years for new descriptions were 1982 (31), 1899 (24), 1968 (16), and 1898 (15) (Fig. 1).

FIGURE 1. Rate of first formal descriptions of new orchid bee taxa by decade. Dates are from the oldest available nomina, 1758, in the sense of the Code, and until 2009.



The checklist is sorted alphabetically first by genus-group nomen and then by species-group nomen. Following the taxon nomen we list the original combination or spelling when different from the currently valid combination. Taxa originally proposed as new subspecies including varieties (n. var.) and forms (n. form) deemed subspecific (Article 45.6.4. of the Code) are indicated by listing the specific nomen followed by “ssp.” (subspecies). We do not recognize any valid subspecific taxa in this checklist. Following the nomen of the taxon, we provide the authority, year of publication, and reference, including page numbers in the reference and reference to illustrations in the original publication. We have here added references (including figure number) to any subsequent illustration of actual type specimens. Following this are status of type (holotype, syntype, lectotype, neotype), gender of type specimen, reference for subsequent lecto- or neotype designation, and type repository including museum code if provided (Table 1 for list of acronyms). Following this are listed type country (in capitals), state, and locality with spelling corrected. The collector of the type specimen is included when known. Following this we add any comments that may be to the entry, including observations on variant spelling, information different from the original publication, or taxonomic observations. The genera Euglossa and Eulaema have often been divided into subgenera, and these are included last (see also Appendix 1).

Taxonomic notes In the present work, we strictly respect the rules of the Code, but some different terminologies were used to designate the concepts therein, for reasons explained in detail by Dubois (2000, 2005). We use the term “nomen” (plural “nomina”) for “scientific name” and the term “onomatophore” (Simpson 1940) to replace the expression “namebearing type” found in the Code.

1. Eulaema meriana (Olivier, 1789) (Figs 4A–4D) Apis meriana Olivier, 1789. Apis dimidiata Fabricius, 1793.

The original description of Apis meriana by Olivier (1789: 64) is as follows: “Abeille Mérian Apis Meriana. Nob. Apis hirsuta, nigra, abdomine segmentorum marginibus pallidè flavis; ano rufo. Nob. Merian. Surin. pl. 48. Cette abeille est une des plus grandes que nous connoissions. Ses antennes e sa tête sont noires. Les yeux sont bruns, e la trompe est plus longue que la moitié du corps. Le corcelet est noir e velu. L’abdomen est noir, avec le bord des quatre premiers anneaux d’un jaune pâle, e l’anus fauve. Les pattes sont noires, e los jambes postérieures sont très grosses. Les aîles supérieures sont noires, depuis la base jusque vers leur milieu; le reste est transparent. Les aîles inférieures sont obscures; leur pointe seulement est transparent. Cette abeille se trouve à Cayenne e à Surinam: elle m’a été communiquée par M. Renaud, docteur en Médicine.”

The textual description refers to the bee currently known as Eulaema meriana or to one very similar Eulaema (Eulaema), with metasoma clothed in black and yellow setae on the first terga and in red setae on the last terga (“ano rufo”). Based on this description and on Fabricius’s (1793) description of Apis dimidiata, Moure (1960b) reached the conclusion that both species were the same, and established the synonymy of the latter under the former. The nomen Apis meriana was largely ignored for more than 170 years; the nomen Eulaema dimidiata being the only one used to refer to the bee we now call Eulaema meriana until 1960, when Moure introduced the synonymy. Nevertheless, Moure (1960b) gave no reason for establishing this synonymy. He stated that “probably the type specimen of Olivier was destroyed” (Moure 1960b: 146). As Nemésio (2009a: 163) noticed, J.S. Moure was ambivalent concerning the nature of the synonymy between El. meriana and El. dimidiata, because in his first work (Moure 1960b), he gave the impression that there were two onomatophores, one for each species (which would result in a subjective synonymy), but in his next work on this subject (Moure 2003: 34), he considered that



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Olivier’s (1789) and Fabricius’s (1793) descriptions were based on the same specimen (a primary objective synonymy). In his latter work, Moure insisted that the specimen was lost (Moure 2003). Here, we intend to show both nomina were erected based on different specimens and that the nomen Apis meriana would partially refer, in fact, to the bee currently known as Eulaema polyzona (Mocsáry, 1897) (Fig. 3). Maria Sybilla Merian made several records and illustrations on insects from Suriname and published a detailed report of her findings in 1705. On her plate 48 (here reproduced as Fig. 2) she illustrated two Coleoptera, two lepidopteran caterpillars, and one bee. This bee was specifically and explicitly indicated by Olivier (1789) as one of the specimens upon which he based his description of Apis meriana (“Abeille Mérian”, honoring M. S. Merian), even though the textual description does not entirely match the illustration. The first point to be made clear is that this indication, under the modern Code, is a valid one (if made before 1930, as happens to be the case) and could explicitly establish the onomatophore of the species. As there is a single bee illustrated [although it could be argued that Merian (1705: 48) interpreted one of the caterpillars as the larval stage of the bee—see below, Olivier (1789) was clearly referring to the bee] this specimen must be interpreted as one of the “type specimens”. Article 73.1.4 of the Code clearly apply to this situation, stating that the “designation of an illustration of a single specimen as a holotype is to be treated as designation of the specimen illustrated; the fact that the specimen no longer exists or cannot be traced does not of itself invalidate the designation” (our bold). In addition, it is also clear from Olivier’s text that he saw at least one specimen of the bee we currently treat as Eulaema meriana (or a similar species), and probably this specimen was brought to him by Mr. Renaud. Olivier (1789) did not explicitly write it, but there was no type concept at that time and we argue here that this specimen should be considered as part of the type series, which would result (together with Merian’s plate) in a syntypic series. Moreover, it is uncertain if Moure (2003) was correct, speculating that Olivier (1789) and Fabricius (1793) based their description on the same specimen (although Olivier helped Fabricius with specimens; see Hope 1845). Olivier’s type specimens are the bee illustrated by Merian (1705) and the specimen brought to him by Renaud (now lost), whereas Fabricius’s type specimen is a bee originally deposited at Bosc’s Collection [Renaud’s specimens are apparently not known to be in the Bosc Collection], as explicitly indicated by Fabricius (1793: 316). This latter bee is considered lost since Moure (1960b) and, recently, a search for this specimen carried out by the curator of the Hymenopteran collection of the Muséum National d’Histoire Naturelle, Paris, where Bosc’s Collection is deposited, has been unsuccessful (Nemésio 2009a: 163). Therefore, we reiterate the remark of this specimen presently being lost.

On the identity of Apis meriana Olivier, 1789 In order for us to make any taxonomic decision, attention should be focused on the identity of the specimen illustrated by Merian (1705). The first noticeable difference is, of course, coloration. The description by Olivier (1789) suggests a bee with red setae on the last terga, whereas the drawing by Merian (1705) reveals a bee with a metasoma entirely banded in black and yellow setae only. Although some specimens of the bee currently treated as Eulaema meriana can present yellowish (usually fulvous) setae on the last terga, there is no record of such specimens in the Guianas (Dressler 1979). General shape of the bee illustrated by Merian (1705) also suggests it is not the current El. meriana. But is the yellow/black coloration of Merian’s plate an actual record of the bee or could it be an artifact? One could argue that plates were printed in black and white and colored manually after printing, as happened to be the case in early XVIII century (see Zaharek & Overstreet 2001 for a discussion of XVIII century scientific illustration). Thus, different copies of the book could present the same bee colored differently. The few copies we have seen (two) portrait identical yellow and black bees. A possibly different source is the book seen by Dressler (1979). According to him, “though Olivier believed that he was treating the bee illustrated by Maria Sybilla Merian (1705), Merian’s plate shows a black bee with yellow bands and no red or brown on the terga” (Dressler 1979: 149). More important and decisive, however, is the statement made by Merian (1705: 48) herself (free translation from Dutch): “The caterpillar, which crawls on the fruit, is yellow (and) black, and resembles a clothesbroom, and eats these leaves. Around August 3rd it made a cocoon in which it became, on August 15th, a big black bee with black and yellow bands”. Although clearly making the mistake of considering the caterpillar as a larval stage of the bee, Merian explicitly described the bee as having only black and yellow bands. There is no mention to red. These evidences are strong enough to consider that the specimen actually was not the bee we now call El. meriana.



FIGURE 2. Original illustration from Merian (1705), with several insects found moving around the Genipa americana (Rubiaceae), including supposedly the enigmatic Apis meriana Olivier.



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FIGURE 3. Eulaema polyzona (Mocsáry, 1897), the likely candidate of Merian’s (1705) actual illustration (see Fig. 2). A. Dorsal view, B. Ventral view.

Some Eulaema species, on the other hand, present a pattern that exactly matches Merian’s illustration. By eliminating candidates due to details of the coloration pattern and geographic distribution, we had two alternatives: El. (Apeulaema) boliviensis Friese, 1898 and El. (Eulaema) polyzona (Mocsáry, 1897). The bee illustrated by Merian (1705) seems to be a male, due to the absence of clear corbiculae, if we consider her drawing accurate in this respect. In spite of being a dorsal view, the face of the bee is somewhat inclined, in such a way it is possible to see an entirely black frons and upper clypeus. There are no white or yellowish markings at all, a character found on males of Eulaema (Apeulaema), except some specimens of El. mocsaryi (Friese, 1899) and El. felipei Nemésio, 2010. If Merian’s representation of this character is reliable, it could not be El. boliviensis. More important, there is no record of El. boliviensis from the Guianas (Dressler 1979: 149; Ramírez et al. 2002; Bart De Dijn, pers. comm.). On the other hand, the drawing exactly matches El. polyzona, and this species has been recorded for the Guianas (Dressler 1979: 151; Ramírez et al. 2002; Nemésio & Silveira 2007; Bart De Dijn, pers. comm.). We conclude that the bee illustrated by Merian (1705) is the species currently known as El. polyzona and if the bee illustrated by Merian is selected as the lectotype of Apis meriana, it will result in the synonymy of El. polyzona under Apis meriana. This would be a difficult decision, since the nomen El. meriana has been frequently used (prevailing usage, according to the Code) – although ignoring it clearly means that a wrong decision by Moure (1960b, 2003) will prevail over Fabricius’s (1793) correct recognition of a new species at the time. The situation here presented is a typical example of a dilemma between strict priority and stability (prevailing usage). Selecting the bee illustrated by Merian (1705) as the lectotype of Apis meriana would result in a change in orchid-bee nomenclature that would not be desirable. Firstly, because Eulaema meriana is a common and widespread species in the Amazon and Central America and a widely accepted nomen in the literature; secondly, because the nomen A. meriana was never applied to the bee later described as El. polyzona; finally, because this decision would result in the onomatophore of A. meriana being not in taxonomic accord with the prevailing usage of the nomen (Article 75.6 of the Code), what would result in the express rule by the Code to ask the Commission to set it aside and designate a neotype which is in taxonomic accord with the prevailing usage of the nomen. Fortunately, Olivier (1789) referred to Renaud’s specimen and his textual description allows us to choose this specimen as the lectotype of A. meriana, in order to maintain stability of the usage of the nomen El. meriana. This action is taken here and the specimen collected by Renaud and mentioned by Olivier (1789) is designated as the lectotype of A. meriana. Nevertheless, once this specimen is lost, another source of problems is introduced. The description of Apis meriana by Olivier (1789) does not exclude the possibility that the bee brought by Renaud belongs to a similar but different species—as Eulaema bombiformis (Packard, 1869), for example. If this specimen is found in the future and detailed studies confirm that it belongs to another species, the action taken here will have been worthless, since the nomen A. meriana would be applied to other species than the bee currently treated as El. meriana. The nomen El. dimidiata (Fabricius, 1793) would have to be resurrected. It must be emphasized that it is not possible to declare El. dimidiata as a nomen oblitum to favor El. meriana under the provisions of the Code (Article 23.9), since the nomen El. dimidiata was used many times as valid after 1899.



Thus, to avoid present and future confusion, and because the onomatophores of both Apis meriana Olivier, 1789 and A. dimidiata Fabricius, 1793 are lost, besides selecting Renaud’s specimen as the lectotype of A. meriana (to avoid the use of the plate by Merian to be used, since the syntypic series comprised two species, as shown above) we here designate neotypes for both species. More important, we select the same specimen to act as the neotype of both species, creating an objective synonymy between A. meriana and A. dimidiata, which would be in accord with Moure’s (1960b) action and also in accord with the current taxonomic treatment of both nomina.

Neotype designations and repositories Apis meriana Olivier, 1789: NEOTYPE ♂, with the following label data: “EIA Porto Trombetas, Teófilo 2, Zona Leste, 11517-34171”, “Oriximiná, PA, Brasil, 10/12/2006, R. B. Martines”, “Apis meriana Olivier, 1789, NEOTYPUS” and “Apis dimidiata Fabricius, 1793, NEOTYPUS”, deposited at the Entomological Collection of the Taxonomic Collections of the Universidade Federal de Minas Gerais, UFMG, Belo Horizonte, Brazil. Apis dimidiata Fabricius, 1793: NEOTYPE ♂, with the following label data: “EIA Porto Trombetas, Teófilo 2, Zona Leste, 11517-34171”, “Oriximiná, PA, Brasil, 10/12/2006, R. B. Martines”, “Apis meriana Olivier, 1789, NEOTYPUS” and “Apis dimidiata Fabricius, 1793, NEOTYPUS”, deposited at the Entomological Collection of the Taxonomic Collections of the Universidade Federal de Minas Gerais, UFMG, Belo Horizonte, Brazil. The above designations met all qualifying conditions of Article 75.3 of the Code, as follows: Art. 75.3.1—“a statement that it is designated with the express purpose of clarifying the taxonomic status (…) of a nominal taxon”. Satisfied previously in this text; Art. 75.3.2—“a statement of the characters that the author regards as differentiating from other taxa the nominal species-group taxon for which the neotype is designated (…)”. See descriptions below; Art. 75.3.3—“data and description sufficient to ensure recognition of the specimen designated”. See label data and repositories above, and descriptions below; Art. 75.3.4—“the author’s reasons for believing the name-bearing type specimen(s) (…) to be lost or destroyed, and the steps that had been taken to trace it or them”. See above and also Nemésio (2009a). Art. 75.3.5—“evidence that the neotype is consistent with what is known of the former name-bearing type from the original description and from other sources; however, a neotype may be based on a different sex (…), if necessary or desirable to secure stability of nomenclature”. The neotype Apis meriana is entirely consistent with the description by Olivier (1789) and the current taxonomic use of the nomen, as discussed above; the neotype Apis dimidiata is entirely consistent with the description by Fabricius (1793) and with a specimen identified as such by Fabricius himself (see Nemésio 2009a: 163); Art. 75.3.6—“evidence that the neotype came as nearly as practicable from the original type locality (…)”. Both species were described from Suriname; the specimen selected as neotypes here come from the quite close state of Pará, Brazil, which shares its borders with Suriname. According to the Code (Article 76.3), “the place of origin of the neotype becomes the type locality of the nominal species-group taxon, despite any previously published statement of the type locality”. Thus, Oriximiná (state of Pará, Brazil) becomes the type locality of both Apis meriana Olivier, 1789 and Apis dimidiata Fabricius, 1793. Art. 75.3.7—“a statement that the neotype is (…) property of a recognized scientific or educational institution, cited by name, that maintains a research collection, with proper facilities for preserving name-bearing types, and that makes them accessible for study”. See repositories above.

Apis meriana Olivier, 1789 and Apis dimidiata Fabricius, 1793 Description of the NEOTYPE male (same specimen for both species) (Fig. 4): Measurements: total length ca. 26.0 mm; head width ca. 7.3 mm; eye length ca. 5.0 mm; scutellum width ca. 5.5 mm; scutellum length ca. 3.2 mm; abdominal width ca. 11.0 mm. Color and vestiture. Face and head entirely black (Fig. 4C), mesosoma dark brown with black hairs, T1–T4 dark with greenish metallic hues, first 2/3 of its length clothed in black setae and



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distal 1/3 clothed in yellow setae; T5–T7 brown with predominantly long reddish setae (Fig. 4A); fifth sternum with very sparse setae (Fig. 4B). Legs. Velvet area of mesotibia not reaching the distal portion, leaving only a broad black area around; mesotibial tuft small, occupying the basal first quarter of the velvet area (Fig. 4D); metatibia covered with very sparse black hairs. Diagnosis. Eulaema meriana is similar to the sympatric El. bombiformis (Packard, 1869). Nevertheless, they can easily be distinguished by the presence of dense setae on the fifth sternum of El. bombiformis, leaving only a glabrous median longitudinal line (only sparse and short setae on the fifth sternum in El. meriana).

FIGURE 4. Eulaema meriana (Olivier, 1789) and El. dimidiata (Fabricius, 1793) neotype (same specimen). A. Dorsal view, B. Ventral view, C. Head, frontal view, D. Mesotibia.

2. Euglossa piliventris Guérin-Méneville, 1844 (Figs 5, 6) Euglossa piliventris was described based on male and female specimens from the state of Pará, Brazil, in the eastern portion of the Brazilian Amazon (Guérin-Méneville 1844). Later, Friese (1899: 136–137) mentioned he saw type specimens in “coll. Saussure” (now in Naturalis, Leiden, Netherlands), in addition to (non-type) specimens from Colombia (Bogotá, Muzo), Brazil, Suriname, Cayenne (in French Guiana) and Peru in collections he visited (Rasmussen & Ascher 2008). Moure (1967b) mentioned “type male and female in coll. Saussure”, attributing this information to Friese (1899). Moure (1967b) also mentioned “a paratype female seen by the author at the Museo Civico di Storia Naturale (MSNG), Genova” and considered the geographic distribution of this species as Brazil (states of Amapá, Amazonas and Pará), “Guianas” and Peru. Roubik (2004) recognized two forms among the bees usually treated as Eg. piliventris and considered the female specimen from Pará deposited at MSNG as the “holo-



type” of Eg. piliventris. With the available data, Roubik (2004: 238, 244–246) considered Eg. lugubris Roubik, 2004 as restricted to western Amazon and Eg. piliventris as restricted to the eastern Amazon. Moure et al. (2007) listed only female onomatophores deposited at the ‘Museo Civico di Storia Naturale’ and at the ‘Naturalis’ of Eg. piliventris (as syntypes). Males were apparently not traceable. Finally, Nemésio (2009a: 238, footnote 81) recognized Roubik’s (2004) designation of a lectotype stating that: “Moure (1967b: 405) listed syntypes (male and female) ‘in coll. Saussure’ and a ‘paratype’ at the ‘Museo Civico di Storia Naturale’, Genoa. As a member of the type series, this ‘paratype’ mentioned by Moure was eligible to be designated as a lectotype [it is from Pará, as in the original description by Guérin-Méneville (1844: 458)]. Moure et al. (2007: 240) treated the type material as a syntypic series, but I here recognize Roubik’s (2004: 246) action of selecting this female deposited in Genoa as lectotype as valid (in spite of writing ‘holotype’ instead of ‘lectotype’). Male and female specimens deposited at the ‘Nationaal Natuurhistorisch Museum’, Leiden, the Netherlands, become, thus, paralectotypes”. Although it is correct that the specimen deposited at MSNG is eligible to be designated as a lectotype, Roubik’s (2004) action cannot be validated under the Code, since Article 74.7 of the Code states that lectotype designations after 1999, to be valid, must employ the term “lectotype” or an exact translation (e.g., “lectotypus”, but not “the type”) and must contain an express statement of the taxonomic purpose of the designation, two criteria not met by Roubik (2004): he used “holotype” instead of “lectotype” or an exact translation and his designation contained no express statement of its taxonomic purpose (although it is implicit from his entire work, in which a similar species was described, that fixing the identity of the species Eg. piliventris was essential). To avoid any misinterpretations in the future, and since Roubik (2004) clearly established the identity of Euglossa piliventris to avoid confusion with the similar Eg. lugubris described in that paper, in the aim of nomenclatural stability, we designate in this work the female Eg. piliventris from Brazil, state of Pará, deposited at the Museo Civico di Storia Naturale, Genoa, Italy, as the lectotype of the Eg. piliventris Guérin-Méneville, 1844. The specimens deposited at the Naturalis, Leiden, the Netherlands, become, thus, paralectotypes. A photograph of the lectotype is here provided (Fig. 5), as well as of its labels (Fig. 6).

FIGURE 5. Euglossa piliventris Guérin-Méneville, 1844 lectotype, habitus.



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FIGURE 6. Euglossa piliventris Guérin-Méneville, 1844 lectotype, labels.

3. Exaerete appendiculata (Romand, 1849), Exaerete subcornuta (Romand, 1849), and Euglossa dentata var. maxima Romand, 1849, nom. nud. Chrysantheda appendiculata Romand, 1849, Chrysantheda subcornuta Romand, 1849, and Euglossa dentata var. maxima Romand, 1849, nom. nud.

When describing a supposed new species of Exaerete (as Chrysantheda), Bernard Romand (1849) inadvertently published three nomina for this same species, one of them long overlooked and here interpreted as a nomen nudum. Although it is well accepted in the literature that Chrysantheda appendiculata and Ch. subcornuta are junior synonyms of Exaerete dentata (Linnaeus, 1758) (see Moure 1967b; Kimsey 1979; Kimsey & Dressler 1986; Roubik & Hanson 2004; Moure et al. 2007; Nemésio 2009a), the status of these two species are apparently not well understood and the order of priority (among the junior synonyms) of both has been also usually mistaken. Although the onomatophore of Ch. subcornuta has been treated as a male since Moure (1967b), the gender of the onomatophore of Ch. appendiculata has been given as “unknown” by all authors. Misunderstandings concerning these little known nomina have their origin in the confusing description made by Romand (1849), but an accurate study of his work can elucidate the case. Also, although we kept them as junior synonyms of Exaerete dentata, there is reasonable evidence suggesting that these nomina can belong to other species of Exaerete. Until the eighteenth and mid-nineteenth centuries illustrated publications often had the plates sent to the printer in advance, in such a way that plates were printed before the text referring to them, and the two parts had to be attached at a later stage (see the example of Buffon’s classical works in Zaharek & Overstreet 2001). It could even happen (and did happen more than once) that plates were released before their texts, allowing some authors to name animals solely indicating the illustrations, creating confusion sometimes (see one of these situations in Nemésio & Rasmussen 2009). Information contained in the text was therefore often more current than that presented on



the plates, because late changes could be done in the text but not on the already printed plates. This is exactly what happened in Romand (1849). When preparing his work, Romand described as Chrysantheda appendiculata what he thought to be an undescribed species. This is the nomen printed in the text and on the captions of the Plate 7, species number 1, in the Annales de la Société Entomologique de France, [2]7. Nevertheless, in the period between the printing of the plates and the printing of the text, Romand received a letter from Marchese Massimiliano Spinola, who had also recognized a supposed new species of Chrysantheda and provided the details to Romand. Romand, in turn, sent a letter to the publishers of his work letting them know that he would like to add the information provided by Spinola. The publishers inserted a footnote on pages xxxvii–xxxviii of Romand’s (1849) work, informing that (free translation from French): “Since this note was printed, the Société received, in its meeting of 12th of September of 1849, a letter from Mr. de Romand containing new details about the Chrysantheda he mentioned [Romand, 1849: p. xxxvi–xxxvii]. We reproduce this letter here: ‘I did research on the genus Chrysantheda and I recognize it was established by Mr. Max. Perty in the Delectus Animalium articulatorium, 1830, which I have seen.’ ‘Perty described, after the genus, one species under the name Ch. nitida, which is a female. The only species [or specimen] he has seen.’ ‘Mr. Max. Spinola communicated to me in time the designation of one species under the name Ch. subcornuta, which he considered as the Euglossa dentata of Fabricius, var. maxima, since Chrysantheda is a subdivision of Euglossa, from which it differs a little.’ ‘Thus I am willing to name this species, which drawing I have already sent and that is extremely similar to that informed by Spinola, as Ch. subcornuta Spinola, and I see no need to add a new description because one can find the characters in Fabricius, the notes by Mr. Max. Spinola, and Perty, which terms do apply, except for size, to the species here considered’.” (our bold)

It is important to notice that the letter was sent to the publishers. It seems that Romand’s intention was to have the nomen of the species changed to Chrysantheda subcornuta both in the text and on the plate, since he was kind enough to give priority to the nomen proposed by Spinola. Nevertheless, it also seems that it was too late to carry out such modifications and the publishers opted instead to insert the letter by Romand, ipsis litteris, as footnotes. One hundred and sixty years after the publication of Romand’s work, we ought to apply the rules of the current Code to interpret which nomenclatural acts are valid in Romand’s (1849) text. Although it is obvious that the footnote was inserted in the work by the publishers, the authorship of the text contained in the letter is also obviously attributed to Romand, in such a way he should naturally be considered the author of the letter text also. Romand’s acts, as interpreted currently, were as follows: 1). Romand explicitly refrained from publishing the new species under the nomen Chrysantheda appendiculata and considered the nomen Ch. subcornuta as the nomen of the species. As both nomina were printed (Ch. appendiculata in the main text and on the plate, because the publishers did not change it), Romand’s preference should be explicitly interpreted as giving priority to Ch. subcornuta, in such a way Ch. appendiculata was originally published as a junior synonym of Ch. subcornuta. Romand (1849: xxxvii) even stated that the nomen Ch. appendiculata was used as an “error” on plate 7. Two points deserve attention here: (i) the nomen Ch. subcornuta has priority over Ch. appendiculata and should be cited before the latter nomen in the list of synonyms of Exaerete dentata. If, in the future, the onomatophore of this species is found and it is proved that it is another species and one of these two nomina is the oldest available nomen for this other species, the nomen to be adopted must be Ch. subcornuta. Modern authors (started by Kimsey 1979) have listed Ch. appendiculata over Ch. subcornuta, probably because this nomen appears first in Romand’s text. As Nemésio (2007a) and Dubois et al. (2011) have already pointed out, there is no page (or line) priority in the Code for such cases. In this situation a First Reviser action is unnecessary, because it is clear that Romand treated Ch. appendiculata as the junior synonym; (ii) Ch. subcornuta was also explicitly considered a junior synonym of both Ch. nitida Perty, 1833 (date erroneously listed as 1830 by Romand) and Euglossa dentata Fabricius, var. maxima, because Romand saw “no need to add a new description because one could find the characters in Fabricius, the notes of Spinola, and Perty, which terms do apply, except for size, to the species here considered”. If Ch. subcornuta was described as a junior synonym, then Article 11.6.1 of the Code applies, and authorship should be given to Romand, not to Spinola – ironically, in spite of Romand’s gen-



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tle attitude of giving priority and even listing Ch. subcornuta as a Spinola species (see examples of application of Article 11.6.1 in the Code itself and in Nemésio 2006b). 2). If Chrysantheda subcornuta and Ch. appendiculata were published based on the same specimen (in fact, it was all a matter of having the two nomina “accidentally” printed for the same description, as shown above), then the onomatophore of both nomina should obviously be the same (an objective synonymy). Thus, there is no sense in listing the onomatophore of Ch. subcornuta as a male and the onomatophore of Ch. appendiculata as undetermined sex, as all authors did since the introduction of this error by Kimsey (1979) [Kimsey & Dressler (1986) omitted Ch. appendiculata from their checklist, possibly considering it a nomen nudum?]. The onomatophore of Ch. appendiculata is clearly a male. 3). Since the nomen Euglossa dentata Fabricius, var. maxima (authorship of Ex. dentata erroneously attributed to Fabricius) was only mentioned as an alleged communication from Spinola to Romand and no specific character was mentioned for this nomen, it could not be considered available under the Article 13.1.1 of the Code and it is best treated as a nomen nudum, the interpretation adopted in this work. 4). As a consequence, if future authors consider that both nomina should be considered as junior synonyms of Exaerete dentata (Linnaeus, 1758), the synonymic list of Ex. dentata, on the order of priority, should be as:

Exaerete dentata (Linnaeus, 1758) Apis dentata Linnaeus, 1758. Chrysantheda nitida Perty, 1833. Chrysantheda subcornuta Romand, 1849. Chrysantheda appendiculata Romand, 1849.

On the other hand, we present below some reasons to question the above synonymy of Chrysantheda subcornuta and Ch. appendiculata under Exaerete dentata. On the true identity of Chrysantheda subcornuta Romand, 1849: natural history of orchid bees and the first record ever of their association with orchids Two important features presented in the “description” of Chrysantheda subcornuta (from now on, both nomina Chrysantheda subcornuta and Ch. appendiculata are treated as Ch. subcornuta, the senior objective synonym of them, as shown above) are the size of this species (which may provide indication of the identity) and the nature of the “projection” on the head of the male specimen of Ch. subcornuta described by Romand (1849) that attracted the attention of the Société Entomologique de France (hereafter treated as the Société), as will be shown below. When refraining from presenting a more detailed morphological description of Ch. subcornuta, Romand (1849: xxxviii, footnote) argued that the descriptions contained in Fabricius (for Euglossa dentata var. maxima), in the letter of Spinola, and in Perty (for Ch. nitida) were applicable to Ch. subcornuta, except for size. Romand (1849) did not explicitly state, however, if the difference in size meant that Ch. subcornuta was larger or smaller than described by the mentioned authors. But it can be inferred from his text. Fabricius (1787, 1793, 1804) has never described an orchid bee as “var. maxima”, in such a way that the introduction of such a nomen is here entirely attributed to Spinola and reproduced by Romand in his letter to the Société. Fabricius (1804: 363) did present a diagnosis of Exaerete dentata (Linnaeus, 1758) (under the nomen Euglossa dentata), but presented no measurement for this species (nor did Linnaeus 1758, 1767). We do not know what Spinola communicated to Romand, but Perty (1833: 148), on the other hand, presented objective measurements for Ch. nitida (Lg. 9’’’, Latit. alar. expans. 17’’’, assuming French lines, this corresponds to a length of 20 mm and a wingspan of 38 mm). We can infer the size of Ch. subcornuta from the following: (i) in the main text, when explaining the Fig. 1A of the Pl. 7, showing the habitum of the specimen, it is written “l’insecte parfait de grandeur naturelle” (in a free translation: “insect reproduced in natural size”). From the original plate we record the following measures: body length: 20.6 mm; head width: 5.3 mm; mesoscutum width: 5.8 mm; abdominal width: 7.0 mm; forewing length: 12.6 mm; hind tibia length: 7.5 mm; (ii) the use of the epithet “maxima” by Spinola to designate this bee, in allusion to Exaerete dentata, also supports the idea of a larger bee, since “maxima” in Latin means, when referring to size, the superlative of big, “the maximum size”.



Although size variation in orchid bees has not been uncommonly reported (e.g. Peruquetti 2003, Roubik 2004), and a noticeable size variation has been particularly observed in parasitic species, especially Aglae coerulea Lepeletier de Saint Fargeau & Audinet-Serville, 1825 (AN, pers. obs.), a disparate difference in size between Exaerete dentata (the smallest Exaerete, between 15–18 mm, (Kimsey 1979, Nemésio 2009a) and Ch. subcornuta (about 21 mm) is remarkable, because there are other Exaerete species by far larger than Ex. dentata, especially Ex. frontalis (Guérin-Méneville, 1844) and Ex. trochanterica (Friese, 1903). It is not possible, through the description and the figures displayed in the Plate 7 of Romand’s (1849) work to be sure of which Exaerete species is there represented [although it is reasonable to discard Ex. smaragdina (Guérin-Méneville, 1844) due to the absence of the median tubercle on the scutellum, if the drawing is exact regarding this character]. Only a study of the onomatophore, if found, can solve the question we propose here on the identity of Ch. subcornuta, since no modern author has seen this onomatophore (e.g. Moure 1967b; Kimsey 1979; Kimsey & Dressler 1986; Roubik 2004; Moure et al. 2007; Nemésio 2009a).

Orchid bees and orchids before Darwin A second interesting feature found on the description of Chrysantheda subcornuta is the presence of a projection [an “appendix”, as described by Romand (1849)] on the head of the examined specimen. This feature obviously caught the attention of both Spinola and Romand, since both nomina erected by them (“subcornuta” and “appendiculata”) clearly referred to this “character”. This “appendix” actually was an orchid pollinarium of an orchid flower (see Fig. 7), but Romand (1849: xxxvi–xxxvii) was convinced that it was a natural feature of males of Ch. subcornuta (“une pièce naturelle aux individus mâles”). Romand (1849) mentioned that the “appendix” has a pedicel which, in turn, is attached to a kind of “cavity” on the frons of the bee. According to him, such a “cavity” was absent from female specimens (and also from some males), and even males with this “cavity” usually lacked the “appendix”, because it is “easily detachable, being the reason why Chrysantheda males in European collections do not present the appendix” (Romand 1849: xxxvii).

Figure 7. Original illustration from Romand (1849). A, B and C are the same species; C lacks the "appendix", which is shown in detail in 1, 2 and 3.

It is commonly attributed to Darwin (1877: 170–176, 205–206, 270–271) the earliest observations on the relationships between orchid bees and orchids, although Darwin himself attributed all information to Crüger’s (1865) and Müller’s (1868, 1869) works and especially to his correspondence with both authors (Darwin 1877: v–vi). Orchid bees mentioned by Darwin (1877) are usually treated as Euglossa, but at that time most entomologists treated all orchid bees as Euglossa, including Smith (1874) who certainly identified some specimens for Darwin (Darwin 1877: 175). It is possible, however, to recognize at least two modern genera among the bees reported by Darwin: Euglossa (Glossura) (“a splendid bee, probably a Euglossa, but with the tongue nearly twice as long as the body”—Darwin 1877: 170), and Eulaema [(Euglossa cajennensis—now recognized as a junior synonym of Eulaema cingulata (Fabricius, 1804)—Darwin 1877: 206]. Euglossa (Glossura) piliventris Guérin-Méneville,



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1844 is also explicitly mentioned by Darwin (1877: 206). Nevertheless, although Darwin was aware that some pollinaria could be attached to body parts of insects such as legs, when referring to bees, particularly to orchid bees, he always mentioned pollinaria attached to the thorax. Darwin (and Crüger and Müller) also seemed to not realize that the orchid bees involved in interactions with orchids were males. It is not known whether Darwin had access to Romand’s (1849) work and whether he particularly paid attention to the Exaerete described by Romand. It is not possible to say from Darwin’s (1877) work (or even from other of his works on orchids and bees) if he or his correspondents observed orchids being visited by Exaerete because: (i) Darwin was much more interested in the orchids themselves than in their possible pollinators; (ii) maybe for this reason, details of the bees were usually neglected; and (iii) as the taxonomy of orchid bees at that time tended to consider Euglossa as a monotypic genus for orchid bees, it is impossible to know if one of the Euglossa mentioned by him was actually an Exaerete. Nonetheless, we can wonder if Darwin (or Crüger or Müller) would realize that an Exaerete bee visiting an orchid flower would be a male due to the lack of a corbicula (although male Euglossa and Eulaema also lack a corbicula and this fact apparently did not catch their attention). More important, at least when visiting Aspasia flowers, Exaerete males usually carry pollinaria on the face or head (R. L. Dressler, pers. comm.; Roubik & Hanson 2004: 30), exactly as happened to the male illustrated by Romand (1849). The fact is that Romand failed to recognize the “appendix” as vegetal matter and Crüger, Müller, and Darwin apparently overlooked this misunderstood record and were unable to realize that orchid-bee males, and not females, were attracted to orchids. The nature of the “cavity” described by Romand (1849) is uncertain and his drawing (here reproduced as Fig. 7) is not very helpful. Nevertheless, it could be interpreted as a character that varies from species to species, since Romand failed to find this character in many specimens. The protuberance on the frons of Exaerete frontalis, for example, could be such character, since most species lack it (only Ex. dentata present a similar, but smaller, protuberance). Concerning the identity of the pollinarium described by Romand (1849) as an “appendix” of the bee, R.L. Dressler (pers. comm.) believes it belongs to Aspasia, whereas G. Gerlach (pers. comm.) also suggested Catasetum and Trichocentrum as possible candidates, although D. Roubik (in his review to this manuscript) doubted this information and stated that members of Exaerete do not carry pollinaria of these latter orchid genera.

4. Eufriesea danielis (Schrottky, 1907) Eumorpha combinata var. danielis Schrottky, 1907

Moure (1967b) considered Eufriesea danielis as a junior synonym of Ef. violacea (Blanchard, 1840). This synonymy was followed by Kimsey (1982) and Kimsey & Dressler (1986). Moure (1967b) also stated that the onomatophore of this species was probably lost and was followed in this regard by all subsequent authors. Moure (1999), on the other hand, revalidated Ef. danielis and considered it to be part of a species group also comprising Ef. auriceps (Friese, 1899) and Ef. violascens (Mocsáry, 1898). He considered a specimen deposited at MPSP, numbered 102.887, as the holotype of Ef. danielis (see Moure 1999: 91). Nemésio & Silveira (2007: 886) considered Eufriesea danielis as a junior synonym of Ef. auriceps. Moure et al. (2007) considered Ef. danielis as a valid species and doubted the specimen listed by Moure (1999) is a true holotype, especially because Schrottky’s identification label dated from 1912. Nemésio (2009a) considered Ef. danielis as a synonym of Ef. auriceps again, based on (i) the fact that both onomatophores Ef. auriceps (a male) and Ef. danielis (a female) were collected at the same locality (Asunción, Paraguay) and (ii) the supposed holotype female of Ef. danielis matched the paralectotype female of Ef. violascens, from the state of Santa Catarina, southern Brazil, considered by Moure (1976) as Ef. auriceps. Nemésio (2009a: 30–32) supported Moure’s (1999) interpretation that the bee numbered 102.887 at MPSP is the true onomatophore of Ef. danielis for the reasons presented therein. Nevertheless, a re-examination of the original labels of the specimen numbered 102.887 at MPSP supports Moure et al. (2007) interpretation that this specimen should not be considered as the onomatophore of Eufriesea danielis. The fact that the bee only received a label with its specific identity in 1912 is not a reason strong enough to disqualify this specimen as an onomatophore, as another true onomatophore described in the same paper



(Schrottky 1907) was also only labeled in 1912 (see Nemésio 2009a: 30–32). The reason to disregard this specimen as the onomatophore is the label with the collector datum: “Schrottky leg.”. Schrottky (1907: 57), Moure et al. (2007) and even Nemésio (2009a: 30, footnote 4) reported that the species was described based on a female collected by János Dániel Anisits (Rasmussen et al. 2009), after whom the species was named. Although this specimen was collected in Paraguay and perfectly matches the description provided by Schrottky, it should not be considered the onomatophore as Moure (1999) and Nemésio (2009a) maintained, because the specimen does not have a typical Anisits label (with “J.D. Anisits” printed in small font, vertical, on left label margin). Rather its labels indicated that the specimen was collected by Schrottky himself [see photographs of the labels and of the specimen in Nemésio (2009a: 34)]. As the identity of this species has been problematic [it has been considered a junior synonym of different species, as mentioned above—which led Nemésio (2009a) to designate a neotype to Eufriesea violacea to avoid any future misunderstandings on the identity of this species], a thorough search for the onomatophore of Ef. danielis should be carried out. Type specimens collected by Anisits have been found at the ‘Zoologisches Museum der Humboldt Universität’, Berlin (CR, pers. obs.). It is also possible that specimens collected by D. Anisits are deposited at the ‘Museo Argentino de Ciencias Naturales’, Buenos Aires and at the Museo de La Plata. If, after a careful search, the onomatophore of Ef. danielis is not found, we strongly recommend that the specimen numbered as 102.887, deposited at MPSP, should be designated as the neotype of this species, due to the reasons pointed out in Nemésio (2009a: 30–32) and, especially, because it is the only known specimen belonging to this species identified by Schrottky himself.

5. Euglossa fimbriata Moure, 1968 Euglossa fimbriata Moure, 1968 Euglossa fimbriata Rebêlo & Moure, 1996

Moure et al. (2007) introduced a change in the authorship of Euglossa fimbriata, which was until then considered to be described by Rebêlo & Moure (1996). Nemésio (2009a: 87–89) disagreed with Moure et al. (2007), arguing that referring to the nomen fimbriata by Moure (1968) was not enough to establish a character to clearly distinguish a species, in such a way that this nomen could not be available from Moure (1968), even under the more flexible rules of the second edition of the Code, then in force. Both interpretations, however, provoked informal debate and we decided to submit the matter to zoologists with expertise in nomenclature (see Acknowledgment section). Unfortunately, this informal consultation resulted in no consensus following much debate. Finally, we decided to submit the situation informally to the Secretariat of the International Commission on Zoological Nomenclature, initially through Ellinor Michel and subsequently through David Notton. Eventually, Notton informed us that the matter provoked discussion at the ICZN Secretariat, but that they came to the view that Euglossa fimbriata is available from Moure (1968) reasoning the following: ·

· ·

·

Moure (1968) included a sentence with a character which is purported to distinguish Euglossa cyanaspis from Euglossa fimbriata, each from the other. (our translation “The length of the postglandular hairs is much shorter than in [Eg.] fimbriata, which distinguishes it from this latter species”). So there is a character which is purported to distinguish Eg. fimbriata and Article 13.1.1 is (minimally) satisfied. Article 13.1.1 has generally been interpreted to be satisfied by one character. It does not matter if the character is relative. It does not matter if the character does not in fact distinguish the species or cannot be objectively perceived, only that it has been purported to distinguish the species. Moure considered the nomen valid in his 1968 paper and so the nomen satisfies Article 11.5. That he did not consider it valid or available in subsequent publications does not prevent it being available from 1968. Possibly Moure did not intend to describe Eg. fimbriata in 1968, however, an author’s intentions are impossible to prove. The availability of nomina can be disclaimed under Article 8.3, but Moure (1968) did not do that, rather he used the nomen as valid, and hence available. It is only after 1999 that authors must make their intention to establish new nomina explict (Article 16.1). Therefore Moure’s intention, whatever that may have been, does not prevent the nomen from being available from 1968. Euglossa fimbriata Moure, 1968 has a separate type, a Bolivian specimen labelled by Moure as Holotype. Nevertheless, ICZN (recommendation 73F) recommends that where no holotype or syntype was fixed in the original publication for a nominal species-group taxon established before 2000, and when it is possible that the nominal species-group taxon was



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based on more than one specimen, an author should proceed as though syntypes may exist and, where appropriate, should designate a lectotype rather than assume a holotype (see also Article 74.6). Recognition of this specimen as lectotype (the action taken here) is in accordance with Article 72.4.1.1 and recommendation 73F. Thus Eg. fimbriata Moure 1968 is a quite different nomen from Eg. fimbriata Rebêlo & Moure 1996 (nec Moure 1968) which has a different type, different description and different authorship and date. The nomenclatural situation is therefore: · Euglossa fimbriata Moure, 1968–available · Euglossa fimbriata Rebêlo & Moure, 1996–available, but permanently invalid as a junior primary homonym of Eg. fimbriata Moure, 1968.

In spite of not being a formal decision by the Commission, we opted for accepting the above interpretation, since it was the most close to a “consensus” they could find. Besides David Notton, two other members of the ICZN Secretariat actively took part on this discussion: Steve Tracy and Svetlana Nikolaeva, through the enquiry service of the ICZN Secretariat. The specimen collected in Bolivia (Santa Cruz, Santiago) in December, 1959, deposited in Moure’s Collection and numbered DZUP 025524 at ‘Universidade Federal do Paraná’ is, thus, the lectotype (color photographs available in Nemésio 2009a: 89).

6. Euplusia yepezi Moure, 2000, nom. nud. Euplusia yepezi Moure, 2000: Errata et Addendum [inserted as a separate sheet in Revista Brasileira de Zoologia, 17 (2)].

Moure (1999) published a note on the taxonomy of Eufriesea (as Euplusia), describing new species and taking nomenclatural acts. One of these acts was to consider Eulaema nigrita raymondi (Schrottky, 1907) as a valid species of Eufriesea (Moure 1999: 94–95 incorrectly spelled as “raymoni”). Following this, Moure (1999) provided a detailed description of the onomatophore [deposited at MPSP with label 102.942, lectotype of Centris nigrita (Lep.) var. raymondi Schrottky, 1907 (Eulaema nigrita raymondi)—photographs of this specimen and labels made available by Nemésio 2009a: 39]. Nevertheless, Moure (2000) later corrected his interpretations concerning this species through a sheet inserted in the journal “Revista Brasileira de Zoologia”, volume 17, number 2 (no page number), titled “Errata et Addendum”, which was distributed on the 30th of June, 2000. In this “Errata et Addendum”, Moure (2000) mentioned that a serious mistake had been made—he did not mention if this mistake was his or the printers’—and reinterpreted Centris nigrita (Lep.) var. raymondi Schrottky, 1907 as Eulaema (Apeulaema) nigrita Lepeletier de Saint Fargeau, 1841. Moure (2000) stated that the rest of the text below Euplusia raymondi on pages 94–96 of the original paper (Moure 1999) referred to the description of a new species, Euplusia yepezi. Moure (2000) also stated that the abstract should be corrected to include a paragraph stating that “one male specimen from Rancho Grande, Aragua, Venezuela, is described as Euplusia yepezi sp. n. …”. This “Errata et Addendum” by Moure (2000), however, was overlooked by many subsequent authors, as Nemésio & Silveira (2007: 886) pointed out, and Eufriesea yepezi (Moure, 2000) was listed neither by Ramírez et al. (2002) nor by Roubik & Hanson (2004) in their checklists. Moure et al. (2007), in turn, considered this species as a junior synonym of Eufriesea venezolana (Schrottky, 1913), a position followed by Nemésio (2009a: 232) in his checklist. A re-study of the Code, its criteria for nomina availability and the dates of publication of Moure’s nomenclatural acts, however, show that this interpretation is incorrect and that Euplusia yepezi should be considered a nomen nudum. Article 21.6 of the Code, expressly states that if the date of publication specified in a work is a range of dates, the work is to be dated from the final day of the range...”. (our bold). It obviously means that the date of publication of Euplusia yepezi is 2000, not 1999, because the publication was interrupted and continued at a later date— and in particular the nomen of the species itself and a tentative onatomophore were only published in 2000. This date, however, introduces a new problem, as the following is mandatory for names published after 1999: “Article 16.4. Species-group names: fixation of name-bearing types to be explicit. Every new specific and subspecific name published after 1999, except a new replacement name (a nomen novum), for which the name-bearing type of the nominal taxon it denotes is fixed automatically [Art. 72.7], must be accompanied in the original publication [Art. 16.4.1] by the explicit fixation of a holotype, or syntypes, for the nominal taxon…”.



Moure (2000) did not fix a holotype. When Moure (2000) stated that “one male specimen from Rancho Grande, Aragua, Venezuela, is described as Euplusia yepezi sp. n. …” the assumption that this specimen is the holotype is implicit, not explicit. This assumption becomes confusing when Moure (2000, last line of the “Errata et Addendum”) textually stated that (translated from Portuguese) “page 95 [in Moure 1999] and subsequent ones remain as they are”. Moure (1999: 95) explicitly stated (translated from Portuguese) “a specimen from Rancho Grande, 1100 m, Aragua, Venezuela, [collected] on 21-VIII-1974, F. Fernando Y. and C. J. Rosales leg. Received from Dr. Yepes when of [my] visit to Maracay in 1980 and compared to the type at the Museu de Zoologia of the Universidade de São Paulo”. Therefore Moure (1999, 2000) did not explicitly consider the specimen from Venezuela as the onomatophore nor explicitly designated this specimen as onomatophore as mandatory by the Code for species described after 1999. By maintaining the original paper as it was, from page 95 onward, Moure (2000) contradicted Moure (1999) and did not explicitly designate an onomatophore, as the specimen implicitly regarded as the onomatophore in Moure (2000) was confusingly compared to a “type” at the MPSP. Given Moure’s (2000) changes, this MPSP type could not be interpreted as the type of Centris nigrita var. raymondi Schrottky, 1907 because Moure (2000) changed what was written at the end of page 94, removing the reference to the type specimen of C. nigrita var. raymondi. Thus, this “type” specimen compared to the specimen from Rancho Grande, Venezuela, is a non-explicit specimen under the new wording provided by Moure (2000). Our conclusion is that Euplusia yepezi is a nomen nudum and the specimen deposited at the DZUP should not be considered a primary onomatophore.

7. Updated catalogue of orchid bees Genus Aglae Lepeletier de Saint Fargeau & Audinet-Serville, 1825 Aglae Lepeletier de Saint Fargeau & Audinet-Serville, 1825: 105. Type species: Aglae coerulea Lepeletier de Saint Fargeau & Audinet-Serville, 1825, monobasic. Aglaa Schulz, 1906: 258, unjustified emendation of Aglae Lepeletier de Saint Fargeau & Audinet-Serville, 1825. Aglae coerulea Lepeletier de Saint Fargeau & Audinet-Serville, 1825: 105, Lectotype ♀, designated by Moure, 1967b: 413, MRSN: FRENCH GUIANA, Cayenne: Original spelling appears to be coerulea, not caerulea as currently used.

Genus Eufriesea Cockerell, 1908 Plusia Hoffmannsegg, 1817: 52–53 (not Ochsenheimer, 1816; Lepidoptera). Type species: Plusia superba Hoffmannsegg, 1817, monobasic. Eumorpha Friese, 1899: 126 (not Hübner, 1807; Lepidoptera). Type species: Euglossa pulchra Smith, 1854, by designation of Cockerell, 1908: 41. Eufriesea Cockerell, 1908: 41, replacement for Eumorpha Friese, 1899. Type species: Euglossa pulchra Smith, 1854, autobasic (see Eumorpha Friese, 1899). Eufriesia Lutz & Cockerell, 1920: 544, unjustified emendation of Eufriesea Cockerell, 1908. Euplusia Moure, 1943: 189–190, replacement for Plusia Hoffmannsegg, 1817. Type species: Plusia superba Hoffmannsegg, 1817, autobasic (see Plusia Hoffmannsegg, 1817). Paleoeuglossa Poinar, 1998: 30. Type species: Paleoeuglossa melissiflora Poinar, 1998, by original designation. (According to D.W. Roubik, not an Eufriesea due to the presence of scutellar tuft).

Eufriesea aeneiventris [Euglossa] (Mocsáry, 1896: 5–6), image: Nemésio, 2009a: 35a–f, Lectotype ♀, designated by Moure, 1967b: 406, HNHM: BRAZIL, Espírito Santo. Eufriesea andina [Euglossa (Eumorpha)] (Friese, 1925: 27–28), Lectotype ♀, designated by Moure, 1967b: 406, ZMHB: PERU, Cusco, Vilcanota. Eufriesea anisochlora [Euplusia] (Kimsey, 1977: 8–10, figs. 10, 22, 34), Holotype ♂, UCDC: PANAMA, Panama, Cerro Jefe, leg. Dressler.



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Eufriesea atlantica Nemésio, 2008 (“2007”): 148–149, image: Nemésio, 2009a: 40a–f, Holotype ♂, UFMG: BRAZIL, Minas Gerais, Marliéria, Parque Estadual do Rio Doce, leg. Nemésio. Eufriesea auriceps [Euglossa (Eulema!)] (Friese, 1899: 156), Holotype ♂, ZMHB: PARAGUAY, Central, Asunción. danielis [Eumorpha combinata ssp.] (Schrottky, 1907: 55, 56–57), image: Nemésio, 2009a: 16a–f (false type), Holotype ♀, ?; MPSP (=false type): PARAGUAY, Central, Asunción, leg. Anisits: Not synonymized in Moure et al. (2007). nordestina [Euplusia] (Moure, 1999: 100–101), image: Nemésio, 2009a: 21a–f, Holotype ♀, DZUP: BRAZIL, Paraíba, Santa Luzia, 300 m, leg. Portela: Not synonymized in Moure et al. (2007). aridicola [Euplusia] (Moure, Neves, & Viana, 2001: 842–844, figs. 1–4), image: Nemésio, 2009a: 20a–f, Holotype ♂, DZUP: BRAZIL, Bahia, Barra, leg. Neves & Viana: Not synonymized in Moure et al. (2007). Eufriesea auripes [Euglossa (Eulema!)] (Gribodo, 1882: 266–267), Holotype ♂, MSNG: FRENCH GUIANA, Cayenne. Eufriesea bare González & Gaiani, 1989: 151–152, fig. 2, Holotype ♂, MIZA: VENEZUELA, Amazonas, San Carlos de Río Negro, leg. Chacón & Yépez. Eufriesea boharti [Euplusia] (Kimsey, 1977: 10, figs. 11, 23, 35), Holotype ♂, UCDC: VENEZUELA, Aragua, Rancho Grande, leg. Bohart. Eufriesea brasilianorum [Euglossa (Eumorpha) combinata ssp.] (Friese, 1899: 147), image: Nemésio, 2009a: 43a–b, 45a–d, Holotype ♀, ZMHB: BRAZIL, Espírito Santo. Eufriesea buchwaldi [Euglossa (Eufriesea)] (Friese, 1923: 27–28), Lectotype ♀, designated by Moure, 1967b: 407, ZMHB: PERU. Eufriesea chaconi González & Gaiani, 1989: 149–150, fig. 1, Holotype ♂, MIZA: VENEZUELA, Bolívar, La Paragua, E of Chigua river, 425m, leg. Instituto de Zoología Agrícola Expedition. Eufriesea chalybaea [Euglossa (Eufriesea)] (Friese, 1923: 27), Lectotype ♂, designated by Moure, 1967b: 407 (cf. ICZN Art. 74.6), AMNH (type status unconfirmed): BOLIVIA, Cochabamba, Tarata. Eufriesea chrysopyga [Euglossa] (Mocsáry, 1898: 497–498), Holotype; invalid lectotype designation of holotype ♀, Moure, 1967b: 407, HNHM: BOLIVIA, La Paz, Cordillera Real, Zongo (“Songo”). Eufriesea coerulescens [Euglossa] (Lepeletier de Saint Fargeau, 1841: 11), Lectotype ♂, designated by Moure, 1967b: 407 (cf. ICZN Art. 74.6), MNHN: MEXICO: Original spelling appears to be coerulescens, not caerulescens as currently used. simillima [Euplusia] (Moure & Michener in Moure, 1965: 275–277), Holotype ♂, SEMC: MEXICO, Chihuahua, Maguarichi, leg. Wood & Karren. Eufriesea combinata [Euglossa] (Mocsáry, 1897: 446), image: Nemésio, 2009a: 14a–f, Holotype; invalid lectotype designation of holotype ♀, Moure, 1967b: 407, HNHM: BOLIVIA, Potosí, San Antonio. Eufriesea concava [Euglossa (Eulema!) mexicana ssp.] (Friese, 1899: 151), image: Nemésio, 2009a: 5a–d, Lectotype ♂, designated by Moure, 1967b: 407, ZMHB: NICARAGUA. Eufriesea convexa [Euglossa (Eulema!) mexicana ssp.] (Friese, 1899: 151), Holotype ♀, ZMHB: PERU, Loreto, Pebas (not BRAZIL, Tebas). Eufriesea corusca [Euplusia] (Kimsey, 1977: 10–12, figs. 12, 24, 36), Holotype ♂, UCDC: PANAMA, Colón, Barro Colorado Island, leg. Dressler. Eufriesea dentilabris [Euglossa (Eulema!)] (Mocsáry, 1897: 443–444), image: Nemésio, 2009a: 46a–d, Holotype ♂, HNHM: BRAZIL, Espírito Santo. Eufriesea distinguenda [Euglossa (Eulema!)] (Gribodo, 1882: 267–268), image: Nemésio, 2009a: 24a–e, Holotype ♀, MSNG: FRENCH GUIANA, Cayenne. Eufriesea dressleri [Euplusia] (Kimsey, 1977: 12, figs. 3, 13, 25, 37), Holotype ♂, UCDC: PANAMA, Colón, Piña, leg. Dressler. Eufriesea duckei [Euglossa (Eulaema)] (Friese, 1923: 28), image: Nemésio, 2009a: 6a–d, Holotype ♀, AMNH: BRAZIL, Amapá, Macapá, leg. Ducke. Eufriesea eburneocincta [Euplusia] (Kimsey, 1977: 12–13, figs. 4, 14, 26, 38), Holotype ♂, ? (not in USNM, UCDC, UF; paratypes in UCDC, UF): GUYANA, Pomeroon–Supenaam, Dawa, leg. Hamer. Eufriesea elegans [Eulaema] (Lepeletier de Saint Fargeau, 1841: 13), Lectotype ♀, designated by Moure, 1967b: 407 (cf. ICZN Art. 74.6), MRSN: FRENCH GUIANA, Cayenne.



Eufriesea excellens [Euglossa (Eumorpha)] (Friese, 1925: 27), Lectotype ♂, designated by Moure, 1967b: 408 (cf. ICZN Art. 74.6), ZMHB: ECUADOR, Guayas, Guayaquil, leg. Buchwald. Eufriesea fallax [Euglossa] (Smith, 1854: 381), Lectotype ♂, designated by Moure, 1967b: 408, BMNH 17b1175: BRAZIL, Pará. Eufriesea flaviventris [Euglossa (Eulema!)] (Friese, 1899: 152), Lectotype ♀, designated by Moure, 1967b: 408, HNHM: BRAZIL. Eufriesea formosa [Euglossa (Eumorpha)] (Mocsáry, 1908: 581), Lectotype ♀, designated by Moure, 1967b: 408 (cf. ICZN Art. 74.6), HNHM: BRAZIL, Amazonas, Tefé. Eufriesea fragrocara [Euplusia] (Kimsey, 1977: 13, figs. 5, 15, 27, 39), Holotype ♂, UCDC: PERU, Huánuco, Llullapichis, Río Pachitea, leg. Dressler. Eufriesea fuscatra [Euplusia danielis ssp.] (Moure, 1999: 94), Holotype ♂, DZUP: BRAZIL, Tocantins, Santa Isabel do Morro, Ilha do Bananal, leg. Alvarenga: Locality originally in the state of Goiás. Eufriesea heideri Nemésio & Bembé, 2008: 243–244, figs. 1a, 1c, 1e, 2, Holotype ♂, UFMG: BOLIVIA, Cochabamba, Chapare, Chimoré river, close to Entre Ríos, 310 m, leg. Heider. Eufriesea kimimari González & Gaiani, 1989: 150, fig. 2, Holotype ♂, MIZA: VENEZUELA, Táchira, Río Frio, leg. Romero. Eufriesea laniventris [Euglossa] (Ducke, 1902a: 403), Lectotype ♂, designated by Moure, 1967b: 408, MPEG: BRAZIL, Pará, Belém, leg. Ducke: The nomen appeared first both in a key and accompanied by a brief description (October 1st), thus giving this paper priority over the actual description, Ducke, 1902b: 567, 575 (December). Eufriesea limbata [Euglossa (Eulema!)] (Mocsáry, 1897: 442), Holotype ♀, HNHM: BRAZIL, Piauí. basalis [Eulema!] (Friese, 1898: 203–204), Lectotype ♀, designated by Moure, 1967b: 407, ZMHB: SURINAME, leg. Fruhstorfer. Eufriesea longipennis [Euglossa (Eumorpha)] (Friese, 1925: 28), Lectotype ♀, designated by Moure, 1967b: 408 (cf. ICZN Art. 74.6), ZMHB: ECUADOR, Guayas, Guayaquil, leg. Buchwald. Eufriesea lucida [Euplusia] (Kimsey, 1977: 13–15, figs. 16, 28, 40), Holotype ♂, UCDC: COLOMBIA, Valle del Cauca, Valle Anchicaya, leg. Kennedy. Eufriesea lucifera Kimsey, 1977: 18, figs. 17, 29, 41, Holotype ♂, UCDC: PANAMA, Panama, El Llano–Cartí Road, 19 km N of El Llano, leg. Siri. Eufriesea macroglossa [Euplusia] (Moure, 1965: 274–275), Holotype ♂, SEMC: COSTA RICA, Cartago, Turrialba, leg. Dodson. Eufriesea magrettii [Euglossa (Eumorpha)] (Friese, 1899: 148), Lectotype ♂, designated by Moure, 1967b: 408, MSNG: VENEZUELA. fulvohirta [Euglossa (Eumorpha) magrettii ssp.] (Friese, 1899: 148), Lectotype ♂, designated by Moure, 1967b: 408, MSNG: VENEZUELA. Eufriesea mariana [Euglossa Mariana] (Mocsáry, 1896: 4–5, pl. 1, fig. 4), Holotype ♀, HNHM: BOLIVIA, El Beni, Bueyes. superba [Euglossa] (Mocsáry, 1898: 498), Holotype ♂, HNHM: BOLIVIA, La Paz, Cordillera Real, Zongo (“Songo”): Junior homonym of superba Hoffmannsegg, 1817. tucumana [Euglossa (Eumorpha) mariana ssp.] (Schrottky, 1902: 117), Holotype ♀, MACN: ARGENTINA, Tucumán. pulcherrima [Euglossa (Eufriesea)] (Friese, 1923: 27), nom. nov. for Euglossa superba Mocsáry 1898, nec Hoffmannsegg 1817. Eufriesea melissiflora [Paleoeuglossa] (Poinar, 1998: 31–34, figs. 1–5), Holotype ♀, WACA: DOMINICAN REPUBLIC (amber inclusion). According to D.W. Roubik, not an Eufriesea due to the presence of scutellar tuft. Eufriesea mexicana [Euglossa (Eulema!) Mexicana] (Mocsáry, 1897: 444–445), Lectotype ♀, designated by Moure, 1967b: 408, HNHM: MEXICO, Veracruz, Presidio. Eufriesea micheneri Ayala & Engel, 2008: 228–234, figs. 1–9, Holotype ♂, UNAM: MEXICO, Jalisco, Mascota, leg. Ayala. Eufriesea mussitans [Apis] (Fabricius, 1787: 301), image: Nemésio, 2009a: 50a–d, Lectotype ♀, designated by Moure, 1967b: 408 (cf. ICZN Art. 74.6), ZMUC: FRENCH GUIANA, Cayenne.



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inermis [Euglossa (Eulema!) mexicana ssp.] (Friese, 1899: 151), Lectotype ♂, designated by Moure, 1967b: 408, AMNH (type status unconfirmed): VENEZUELA. Eufriesea nigrescens [Euglossa (Eufriesea) magretti ssp.] (Friese, 1923: 27), nom. nov. for Euglossa nigrita Friese 1899, nec Lepeletier de Saint Fargeau 1841. nigrita [Euglossa (Eumorpha) magrettii ssp.] (Friese, 1903: 574), Lectotype ♀, designated by Moure, 1967b: 409, AMNH (type status unconfirmed): ECUADOR, Bolívar, Balsapamba (Balzapamba), leg. Schulz (durch A Schulz). Eufriesea nigrohirta [Euglossa (Eumorpha)] (Friese, 1899: 142–143), Nemésio, 2005: 41, ♂; image: Nemésio, 2009a: 52a–d, 53d, Holotype ♀, ZMHB: BRAZIL, Pará. faceta [Euplusia] (Moure, 1999: 98–99), image: Nemésio, 2009a: 53a, 54a–d, Holotype ♀, DZUP: BRAZIL, Bahia, Mucugê, 984m, leg. Benton. Eufriesea opulenta [Euglossa (Eumorpha)] (Mocsáry, 1908: 582), Lectotype ♂, designated by Moure, 1967b: 409 (cf. ICZN Art. 74.6), HNHM: PERU, Junín, Chanchamayo. Eufriesea ornata [Eulema!] (Mocsáry, 1896: 3–4), image: Nemésio, 2009a: 38, Holotype ♂, HNHM: BRAZIL, Pará: Purchased through Dr. O. Staudinger’s insect dealership, Dresden, Germany. Eufriesea pallida [Euplusia] (Kimsey, 1977: 15, figs. 6, 8, 18, 30, 42), Holotype ♂, UCDC: MEXICO, Chiapas, Tuxtla Gutiérrez, leg. Dodson. Eufriesea pretiosa [Euglossa (Eulema!)] (Friese, 1903: 575), Syntype ♀, Moure, 1967b: 408 (cf. ICZN Art. 74.6), ?; “AMNH” (false type): COLOMBIA, Valle del Cauca, Cauca–Tal, near Popayán: Friese cited Ecuador and Caucatal as the type locality. However, Cauca–Tal is in Colombia. The presumably false type from AMNH is from Ecuador (Tungurahua, Santa Ines). Eufriesea pulchra [Euglossa] (Smith, 1854: 381), image: Nemésio, 2009a: 7a–d, Holotype ♀, BMNH 17b948: BRAZIL, Pará, Tapajós. Eufriesea purpurata [Euglossa] (Mocsáry, 1896: 5, fig. 5), Lectotype ♀, designated by Moure, 1967b: 409, HNHM: PERU, Loreto, Iquitos. Eufriesea pyrrhopyga Faria & Melo, 2011: 36–37, figs. 1–5, Holotype ♀, DZUP: BRAZIL, Pernambuco, Recife, leg. Lopes. Eufriesea rufocauda [Euplusia] (Kimsey, 1977: 15–16, figs. 7, 19, 31, 43), Holotype ♂, UCDC: PANAMA, Panama, Cerro Campana, leg. Kimsey. Eufriesea rugosa [Euglossa (Eumorpha)] (Friese, 1899: 152–153), Lectotype ♀, designated by Moure, 1967b: 409, ZMHB: MEXICO. Eufriesea schmidtiana [Euglossa (Eulema!)] (Friese, 1925: 26), Lectotype ♂, designated by Moure, 1967b: 409, ZMHB: COSTA RICA, Heredia, Puerto Viejo de Sarapiquí, leg. Schmidt (Albert). Eufriesea smaragdina [Centris] (Perty, 1833: 150, pl. 28, fig. 13), image: Nemésio, 2009a: 30 (org. plate), 33 (org. plate), 55a–d, 57, Holotype ♂, ZSM: BRAZIL, Minas Gerais, leg. Spix et al. Eufriesea superba [Plusia] (Hoffmannsegg, 1817: 53), Holotype ♂, ?: BRAZIL, Pará. manni [Eulaema] (Cockerell, 1912: 41–42), Holotype ♂, USNM 58113: BRAZIL, Rondônia, Abunã, leg. Mann & Baker. Eufriesea surinamensis [Apis] (Linnaeus, 1758: 579 (not 575)), image: Nemésio, 2009a: 28b, d, 29b, d, f, 60a–f, Holotype ♀, NHRS: SURINAME, leg. Rolander: (not Linnaeus, 1758: 575). See Linnaeus, 1767: 961 for First Reviser action and van der Vecht, 1959: 211 for a discussion of the priority of this nomen. tropica [Apis] (Linnaeus, 1758: 579), image: Nemésio, 2009a: 28a, 29a, c, e, Holotype ♀, UUZM: “Calidis regionibus” (possibly SURINAME). abdomenflavum [Apis abdomen flavum] (De Geer, 1773: 574, pl. 28, figs. 9, 10), image: Nemésio, 2009a: 34 (org. plate), Holotype ♀, NHRS: SURINAME: Possibly a syntype. angulata [Euglossa] (Mocsáry, 1897: 443), Lectotype ♀, designated by Moure, 1967b: 406, HNHM: BRAZIL, Piauí. bruesi [Eulaema] (Cockerell, 1914: 307–308), image: Nemésio, 2009a: 30c, Holotype ♀, USNM 58114: ECUADOR, Guayas, Guayaquil, leg. Brues. amabilis [Eulaema] (Cockerell, 1917a: 476–477), image: Nemésio, 2009a: 30b, 59, Holotype ♂, USNM 23155: BRAZIL, Amazonas, Manaus, leg. Merrill. tectora [Euplusia] (Kimsey, 1977: 16, figs. 20, 32, 44), Holotype ♂, UCDC: PANAMA, Colón, Frijoles, leg.



Dressler. Eufriesea theresiae [Euglossa (Eumorpha) Theresiae] (Mocsáry, 1908: 581–582), image: Nemésio, 2011a: 1, 2, Lectotype ♀, designated by Moure, 1967b: 409–410 (cf. ICZN Art. 74.6), HNHM: BRAZIL, Amazonas, Tefé. Eufriesea velutina [Euplusia] (Moure, 1999: 101–103), Holotype ♀, DZUP: VENEZUELA, Mérida. Eufriesea venezolana [Centris] (Schrottky, 1913: 708), Syntype ♂, MPSP: VENEZUELA. Eufriesea venusta [Euplusia] (Moure, 1965: 273–274), Holotype ♀, SEMC: PANAMA, Panama, Cerro Campana, leg. Hanson. Eufriesea vidua [Euplusia] (Moure, 1976: 277–281), Holotype ♀, HNHM: SURINAME. xantha [Euplusia] (Kimsey, 1977: 17–18, figs. 21, 33, 45), Holotype ♂, USNM: GUYANA, Upper Demerara– Berbice, Mazaruni–Potaro river, Rockstone, leg. Williams. Eufriesea violacea [Euglossa] (Blanchard, 1840: 405, pl. 7, fig. 3), image: Nemésio, 2009a: 61 (org. plate), 63a–d, Neotype ♂, Nemésio, 2009a: 69, fig. 63, UFMG: BRAZIL, Minas Gerais, Viçosa, leg. Soares. Eufriesea violascens [Euglossa] (Mocsáry, 1898: 497), image: Nemésio, 2009a: 15a–f, Lectotype ♀, designated by Moure, 1967b: 410, HNHM: BOLIVIA, La Paz, Cordillera Real, Zongo (“Songo”).

Genus Euglossa Latreille, 1802 Euglossa Latreille, 1802b: 436 Type species: Apis cordata Linnaeus, 1758, by designation of Taschenberg, 1883: 85: Published in April, before Latreille, 1802a: 385 which was published in November. Cnemidium Perty, 1833: 148-149 (not Goldfuss, 1826; Porifera). Type species: Cnemidium viride Perty, 1833, monobasic.

Six subgenera are recognized in Euglossa: Euglossa Latreille, 1802b: 436. Type species: Apis cordata Linnaeus, 1758, by designation of Taschenberg, 1883: 85: Published in April, before Latreille, 1802a: 385 which was published in November. Euglossa (Glossura) Cockerell, 1917b: 144. Type species: Euglossa piliventris Guérin-Méneville, 1844, by original designation. Euglossa (Euglossella) Moure, 1967b: 401, replacement for Cnemidium Perty, 1833. Type species: Cnemidium viride Perty, 1833, autobasic (see Cnemidium Perty, 1833). Euglossa (Dasystilbe) Dressler, 1978b: 193. Type species: Euglossa villosa Moure, 1968, by original designation. Euglossa (Glossurella) Dressler, 1982b: 131. Type species: Euglossa bursigera Moure, 1970, by original designation. Euglossa (Glossuropoda) Moure, 1989: 387. Type species: Euglossa intersecta Audouin, 1824, by original designation. Euglossa alleni [Euglossa (Euglossa)] Moure, 1968: 35–37, Holotype ♂, USNM 70774: COSTA RICA, Puntarenas, Palmar, leg. Allen & Allen. Euglossa. Euglossa allosticta [Euglossa (Glossura)] Moure, 1969: 242–246, Holotype ♂, USNM 70776: PANAMA, Colón, Gamboa, leg. Dressler: Placed in the subgenus Glossura by Moure et al. (2007). Glossura. Euglossa amazonica Dressler, 1982c: 146, fig. 1g, Holotype ♂, USNM: BRAZIL, Pará, Belém, leg. Dressler. Euglossa. Euglossa analis Westwood, 1840: 262, pl. 19, fig. 2, image: Nemésio, 2009a: 72 (org. plate), 86, 87, 88a–f, Lectotype ♂, designated by Moure, 1967b: 401, OUMNH: BRAZIL, Amazonas. Euglossa. bicolor Ducke, 1902a: 401, 402, Lectotype ♂, designated by Moure, 1967b: 401–402, MPEG: BRAZIL, Pará, Belém, leg. Ducke: The nomen appeared first both in a key and accompanied by a brief description (October 1st), thus giving this paper priority over the actual description, Ducke, 1902b: 565–566, 570 (December). Euglossa. Euglossa anodorhynchi Nemésio, 2006: 206–208, fig. 1, image: Nemésio, 2009a: 89a–d, Holotype ♂, UFMG: BRAZIL, Santa Catarina, Joinville, leg. Soares. Euglossa. Euglossa aratingae Nemésio, 2009a: 106–108, figs. 84b, d, f, h, 91a–d, Holotype ♂, UFMG: BRAZIL, São Paulo, Ribeirão Preto, leg. Garófalo, Camillo & Serrano. Euglossa.



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Euglossa asarophora [Euglossa (Glossura)] Moure & Sakagami in Moure, 1969: 239–242, Holotype ♂, DZUP: PANAMA, Chiriquí, 2500–4000ft., leg. Champion: Although the holotype is stated to be in BMNH it is actually in DZUP; Placed in the subgenus Glossura by Moure et al. (2007). Glossura. Euglossa atroveneta [Euglossa (Euglossa)] Dressler, 1978a: 177–178, figs. 2h, 3h, Holotype ♂, USNM: GUATEMALA, Santa Rosa, leg. Tan. Euglossa. Euglossa augaspis Dressler, 1982b: 133, fig. 1a, image: Nemésio, 2009a: 93b–d, Holotype ♂, USNM: BRAZIL, Amazonas, Manaus, leg. Dressler. Glossurella. Euglossa aureiventris [Euglossa (Euglossa) cordata ssp.] Friese, 1899: 135, Lectotype ♀, designated by Moure, 1967b: 401, NMW: BRAZIL. Euglossa. Euglossa auriventris [Euglossa cordata ssp.] Friese, 1925: 29, Syntype ♀, ZMHB: BRAZIL, Acre, Rio Branco; ECUADOR, Guayas, Guayaquil. Euglossa. Euglossa avicula Dressler, 1982c: 146–147, fig. 1h, image: Nemésio, 2009a: 85a, c, e, 94a–c, Holotype ♂, USNM: BRAZIL, Espírito Santo, Conceição da Barra, leg. Dressler. Euglossa. Euglossa bembei Nemésio, 2011d: 44–45, fig. 1a, c, e, g, Holotype ♂, UFMG: BRAZIL, Minas Gerais, Santa Maria do Salto, leg. Nemésio. Euglossa. Euglossa bidentata Dressler, 1982a: 122, fig. 1a, Holotype ♂, USNM: PERU, Huánuco, Llullapichis, Río Pachitea, leg. Dressler. Euglossa. Euglossa bigibba [Euglossa (Euglossella)] Dressler, 1982a: 128–129, fig. 3b, Holotype ♂, HNHM: PERU, Loreto, Pebas. Euglossella. Euglossa bursigera [Euglossa (Euglossella)] Moure, 1970: 151–152, figs. 1c, 2c, Holotype ♂, USNM 70775: PANAMA, Colón, Barro Colorado Island, leg. Hanson. Glossurella. cupreicolor [Euglossa (Euglossella) bursigera ssp.] Moure, 1970: 152, Holotype ♂, DZUP: COSTA RICA, San José, Hacienda Pozo Azul, leg. Wille & Ramírez. Glossurella. Euglossa carinilabris Dressler, 1982b: 134, fig. 1b, image: Nemésio, 2009a: 95b–d, Holotype ♂, USNM: BRAZIL, Bahia, Itabuna, leg. Dressler: Junior synonym of stellfeldi in Moure et al. (2007). Glossurella. Euglossa carolina Nemésio, 2009a: 111–113, figs. 73b, d, 96a–d, Holotype ♂, UFMG: BRAZIL, São Paulo, Ribeirão Preto, leg. Serrano & Augusto. Euglossa. Euglossa chalybeata [Euglossa ignita ssp.] Friese, 1925: 29, image: Nemésio, 2009a: 2b, 65a–d, Lectotype ♂, designated by Moure, 1967b: 402, ZMHB: BRAZIL, Pará, Belém, leg. Ducke. Glossura. Euglossa championi Cheesman, 1929: 148–149, figs. 6, 7, Lectotype ♂, designated by Moure, 1967b: 402, BMNH 17b952: PANAMA, Chiriquí, leg. Champion. Euglossa. Euglossa charapensis Cockerell, 1917b: 146, Holotype ♀, USNM 23142: PERU, Cajamarca, Río Charape, leg. Townsend: A species distinct from aureiventris Friese. Euglossa. Euglossa chlorina Dressler, 1982c: 141–142, fig. 1a, Holotype ♂, USNM: VENEZUELA, Miranda, Caracas, leg. Dressler. Euglossa. Euglossa cognata [Euglossa (Euglossella)] Moure, 1970: 156–157, figs. 1f, 2f, image: Nemésio, 2009a: 98a–d, Holotype ♂ nomen protectum, DZUP: BRAZIL, Pará, Mocajuba, Mangabeira, leg. Rego. Euglossa. bureaui Dominique, 1898: 58, Lectotype ♂, designated by Rasmussen et al. 2007: 60, nomen oblitum, MHNAN: FRENCH GUIANA, Saint–Laurent du Maroni, Íle Portal, leg. Constant Bar and family. Euglossa. Euglossa cordata [Apis] (Linnaeus, 1758: 575), image: Nemésio, 2009a: 67a–e, 70a, Holotype ♀, UUZM: “Indiis” (possibly WEST INDIES or SURINAME). Euglossa. Euglossa cosmodora [Euglossa (Euglossella)] Hinojosa-Díaz & Engel, 2007b: 94–100, figs. 1–12, Holotype ♂, SEMC: PERU, Junín, Villa Rica–Oxapampa Road, leg. Brooks. Euglossella. Euglossa cotylisca Hinojosa-Díaz & Engel, 2007a: 2–5, figs. 1, 2, Holotype ♂, BMNH: COLOMBIA, Santander (Quaternary copal inclusion). Euglossa. Euglossa crassipunctata [Euglossa (Euglossa)] Moure, 1968: 40–43, image: Nemésio, 2009a: 75a, c, e, 99a–c, Holotype ♂, USNM 70777: PANAMA, Colón, Gamboa, leg. Dressler. Glossurella. Euglossa crininota [Euglossa (Euglossa)] Dressler, 1978a: 176–177, figs. 2g, 3g, Holotype ♂, USNM: MEXICO, Veracruz, Córdoba, leg. Spangler. Euglossa. Euglossa cyanaspis [Euglossa (Euglossa)] Moure, 1968: 46–49, image: Nemésio, 2009a: 73a, c, Holotype ♂, USNM 70778: PANAMA, Colón, Diablo Heights, leg. Dressler. Euglossa. Euglossa cyanea [Euglossa (Euglossa) variabilis ssp.] Friese, 1899: 135, Lectotype ♀ (or possibly ♂), designated



by Moure, 1967b: 402, HNHM: BOLIVIA, Potosí, San Antonio. Euglossella. Euglossa cyanochlora [Euglossa (Euglossella)] Moure, 1996 (“1995”): 468–469, image: Nemésio, 2009a: 100, 101, 103a–d, Holotype ♀, MNRJ: BRAZIL, Bahia, Itamaraju, leg. Roppa & Becker. Glossuropoda. Euglossa cyanura Cockerell, 1917b: 146, Holotype ♀, USNM 23144: PANAMA, Colón, Portobello, leg. Busck. Euglossella. Euglossa cybelia [Euglossa (Euglossa)] Moure, 1968: 26–29, image: Nemésio, 2009a: 9b–d, Holotype ♂, USNM 70779: PANAMA, Panama, Cerro Campana, leg. Dressler. Euglossa. Euglossa deceptrix [Euglossa (Euglossa)] Moure, 1968: 58–61, image: Nemésio, 2009a: 11a–d, Holotype ♂, USNM 70780: PANAMA, Panama, Cerro Campana, leg. Dressler. Euglossa. Euglossa decorata Smith, 1874: 444–445, image: Nemésio, 2009a: 105a–e, Holotype ♀, BMNH 17b949: BRAZIL, Amazonas, São Paulo de Olivença. Euglossella. ruficauda [Euglossa decorata ssp.] Cockerell, 1918: 688, Holotype ♀, AMNH: GUYANA, Cuyuni–Mazaruni, Kalacoon, leg. Tee Van. Euglossella. Euglossa despecta [Euglossa (Euglossa)] Moure, 1968: 55–58, image: Nemésio, 2009a: 106b–d, Holotype ♂, USNM 70781: PANAMA, Colón, Barro Colorado Island, leg. Dressler. Euglossa. violaceifrons [Euglossa (Euglossa)] Rebêlo & Moure, 1996(“1995”): 465, figs. 3k, 5k, 7k, Holotype ♂, DZUP? (not located, nor in RPSP and possibly lost): BRAZIL, São Paulo, Cajuru, Fazenda Santa Carlota, leg. Rebêlo: Not synonymized in Moure et al. (2007). Euglossa. Euglossa dissimula [Euglossa (Euglossa)] Dressler, 1978a: 173–174, figs. 2e, 3e, Holotype ♂, USNM: PANAMA, Panama, Cerro Campana, leg. Dressler. Euglossa. Euglossa dodsoni [Euglossa (Euglossa)] Moure, 1965: 266–269, Holotype ♀, SEMC: COSTA RICA, Cartago, 14 km E Turrialba, 650 m, leg. Dodson. Glossurella. Euglossa dressleri [Euglossa (Euglossa)] Moure, 1968: 29–32, Holotype ♂, USNM 70782: PANAMA, Colón, Gamboa, leg. Dressler. Euglossa. Euglossa erythrochlora [Euglossa (Euglossa)] Moure, 1968: 32–35, Holotype ♂, USNM 70783: COSTA RICA, Puntarenas, Las Cruces, San Vito de Java, leg. Dressler. According to D.W. Roubik, molecular data (CO1) shows this species is indistinguishable from hemichlora, and both hybridize in Coíba Island (PANAMA). Euglossa. Euglossa erythrophana [Euglossa (Glossura) gorgonensis ssp.] Dressler, 1978a: 170–171, figs. 2c, 3c, 4b, Holotype ♂, USNM 70773: COSTA RICA, Puntarenas, Golfito, leg. Dressler: Placed in the subgenus Glossura by Moure et al. (2007). Glossurella. Euglossa fimbriata [Euglossa (Euglossa)] Moure, 1968: 48–49 Nemésio, 2009a: 76a–d, 77, Holotype ♂, DZUP: BOLIVIA, Santa Cruz, Santiago. Euglossa. Euglossa flammea [Euglossa (Glossura)] Moure, 1969: 231–234, figs. 1–4, Holotype ♂, USNM 70784: PANAMA, Colón, Gamboa, leg. Dressler. Glossura. Euglossa fuscifrons Dressler, 1982b: 134–135, fig. 1c, Holotype ♂, USNM: ECUADOR, Pastaza, Veracruz, leg. Velástegui. Glossurella. Euglossa gibbosa Dressler, 1982c: 147–148, fig. 1i, Holotype ♂, USNM: ECUADOR, Santo Domingo, Santo Domingo, leg. Dressler. Euglossa. Euglossa gorgonensis Cheesman, 1929: 146–147, fig. 4, Syntype (2) ♀, BMNH 17b947: COLOMBIA, Valle del Cauca, Gorgona Island, leg. St. George Expedition. Glossurella. Euglossa granti Cheesman, 1929: 147–148, fig. 5, Holotype ♀, BMNH 17b945: COLOMBIA, Valle del Cauca, Gorgona Island, leg. St. George Expedition. Euglossella. Euglossa hansoni [Euglossa (Euglossa)] Moure, 1965: 269–270, Holotype ♀, SEMC: PANAMA, Colón, Piña, leg. Hanson. Euglossa. Euglossa hemichlora [Euglossa variabilis ssp.] Cockerell, 1917b: 146, image: Nemésio, 2009a: 78, 79b, d, f, Holotype ♀, USNM 23145: ECUADOR, Imbabura, Parámba, leg. Rosenberg. Euglossa. auricollis [Euglossa cordata ssp.] Friese, 1923: 26, Lectotype ♀, designated by Moure, 1967b: 401, ZMHB: ECUADOR, Guayas, Guayaquil, leg. Buchwald: Not synonymized in Moure et al. (2007). Euglossa. azureoviridis [Euglossa variabilis ssp. azureovirida] Friese, 1930: 137, Lectotype ♂, designated by Moure, 1967b: 401, ZMHB: COSTA RICA, San José, leg. Schmidt (H.): Emended to azureoviridis by Moure, 1967b: 401. Euglossa.



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nigrescens [Euglossa variabilis ssp.] Friese, 1930: 137, Lectotype ♀, designated by Moure, 1967b: 404, ZMHB: COSTA RICA, San José, leg. Schmidt (H.). Euglossa. gaianii Dressler, 1982c: 142–143, fig. 1b, image: Nemésio, 2009a: 79a, c, e, Holotype ♂, USNM: FRENCH GUIANA, Kourou, 12 km SW Kourou, leg. Roubik: Not synonymized in Moure et al. (2007). Euglossa. Euglossa heterosticta [Euglossa (Euglossa)] Moure, 1968: 52–55, image: Nemésio, 2009a: 85b, d, f, Holotype ♂, USNM 70785: PANAMA, Panama, Cerro Campana, leg. Dressler. Euglossa. Euglossa hyacinthina Dressler, 1982b: 135, fig. 1d, Holotype ♂, USNM: PANAMA, Chiriquí, Fortuna Dam, leg. Dressler. Glossurella. Euglossa ignita Smith, 1874: 444, image: Nemésio, 2009a: 110a, 111, 112a–d, Holotype ♂, BMNH 17b950: JAMAICA.According to D.W. Roubik, this species does not occur in Jamaica, suggesting a possible label error. Glossura. bari Dominique, 1898: 58, Lectotype ♂, designated by Rasmussen et al. 2007: 60, MHNAN: FRENCH GUIANA, Saint–Laurent du Maroni, Íle Portal, leg. Constant Bar and family. Glossura. chlorosoma [Euglossa ignita ssp.] Cockerell, 1918: 688, image: Nemésio, 2009a: 113a–d, Holotype ♀, AMNH: GUYANA, Cuyuni–Mazaruni, Bartica, leg. Tee Van. Glossura. Euglossa igniventris Friese, 1925: 29–30, Lectotype ♀, designated by Moure, 1967b: 403 (cf. ICZN Art. 74.6), ZMHB: COSTA RICA, San José, La Caja, leg. Schmidt (H.). Euglossa. cupreiventris Cheesman, 1929: 146, fig. 3, Holotype ♀, BMNH 17b946: PANAMA, leg. St. George Expedition. Euglossa. Euglossa imperialis [Euglossa (Glossura) piliventris ssp.] Cockerell, 1922: 6, image: Nemésio, 2009a: 114, 116a– d, Holotype ♀, USNM 24882: PANAMA, Bocas del Toro, Río Trinidad, leg. Busck. Glossura. Euglossa inflata [Euglossa (Glossuropoda)] Roubik, 2004: 236–239, figs. 1, 2, Holotype ♂, USNM: FRENCH GUIANA, Kourou, 14 km SW, 9 km S Kourou, leg. Roubik. Glossuropoda. Euglossa intersecta Audouin, 1824: 340–341, Syntype unknown sex, ?: FRENCH GUIANA, Cayenne. Glossuropoda. brullei Lepeletier de Saint Fargeau, 1841: 10, Lectotype ♀, designated by Moure, 1967b: 402 (cf. ICZN Art. 74.6), MNHN: “SENEGAL” in error. Glossuropoda. romandii Guérin-Méneville, 1844: 458–459, Lectotype ♂, designated by Moure, 1967b: 405, MHNG: Unknown locality. Glossuropoda. Euglossa iopoecila [Euglossa chalybeata ssp.] Dressler, 1982a: 126–127, image: Nemésio, 2009a: 117b–d, Holotype ♂, USNM: BRAZIL, Paraná, Alexandra, leg. Dressler. Glossura. Euglossa ioprosopa Dressler, 1982a: 124–125, fig. 3c, image: Nemésio, 2009a: 118b–d, Holotype ♂, USNM: PERU, Huánuco, Llullapichis, Río Pachitea, leg. Dressler. Euglossa. Euglossa iopyrrha Dressler, 1982a: 123–124, fig. 1c, image: Nemésio, 2009a: 120, 121a–d, Holotype ♂, USNM: BRAZIL, Pará, Óbidos, leg. Brazilino. Euglossa. Euglossa jacquelynae [Euglossa (Euglossella)] Nemésio, 2007: 22–24, figs. 1a–d, image: Nemésio, 2009a: 122a– d, Holotype ♂, UFMG: BRAZIL, Goiás, Caldas Novas, leg. Silva. Euglossella. Euglossa jamaicensis [Euglossa (Euglossa)] Moure, 1968: 43–46, Holotype ♂, DZUP: JAMAICA. Euglossa. Euglossa juremae [Euglossa (Glossuropoda)] Moure, 1989: 389, image: Nemésio, 2009b: 1b, d, f, h, 2b, d, f, Holotype ♂, MNRJ: BRAZIL, Pará, Vigia, leg. Silva: Junior synonym of hugonis in Roubik (2004) and Moure et al. (2007). Reinstated as valid in Nemésio (2009b). Glossuropoda. Euglossa laevicincta Dressler, 1982b: 135–136, fig. 1e, Holotype ♂, USNM: FRENCH GUIANA, Kourou, 19 km SW Kourou, leg. Michener. Glossurella. Euglossa laurensi Bembé, 2008: 60–63, figs. 1, 2a, 3, Holotype ♂, ZSM: BOLIVIA, Cochabamba, Villa Tunari, leg. Bembé. Euglossa. Euglossa lazulina [Euglossa cordata ssp.] Friese, 1923: 26, Lectotype ♀, designated by Moure, 1967b: 404 (cf. ICZN Art. 74.6), ZMHB: BRAZIL, Pará, Óbidos. Unplaced. Euglossa leucotricha [Euglossa (Euglossa)] Rebêlo & Moure, 1996 (“1995”): 462, figs. 3g, 5g, 7g, image: Nemésio, 2009a: 123a–d, Holotype ♂, RPSP: BRAZIL, São Paulo, Dumont, Boa Vista, leg. Camilo & Macário. Euglossa. Euglossa liopoda Dressler, 1982c: 143, fig. 1c, image: Nemésio, 2009a: 124b–d, Holotype ♂, USNM: VENEZUELA, Bolívar, Chicanán river, leg. Dressler. Euglossa. Euglossa lugubris [Euglossa (Glossura)] Roubik, 2004: 244–245, fig. 10, Holotype ♂, USNM: PERU, Madre de



Díos, 30 km SW Puerto Maldonado, leg. Frisbie. Glossura. Euglossa macrorhyncha Dressler, 1982b: 136–137, fig. 1f, Holotype ♂, USNM: ECUADOR, Santo Domingo, Santo Domingo, leg. Dressler. Glossurella. Euglossa maculilabris [Euglossa (Euglossa)] Moure, 1968: 23–26, Holotype ♂, USNM 70786: PANAMA, Panama, Cerro Campana, leg. Dressler. Euglossa. Euglossa magnipes Dressler, 1982c: 148, fig. 1j, Holotype ♂, USNM: PERU, Huánuco, Llullapichis, Río Pachitea, leg. Dressler. Euglossa. Euglossa mandibularis [Euglossa (Euglossa)] Friese, 1899: 137–138, image: Nemésio, 2009a: 125a–d, 128, Lectotype ♀, designated by Moure, 1967b: 404, ZMHB: BRAZIL, Rio Grande do Sul / Santa Catarina, Santa Cruz: Uncertain locality. Euglossella. bernardina [Euglossa mandibularis ssp.] Cockerell, 1917b: 144–145, image: Nemésio, 2009a: 130b–d, Holotype ♂, USNM 23143: PARAGUAY, Cordillera, San Bernardino, leg. Fiebrig. Euglossella. aenescens Friese, 1925: 28–29, image: Nemésio, 2009a: 131a–d, Lectotype ♂, designated by Moure, 1967b: 401, ZMHB: BRAZIL, Minas Gerais, Passa Quatro, leg. Zikán. Euglossella. Euglossa marianae Nemésio, 2011b: 63, figs. 2a–e, Holotype ♂, UFMG: BRAZIL, Minas Gerais, Marliéria, Parque Estadual do Rio Doce, leg. Nemésio. Euglossa. Euglossa melanotricha [Euglossa (Euglossa)] Moure in Sakagami, Laroca, & Moure, 1967: 47–50, image: Nemésio, 2009a: 132a–d, Holotype ♂, DZUP: BRAZIL, Minas Gerais, Araxá, leg. Elias. Euglossa. Euglossa meliponoides [Euglossa (Euglossa)] Ducke, 1902b: 563–564, 569, stat. resur., Lectotype ♂, designated by Moure, 1967b: 404, MPEG: BRAZIL, Pará, Itaituba, leg. Ducke. Lectotype is given as ♀ in all previous catalogues, but it is a ♂ (Nemésio, in prep.). Euglossella. Euglossa micans [Euglossa (Euglossa)] Dressler, 1978a: 175–176, figs. 2f, 3f, Holotype ♂, UF: COSTA RICA, Heredia, Puerto Viejo de Sarapiquí, La Selva, leg. Dodson: Although the holotype is stated to be in Marie Selby Botanical Gardens it is actually in UF. Euglossa. Euglossa milenae Bembé, 2007: 59–62, 143, figs. 5f, 7a, 9e, 11a, 14a–b, image: Nemésio, 2009a: 12a–d, Holotype ♂, ZSM: BOLIVIA, Cochabamba, Villa Tunari, leg. Bembé. Euglossa. Euglossa mixta [Euglossa (Euglossa) variabilis ssp.] Friese, 1899: 135, image: Nemésio, 2009a: 134a–d, 137a, Lectotype ♂, designated by Moure, 1967b: 404, HNHM: PANAMA, Chiriquí. Euglossa. Euglossa modestior Dressler, 1982c: 143–144, fig. 1d, image: Nemésio, 2009a: 148b–d, Holotype ♂, USNM: PERU, Loreto, Iquitos, leg. Dressler. Euglossa. Euglossa moronei Engel, 1999: 4–8, figs. 1, 2, 4, 5, Holotype ♀, “Ettore Morone amber collection”, Turin, Italy: DOMINICAN REPUBLIC (amber inclusion). Unplaced. Euglossa mourei Dressler, 1982c: 148–150, fig. 1k, Holotype ♂, USNM: COLOMBIA, Amazonas, Leticia, leg. Dressler. Euglossa. Euglossa nanomelanotricha Nemésio, 2009a: 143–145, figs. 138, 139a–d, Holotype ♂, UFMG: BRAZIL, Paraíba, Cabedelo, leg. Peixoto. Euglossa. Euglossa natesi [Euglossa (Glossura)] Parra-H, Ospina-Torres, & Ramírez, 2006: 30–34, figs. 1–10, Holotype ♂, ICN (Laboratorio de Investigaciones en Abejas): COLOMBIA, Nariño, Barbacoas, La Tajada, leg. Parra–H. Glossura. Euglossa nigropilosa [Euglossa (Euglossa)] Moure, 1965: 270–273, Holotype ♀, SEMC: ECUADOR, Tungurahua, Topo, 700–1400 m, leg. Dodson. Euglossa. Euglossa nigrosignata [Euglossa (Glossura)] Moure, 1969: 234–238, Holotype ♂, USNM 70788: PANAMA, Coclé, El Valle de Antón, leg. Dressler: Placed in the subgenus Glossuropoda by Roubik (2004). Glossuropoda. Euglossa obrima Hinojosa-Díaz, Melo, & Engel, 2011: 14–26, figs. 1, 3, 4, 6, 7, 9–15, 17, 19, 21–26, Holotype ♂, SEMC: MEXICO, Veracruz, 34 km N Catemaco, leg. Wenzel. Dasytilbe. Euglossa obtusa [Euglossa (Glossura)] Dressler, 1978a: 171–173, figs. 2d, 3d, Holotype ♂, USNM: MEXICO, Veracruz, Los Tuxtlas near Montepío, leg. Dressler. Glossurella. Euglossa occidentalis [Euglossa (Glossura)] Roubik, 2004: 246–248, fig. 12, Holotype ♂, QCAZ: ECUADOR, Napo, Yasuní National Park, leg. Roubik. Glossura. Euglossa oleolucens [Euglossa (Glossura)] Dressler, 1978a: 169–170, figs. 2b, 3b, 4a, Holotype ♂, USNM: COSTA RICA, Puntarenas, Las Cruces, San Vito de Java, leg. Dressler. Glossurella. Euglossa orellana [Euglossa (Glossura)] Roubik, 2004: 248–250, fig. 13, image: Nemésio, 2009a: 66a–b, HoloNOMENCLATURAL ISSUES IN THE ORCHID BEES


27

type ♂, QCAZ: ECUADOR, Napo, Yasuní National Park, leg. Roubik. Glossura. Euglossa paisa [Euglossa (Glossurella)] Ramírez, 2005: 53–57, figs. 1–8, Holotype ♂, IAVH: COLOMBIA, Antioquia, Anorí, leg. Arias. Glossurella. Euglossa parvula Dressler, 1982b: 137–138, fig. 1g, Holotype ♂, USNM: COLOMBIA, Amazonas, Leticia, leg. Dressler. Glossurella. Euglossa perfulgens [Euglossa (Euglossa)] Moure, 1967a: 388–391, Holotype ♀, DZUP: BRAZIL, Amazonas, Tefé, leg. Carvalho. Euglossella. Euglossa perpulchra [Euglossa (Euglossella)] Moure & Schlindwein, 2002: 586–588, image: Nemésio, 2009a: 140a–d, Holotype ♂, DZUP: BRAZIL, Pernambuco, Igarassu, Reserva Ecológica Charles Darwin, leg. Schlindwein & Martini. Euglossella. Euglossa perviridis Dressler, 1985: 78–79, Holotype ♂, USNM: PERU, Madre de Díos, 30 km SW Puerto Maldonado, leg. Pearson. Euglossella. Euglossa pictipennis Moure, 1943: 191, image: Nemésio, 2009a: 80a–c, Holotype ♀, DZUP: BRAZIL, São Paulo, Batatais, leg. Rodrigues. Euglossa. Euglossa piliventris Guérin-Méneville, 1844: 458, Lectotype ♀, designated in present work, MSNG: BRAZIL, Pará. Glossura. Euglossa platymera Dressler, 1982c: 144–145, fig. 1e, Holotype ♂, USNM: BRAZIL, Amazonas, Manaus, leg. Elias. Euglossa. Euglossa pleosticta Dressler, 1982c: 150, fig. 1l, Holotype ♂, USNM: BRAZIL, Espírito Santo, Linhares, leg. Dressler. Euglossa. Euglossa polita Ducke, 1902a: 400, 402, Lectotype ♀, designated by Moure, 1967b: 405, MPEG: BRAZIL, Pará, Belém, leg. Ducke: The nomen appeared first both in a key and accompanied by a brief description (October 1st), thus giving this paper priority over the actual description, Ducke, 1902b: 564–565, 571 (December); All specimens in this type series were labeled types and should be considered syntypes of which a lectotype has been selected (incl. also paralectotype specimen BMNH 17b 944); Lectotype is given as ♂ in all previous catalogues, but it is a ♀ (Nemésio, in prep.). Unplaced. Euglossa prasina Dressler, 1982b: 138, fig. 1h, Holotype ♂, USNM: COLOMBIA, Amazonas, Leticia, leg. Dressler. Glossurella. Euglossa purpurea [Euglossa (Euglossa) variabilis ssp.] Friese, 1899: 135, Lectotype ♂, designated by Moure, 1967b: 405, HNHM: PANAMA, Chiriquí. Euglossa. Euglossa retroviridis Dressler, 1982a: 123, fig. 1b, Holotype ♂, USNM: COLOMBIA, Amazonas, Leticia, leg. Dressler. Euglossa. Euglossa roderici Nemésio, 2009a: 149–150, figs. 143a–d, 144, Holotype ♂, UFMG: BRAZIL, São Paulo, São Sebastião, leg. Mateus. Euglossa. Euglossa roubiki Nemésio, 2009a: 151–152, figs. 145a–d, 146, Holotype ♂, UFMG: BRAZIL, Bahia, Porto Seguro, leg. Cordeiro. Glossura. Euglossa rufipes [Euglossa (Glossurella)] Rasmussen & Skov, 2006: 55–61, figs. 1, 3, 5, 7, 8, 9, Holotype ♂, MUSM: PERU, San Martín, Tarapoto–Yurimaguas road km 20, leg. Rasmussen: Placed in the subgenus Glossura by Moure et al. (2007). Glossura. Euglossa rugilabris [Euglossa (Glossura)] Moure, 1967a: 391–394, image: Nemésio, 2009b: 4a–f, 5, Holotype ♀, DZUP: BRAZIL, Amazonas, Tabatinga, leg. Oliveira. Glossuropoda. hugonis [Euglossa (Glossuropoda)] Moure, 1989: 387–389, image: Nemésio, 2009b: 1a, c, e, g, 2a, c, e, Holotype ♂, MNRJ: BRAZIL, Amazonas, Tabatinga, leg. Silva. Glossuropoda. Euglossa samperi [Euglossa (Glossurella)] Ramírez, 2006: 62–65, figs. 1–5, 9–12, Holotype ♂, QCAZ: ECUADOR, Esmeraldas, Bilsa Biological Station, leg. Ramírez. Glossurella. Euglossa sapphirina [Euglossa (Euglossa)] Moure, 1968: 38–40, pl. 1, image: Nemésio, 2009a: 75b, d, f, Holotype ♂, USNM 70787: PANAMA, Colón, Gamboa, leg. Dressler. Glossurella. Euglossa securigera Dressler, 1982c: 145, fig. 1f, image: Nemésio, 2009a: 147b–d, Holotype ♂, USNM: BRAZIL, Espírito Santo, Conceição da Barra, leg. Dressler. Euglossa. Euglossa singularis Mocsáry, 1899: 169, Holotype ♀, HNHM: SURINAME. Euglossella. apiformis Schrottky, 1911: 39, Syntype ♀, ?: PERU, Cusco, leg. Garlepp. Euglossella. Euglossa solangeae Nemésio, 2007: 25–27, figs. 2a–e, 2g, image: Nemésio, 2009a: 81m, n, o, 149a–d, Holotype



♂, UFMG: BRAZIL, São Paulo, Cananéia, Ilha do Cardoso, leg. Augusto. Unplaced.

Euglossa sovietica [Euglossa (Euglossa)] Nemésio, 2007: 27–30, figs. 3a, 3c, 3e, 3g, Holotype ♂, UFMG: BRAZIL, Acre, Mâncio Lima, Parque Nacional da Serra do Divisor, leg. Morato. Euglossa. Euglossa stellfeldi Moure, 1947: 11–14, image: Nemésio, 2009a: 81g, h, i, 152a–d, Holotype ♀, MNCE: BRAZIL, Paraná, Caiobá. Glossura. annectans [Euglossa (Glossura)] Dressler, 1982a: 127–128, fig. 3a, image: Nemésio, 2009a: 82m, n, o, 151b– d, Holotype ♂, USNM: BRAZIL, Rio de Janeiro, Rio de Janeiro, Parque Nacional da Floresta da Tijuca, leg. Dressler: Not synonymized in Moure et al. (2007). Glossura. Euglossa stilbonota Dressler, 1982b: 138–139, fig. 1i, Holotype ♂, USNM: FRENCH GUIANA, Kourou, 19 km SW Kourou, leg. Roubik. Glossurella. Euglossa tiputini [Euglossa (Glossura)] Roubik, 2004: 241–244, fig. 9, Holotype ♂, USNM: ECUADOR, Napo, Yasuní National Park, leg. Roubik. Glossura. Euglossa townsendi Cockerell, 1904: 24, image: Nemésio, 2009a: 84a, c, e, g, Lectotype ♀, designated by Moure, 1967b: 405, AMNH: MEXICO, Veracruz, San Rafael (ex. nest), leg. Townsend: Cockerell did not follow his usual practice of labeling a single specimen as the type and remaining specimens as co–types. All specimens in this type series were labeled types and should be considered syntypes of which a lectotype has been selected (incl. also paralectotype specimen BMNH 17b 943). Euglossa. Euglossa tridentata [Euglossa (Euglossella)] Moure, 1970: 152–155, figs. 1d, 2d, Holotype ♂, USNM 70789: PANAMA, Colón, Summit Gardens, leg. Dressler. Euglossa. Euglossa trinotata Dressler, 1982b: 139–140, fig. 1j, Holotype ♂, USNM: COLOMBIA, Valle del Cauca, Buenaventura, Pulpapel, leg. Kennedy. Glossurella. Euglossa truncata [Euglossa (Euglossa)] Rebêlo & Moure, 1996 (“1995”): 457–459, figs. 2c, 4c, 6c, image: Nemésio, 2009a: 153a–d, Holotype ♂, RPSP: BRAZIL, São Paulo, Cajuru, Fazenda Santa Carlota, leg. Rebêlo. Euglossa. Euglossa turbinifex [Euglossa (Glossura)] Dressler, 1978a: 168–169, figs. 2a, 3a, Holotype ♂, USNM: PANAMA, Colón, Santa Rita ridge, leg. Dressler. Glossurella. Euglossa ultima [Euglossa (Euglossa)] Moure, 1968: 61–64, Holotype ♂, HNHM: VENEZUELA, Mérida. Euglossa. Euglossa urarina [Euglossa (Euglossella)] Hinojosa-Díaz & Engel, 2007: 100–103, figs. 13–22, Holotype ♂, UF: PERU, Loreto, Iquitos, leg. Dodson. Euglossella. Euglossa variabilis [Euglossa (Euglossa)] Friese, 1899: 135, Lectotype ♂, designated by Moure, 1967b: 406, ZMHB: FRENCH GUIANA, Cayenne. Euglossa. Euglossa villosa [Euglossa (Euglossa)] Moure, 1968: 17–20, Holotype ♂, USNM 70790: PANAMA, Coclé, El Valle de Antón, leg. Dressler. Dasytilbe. Euglossa villosiventris [Euglossa (Euglossa)] Moure, 1968: 20–23, Holotype ♂, USNM 70791: PANAMA, Panama, Cerro Jefe, leg. Dressler. Euglossa. Euglossa violascens Dominique, 1898: 58, Syntype unknown sex, ?: FRENCH GUIANA, Saint–Laurent du Maroni, Íle Portal, leg. Constant Bar and family. Unplaced. Euglossa viridifrons Dressler, 1982b: 140, fig. 1k, Holotype ♂, USNM: BRAZIL, Pará, Belém, leg. Dressler: Placed in the subgenus Glossura by Moure et al. (2007). Glossura. Euglossa viridis [Cnemidium] (Perty, 1833: 149, pl. 28, fig. 9), image: Nemésio, 2009a: 33 (org. plate), 155a–c, Holotype ♂, ZSM: BRAZIL, Amazonas, leg. Spix et al. Euglossella. affinis Dominique, 1898: 58, Lectotype ♂, designated by Rasmussen et al., 2007: 60, MHNAN: FRENCH GUIANA, Saint–Laurent du Maroni, Íle Portal, leg. Constant Bar and family. Euglossella. azurea Ducke, 1902a: 401, 402, Lectotype ♀, designated by Moure, 1967b: 401, MPEG: BRAZIL, Amapá, Macapá: The nomen appeared first both in a key and accompanied by a brief description (October 1st), thus giving this paper priority over the actual description, Ducke, 1902b: 565, 570 (December). Euglossella. Euglossa viridissima [Euglossa (Euglossa)] Friese, 1899: 136, Lectotype ♂, Moure, 1967b: 406, NMW: MEXICO. Euglossa.



29

Genus Eulaema Lepeletier de Saint Fargeau, 1841 Eulaema Lepeletier de Saint Fargeau, 1841: 11 Type species: Apis dimidiata Fabricius, 1793, by designation of Smith, 1874: 441. Eulaenia Spinola, 1851: 167 Lapsus for Eulaema Lepeletier de Saint Fargeau, 1841. Eulema Smith, 1854: 380 unjustified emendation of Eulaema Lepeletier de Saint Fargeau, 1841. Eulaima Dominique, 1898: 59 Lapsus for Eulaema Lepeletier de Saint Fargeau, 1841.

Two subgenera are recognized in Eulaema Eulaema Lepeletier de Saint Fargeau, 1841: 11. Apeulaema Moure, 1950: 184–186. Type species: Eulaema fasciata Lepeletier de Saint Fargeau, 1841, by original designation. Eulaema atleticana Nemésio, 2009a: 185–187, figs. 184a–d, Holotype ♂, UFMG: BRAZIL, Bahia, Valença, Cajaíba, leg. Neves. Eulaema. Eulaema basicincta [Eulaema (Eulaema)] Moure, 2003 (“2000”): 44–45, pl. 4, fig. 16, Holotype ♂, DZUP: TRINIDAD & TOBAGO, Aripo Valley, leg. Bennett. Eulaema. Eulaema bennetti [Eulaema (Eulaema)] Moure, 1967a: 384–388, Holotype ♂, ?; CIBC collection transferred to CABI Caribbean and Latin America, but there is no holotype: TRINIDAD & TOBAGO, Aripo Valley, leg. Bennett. Eulaema. Eulaema boliviensis [Eulema!] Friese, 1898: 205–206, Lectotype ♀, designated by Moure, 1967b: 411, ZMHB: BOLIVIA: Purchased through Dr. O. Staudinger's insect dealership, Dresden, Germany. Apeulaema. Eulaema bombiformis [Euglossa] (Packard, 1869: 57–58), Holotype ♀, MCZ: ECUADOR, Pichincha–Napo, Quito (probably en route to Napo river), leg. Smithsonian Expedition/Orton. Eulaema. Eulaema bomboides [Euglossa (Eulaema)] (Friese, 1923: 28), Lectotype ♀, designated by Moure, 1967b: 411, AMNH (type status unconfirmed): ECUADOR, Bolívar, Balsapampa (Balzapampa). Eulaema. Eulaema chocoana Ospina-Torres & Sandino-Franco, 1997: 166–172, figs. 1, 2, 3a, 4a, 5, 6a, 7, 8, Holotype ♂, ICN (Laboratorio de Investigaciones en Abejas): COLOMBIA, Nariño, Barbacoas, Pueblo Nuevo, Río Malaunde chiquito, leg. Sandino. Eulaema. Eulaema cingulata [Centris] (Fabricius, 1804: 355), image: Nemésio, 2009a: 159, 163a–d, Lectotype ♀, designated by Moure, 1960: 145, ZMUC: “America meridionali” (possibly GUYANA). Apeulaema. cajennensis Lepeletier de Saint Fargeau, 1841: 14, Lectotype ♂, designated by Moure, 1967b: 411 (cf. ICZN Art. 74.6), ?: FRENCH GUIANA, Cayenne. Apeulaema. fasciata Lepeletier de Saint Fargeau, 1841: 12, image: Nemésio, 2009a: 165a–d, Lectotype ♀, designated by Moure, 1967b: 411, MNHN: FRENCH GUIANA, Cayenne. Apeulaema. pseudocingulata Oliveira, 2006: 122–124, fig. 1f, image: Nemésio, 2009a: 162a–d, Holotype ♂, RPSP: BRAZIL, Amazonas, Arumã, leg. Camargo & Mazucato. Apeulaema. Eulaema felipei Nemésio, 2010: 54–56, figs. 3a–3d, Holotype ♂, UFMG: BRAZIL, Alagoas, Murici, leg. Nemésio. Apeulaema. Eulaema flavescens [Euglossa (Eulema!) dimidiata ssp.] (Friese, 1899: 165), image: Nemésio, 2009a: 174, 175, 180a–d, Lectotype ♀, designated by Moure, 1967b: 411, NMW: VENEZUELA. Eulaema. Eulaema helvola [Eulaema (Eulaema)] Moure, 2003 (“2000”): 60–61, pl. 1, fig. 4, image: Nemésio, 2009a: 185a– f, Holotype ♂, DZUP: BRAZIL, Goiás, Goiânia. Eulaema. Eulaema leucopyga [Eulema!] Friese, 1898: 203, Holotype ♀, ZMHB: COLOMBIA. Eulaema. Eulaema luteola [Eulaema (Eulaema)] Moure, 1967a: 375–378, Holotype ♀, DZUP: COLOMBIA, Valle del Cauca, Monte Redondo, leg. Foerster. Eulaema. Eulaema marcii Nemésio, 2009a: 175–177, figs. 170a–d, Holotype ♂, UFMG: BRAZIL, Minas Gerais, Marliéria, Parque Estadual do Rio Doce, leg. Nemésio. Apeulaema. Eulaema meriana [Apis] (Olivier, 1789: 64), Neotype ♂, designated in present work, UFMG: BRAZIL, Pará, Oriximiná, leg. Martines. Eulaema. dimidiata [Apis] (Fabricius, 1793: 316), Neotype ♂, designated in present work, UFMG: BRAZIL, Pará, Orix



iminá, leg. Martines. Eulaema. quadrifasciata [Euglossa (Eulema!) dimidiata ssp.] (Friese, 1903: 575), Lectotype ♂, designated by Moure, 1967b: 412, ZMHB: COSTA RICA, San José, San Carlos, leg. Burgdorf: Not synonymized in Moure et al. (2007). Eulaema. pallescens [Eulaema (Eulaema)] Moure, 2003 (“2000”): 35–36, pl. 5, fig. 20, Holotype ♂, DZUP: ECUADOR, Esmeraldas, San Mateo: Not synonymized in Moure et al. (2007). Eulaema. Eulaema mimetica [Eulaema (Eulaema)] Moure, 1967a: 379–384, Holotype ♂, DZUP: BRAZIL, Pará, Óbidos, Curusamba, leg. Brazilino. Eulaema. Eulaema mocsaryi [Euglossa (Euglossa) mocsáryi] (Friese, 1899: 161–162), Lectotype ♀, designated by Moure, 1967b: 411 (cf. ICZN Art. 74.6), OUMNH: BRAZIL, Pará: The female paralectotype of fallax Smith is the lectotype by indication of Friese and later Moure. Apeulaema. Eulaema napensis [Eulaema (Eulaema)] Oliveira, 2006: 125, fig. 1e, Holotype ♂, QCAZ: ECUADOR, Francisco de Orellana, Archidona, Jumandi, leg. Sanchez. Eulaema. Eulaema nigrifacies [Eulema! surinamensis ssp.] Friese, 1898: 205, Lectotype ♂, designated by Moure, 1967b: 412 (cf. ICZN Art. 74.6), ZMHB: VENEZUELA, Bolívar, Sierra Parima (as St. Parime on label, see Moure, 1967b: 412). Eulaema. panamensis [Euglossa (Eulema!)] (Mocsáry, 1899: 169–170), Holotype ♀, HNHM: PANAMA, Chiriquí. Eulaema. sarapiquiensis [Euglossa (Eulema!) nigrifacies ssp.] (Friese, 1925: 30), Lectotype ♀, designated by Moure, 1967b: 412 (cf. ICZN Art. 74.6), ZMHB: COSTA RICA, Heredia, Puerto Viejo de Sarapiquí, leg. Schmidt (Albert). Eulaema. Eulaema nigrita Lepeletier de Saint Fargeau, 1841: 14, Lectotype ♀, designated by Moure, 1967b: 412, nomen protectum, ?: FRENCH GUIANA, Cayenne. Apeulaema. analis Lepeletier de Saint Fargeau, 1841: 14–15, Syntype ♂ nomen oblitum, MNHN?: BRAZIL. Apeulaema. raymondi [Centris nigrita ssp.] (Schrottky, 1907: 65), image: Nemésio, 2009a: 22a–f, Lectotype ♀, designated by Moure, 1967b: 412, MPSP: VENEZUELA, Miranda, Caracas, leg. Raymond. Apeulaema. nigriceps [Euglossa (Eulaema) nigrita ssp.] (Friese, 1923: 27), Lectotype ♂, designated by Moure, 1967b: 411–412, ZMHB: COLOMBIA, Valle del Cauca, Cauca–Tal, near Popayán, 1000 m, leg. Fassl. Apeulaema. willei [Eulaema (Apeulaema)] Moure, 1963: 213–215, Holotype ♂, ? (not in MUCR, MZCR): COSTA RICA, Guanacaste, 14 km S of Cañas, Río Blanco, leg. Wille. Apeulaema. Eulaema niveofasciata [Euglossa (Eulema!) dimidiata ssp.] (Friese, 1899: 165), image: Nemésio, 2009a: 187, 189a–d, Lectotype ♀, designated by Moure, 1967b: 412, NMW: BRAZIL, Pernambuco. Eulaema. Eulaema parapolyzona [Eulaema (Eulaema)] Oliveira, 2006: 124–125, fig. 1h, Holotype ♂, SEMC: BOLIVIA, Cochabamba, Chapare, 400 m, leg. Zischaka. Eulaema. Eulaema peruviana [Euglossa (Eulema!)] (Friese, 1903: 575), Lectotype ♀, designated by Moure, 1967b: 412, HNHM: PERU, Cusco, Marcapata. Eulaema. Eulaema polychroma [Euglossa (Eulema!)] (Mocsáry, 1899: 170), Holotype ♂, HNHM: PERU, Cusco, Callanga. Apeulaema. Eulaema polyzona [Euglossa (Eulema!)] (Mocsáry, 1897: 442–443), Lectotype ♀, designated by Moure, 1967b: 412, HNHM: SURINAME. Eulaema. difficilis [Eulema!] Friese, 1898: 206, Lectotype ♀, designated by Moure, 1967b: 411, ZMHB: BRAZIL, Pará, Belém. Eulaema. maroniensis [Eulaima] Dominique, 1898: 59, Lectotype ♂, designated by Rasmussen et al., 2007: 61, MHNAN: FRENCH GUIANA, Saint–Laurent du Maroni, Íle Portal, leg. Constant Bar and family. Eulaema. Eulaema seabrai [Eulaema (Eulaema)] Moure, 1960: 19–22, image: Nemésio, 2009a: 190, 192a–d, Holotype ♀, DZUP: BRAZIL, Rio de Janeiro, Alto da Boa Vista, Tijuca, leg. Seabra. Eulaema. Eulaema sororia Dressler & Ospina-Torres, 1997: 95–100, figs. 1, 2.1, 3, 4, 5, 6, 7, 8, Holotype ♂, SEMC: ECUADOR, Esmeraldas, Pichincha, W of Lita, leg. Dressler et al. Eulaema. Eulaema speciosa [Euglossa (Eulema!)] (Mocsáry, 1897: 445–446), Holotype ♀, HNHM: PANAMA, Chiriquí. Eulaema.



31

semirufa [Eulema!] Friese, 1898: 204–205, Holotype ♂, ZMHB: PANAMA, Chiriquí. Eulaema. Eulaema tenuifasciata [Euglossa dimidiata ssp.] (Friese, 1925: 30), Syntype ♀, AMNH (type status unconfirmed): BRAZIL, Amazonas: Although Friese only listed Amazonas, the type locality is likely Brazil. Eulaema. Eulaema terminata [Eulema!] Smith, 1874: 442, Holotype ♂, BMNH 17b951: TRINIDAD & TOBAGO. Eulaema. stenozona [Eulaema (Eulaema)] Moure, 2003 (“2000”): 38, pl. 3, figs. 10, 12, Holotype ♂, DZUP: TRINIDAD & TOBAGO, Aripo Valley, leg. Bennett: Not synonymized in Moure et al. (2007). Eulaema.

Genus Exaerete Hoffmannsegg, 1817 Exaerete Hoffmannsegg, 1817: 53. Type species: Apis dentata Linnaeus, 1758, monobasic. Chrysantheda Perty, 1833: 147–148. Type species: Chrysantheda nitida Perty, 1833, monobasic. Caliendra Gistel, 1848: viii, unjustified replacement for Chrysantheda Perty, 1833. Type species: Chrysantheda nitida Perty, 1833, autobasic (see Chrysantheda Perty, 1833).

Exaerete azteca Moure, 1964: 15–16, Holotype ♂, SEMC: MEXICO, Hidalgo, 38 miles NE of Jacala, 3100 ft., leg. University of Kansas Mexican Expedition. Exaerete dentata [Apis] (Linnaeus, 1758: 575), image: Nemésio, 2009a: 195, 196, 198a–f, Holotype ♀, UUZM: “Indiis” (possibly WEST INDIES or SURINAME). nitida [Chrysantheda] (Perty, 1833: 148, pl. 28, fig. 8), image: Nemésio, 2009a: 33 (org. plate), 199a–f, Holotype ♀, ZSM: BRAZIL, Piauí, leg. Spix et al. subcornuta [Chrysantheda] (Romand, 1849: xxxvi–xxxviii), Holotype ♂, ?: VENEZUELA, Miranda, Caracas, leg. Sallé: Published after September 12, 1849. appendiculata [Chrysantheda] (Romand, 1849: xxxvi, pl. 7, fig. Ia–c), Holotype ♂, see subcornuta: VENEZUELA, Miranda, Caracas, leg. Sallé: Proposed as a junior synonym of subcornuta, based on the same specimen; Published after September 12, 1849. Exaerete frontalis [Euglossa] (Guérin-Méneville, 1844: 458), image: Nemésio, 2009a: 194b, d, f, 200, 201a–f, Neotype ♂ (lost holotype was ♀), Nemésio, 2009a: 194, figs. 194b, 194d, 194f, UFMG: BRAZIL, Pará, Oriximiná, Porto Trombetas, leg. Martines. lucida Erichson, 1849 (“1848”): 592, Holotype ♂, ZMHB (only pin and labels left): GUYANA, leg. Schomburgk et al.: Possibly syntypes; published March 10, 1849 and not 1848. Exaerete kimseyae Oliveira, 2011: 2–4, figs. 1–7 [figs. of male genitalia of Ex. trochanterica in Kimsey (1979) should also be considered as figs. of holotype Ex. kimseyae, the only known specimen], Holotype ♂, UCDC: PANAMA, Colón, Barro Colorado Island, leg. Kimsey. Exaerete lepeletieri Oliveira & Nemésio, 2003: 117–119, figs. 2, 4, 5, 6, image: Nemésio, 2009a: 194a, c, e, Holotype ♂, INPA: BRAZIL, Acre, Rio Branco, leg. Oliveira. Exaerete salsai Nemésio, 2011c: 14–15, figs. 2–3, Holotype ♂, UFMG: BRAZIL, Bahia, Porto Seguro, leg. Nemésio. Exaerete smaragdina [Euglossa] (Guérin-Méneville, 1844: 458), image: Nemésio, 2009a: 193a–b, 202, 204a–h, 206, Lectotype ♀, designated by Moure, 1967b: 414, MSNG: MEXICO, Yucatán, Campeche. aurata Erichson, 1849 (“1848”): 592, Holotype ♀, ZMHB: GUYANA, leg. Schomburgk et al.: Possibly syntypes; published March 10, 1849 and not 1848. cyanescens [Exaerete smaragdina ssp.] Cockerell, 1926: 657, image: Nemésio, 2009a: 205a–h, Holotype; invalid lectotype designation of holotype ♀, Moure, 1967b: 413, AMNH: TRINIDAD & TOBAGO, leg. Urich. Exaerete trochanterica [Chrysantheda] (Friese, 1900: 66–67), Holotype ♀, ZMHB: BRAZIL, Pará, Belém, leg. Schulz. guaykuru Anjos-Silva & Rebêlo, 2006: 29–33, figs. 1, 2, 7, 12, 17, 22, 23, Holotype ♂, MPSP: BRAZIL, Mato Grosso, Chapada dos Guimarães, 600 m, leg. Anjos–Silva.

Acknowledgments We are indebted to the many curators who replied to our requests and to Bart de Dijn and Remko Leijs, who helped us with translation of Merian’s work and also commented on the Eulaema meriana taxonomic problem. We also



thank Eduardo A. B. Almeida, Gerardo Lamas, and David Notton for reading and commenting on earlier versions (or parts) of this manuscript. Robert L. Dressler and G. Gerlach kindly discussed on the pollinarium of Exaerete subcornuta. Besides the members of ICZN Secretariat mentioned in the text under the section on Euglossa fimbriata, the following zoologists were kind enough to share their views with us on this matter: Alan L. Melo, Angelico Asenjo, Carlos Peña, Eduardo A. B. Almeida, Fernando A. Silveira, James Pitts, José P. Pombal Jr., Lúcio A. O. Campos, Mauro L. Triques, Miguel A. Monné, and Victor H. Gonzalez Betancourt. Fabio Penati kindly photographed the lectotype Euglossa piliventris here illustrated as Figs. 5 and 6. We are also grateful to David W. Roubik (Smithsonian Tropical Research Institute, Balboa, Ancón, Panama) and Eduardo A. B. Almeida (Universidade de São Paulo, Ribeirão Preto, SP, Brazil) as reviewers for useful comments and criticisms on this paper. Information attributed to D.W. Roubik both in the text and in the catalogue above was retrieved from his review. We especially thank Arkady Lelej, one of Zootaxa editors for Hymenoptera, who carefully edited this manuscript (the editor for Apoidea was not eligible since he co-authors this manuscript).

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APPENDIX 1. Valid species of orchid bees (genera Euglossa and Eulaema) placed in subgenera Euglossa (Dasytilbe): Euglossa obrima; Eg. villosa. Euglossa (Euglossa): Euglossa alleni; Eg. amazonica; Eg. analis; Eg. anodorhynchi; Eg. aratingae; Eg. atroveneta; Eg. aureiventris; Eg. auriventris; Eg. avicula; Eg. bembei; Eg. bidentata; Eg. carolina; Eg. championi; Eg. charapensis; Eg. chlorina; Eg. cognata; Eg. cordata; Eg. cotylisca; Eg. crininota; Eg. cyanaspis; Eg. cybelia; Eg. deceptrix; Eg. despecta; Eg. dissimula; Eg. dressleri; Eg. erythrochlora; Eg. fimbriata; Eg. gibbosa; Eg. hansoni; Eg. hemichlora; Eg. heterosticta; Eg. igniventris; Eg. ioprosopa; Eg. iopyrrha; Eg. jamaicensis; Eg. laurensi; Eg. leucotricha; Eg. liopoda; Eg. maculilabris; Eg. magnipes; Eg. marianae; Eg. melanotricha; Eg. micans; Eg. milenae; Eg. mixta; Eg. modestior; Eg. mourei; Eg. nanomelanotricha; Eg. nigropilosa; Eg. pictipennis; Eg. platymera; Eg. pleosticta; Eg. purpurea; Eg. retroviridis; Eg. roderici; Eg. securigera; Eg. sovietica; Eg. townsendi; Eg. tridentata; Eg. truncata; Eg. ultima; Eg. variabilis; Eg. villosiventris; Eg. viridissima. Euglossa (Euglossella): Euglossa bigibba; Eg. cosmodora; Eg. cyanea; Eg. cyanura; Eg. decorata; Eg. granti; Eg. jacquelynae; Eg. mandibularis; Eg. perfulgens; Eg. perpulchra; Eg. perviridis; Eg. singularis; Eg. urarina; Eg. viridis. Euglossa (Glossura): Euglossa allosticta; Eg. asarophora; Eg. chalybeata; Eg. flammea; Eg. ignita; Eg. imperialis; Eg. iopoecila; Eg. lugubris; Eg. natesi; Eg. occidentalis; Eg. orellana; Eg. piliventris; Eg. roubiki; Eg. rufipes; Eg. stellfeldi; Eg. tiputini; Eg. viridifrons. Euglossa (Glossurella): Euglossa augaspis; Eg. bursigera; Eg. carinilabris; Eg. crassipunctata; Eg. dodsoni; Eg. erythrophana; Eg. fuscifrons; Eg. gorgonensis; Eg. hyacinthina; Eg. laevicincta; Eg. macrorhyncha; Eg. obtusa; Eg. oleolucens; Eg. paisa; Eg. parvula; Eg. prasina; Eg. samperi; Eg. sapphirina; Eg. solangeae; Eg. stilbonota; Eg. trinotata; Eg. turbinifex. Euglossa (Glossuropoda): Euglossa cyanochlora; Eg. inflata; Eg. intersecta; Eg. juremae; Eg. nigrosignata; Eg. rugilabris. Euglossa (unplaced): Euglossa lazulina; Eg. moronei; Eg. polita; Eg. solangeae; Eg. violascens. Eulaema (Apeulaema): Eulaema boliviensis; El. cingulata; El. felipei; El. marcii; El. mocsaryi; El. nigrita; El. polychroma. Eulaema (Eulaema): Eulaema atleticana; El. basicincta; El. bennetti; El. bombiformis; El. bomboides; El. chocoana; El. flavescens; El. helvola; El. leucopyga; El. luteola; El. meriana; El. mimetica; El. napensis; El. nigrifacies; El. niveofasciata; El. parapolyzona; El. peruviana; El. polyzona; El. seabrai; El. sororia; El. speciosa; El. tenuifasciata; El. terminata.



39

Index of Latin names The valid species-names (i.e., specific epithets) are not italicized, the synonyms and unavailable names in italic.

abdomenflavum, see surinamensis, Eufriesea aeneiventris, Eufriesea aenescens, see mandibularis, Euglossa affinis, see viridis, Euglossa Aglae alleni, Euglossa allosticta, Euglossa amabilis, see surinamensis, Eufriesea amazonica, Euglossa analis, see nigrita, Eulaema analis, Euglossa andina, Eufriesea angulata, see surinamensis, Eufriesea anisochlora, Eufriesea annectans, see stellfeldi, Euglossa anodorhynchi, Euglossa apiformis, see singularis, Euglossa appendiculata, see dentata, Exaerete aratingae, Euglossa aridicola, see auriceps, Eufriesea asarophora, Euglossa atlantica, Eufriesea atleticana, Eulaema atroveneta, Euglossa augaspis, Euglossa aurata, see smaragdina, Exaerete aureiventris, Euglossa auriceps, Eufriesea auricollis, see hemichlora, Euglossa auripes, Eufriesea auriventris, Euglossa avicula, Euglossa azteca, Exaerete azurea, see viridis, Euglossa azureoviridis, see hemichlora, Euglossa bare, Eufriesea bari, see ignita, Euglossa basalis, see limbata, Eufriesea basicincta, Eulaema bembei, Euglossa bennetti, Eulaema bernardina, see mandibularis, Euglossa bicolor, see analis, Euglossa bidentata, Euglossa bigibba, Euglossa boharti, Eufriesea boliviensis, Eulaema bombiformis, Eulaema bomboides, Eulaema brasilianorum, Eufriesea bruesi, see surinamensis, Eufriesea brullei, see intersecta, Euglossa buchwaldi, Eufriesea


bureaui, see cognata, Euglossa bursigera, Euglossa cajennensis, see cingulata, Eulaema Caliendra, see Exaerete carinilabris, Euglossa carolina, Euglossa chaconi, Eufriesea chalybaea, Eufriesea chalybeata, Euglossa championi, Euglossa charapensis, Euglossa chlorina, Euglossa chlorosoma, see ignita, Euglossa chocoana, Eulaema Chrysantheda, see Exaerete chrysopyga, Eufriesea cingulata, Eulaema Cnemidium, see Euglossa cognata, Euglossa coerulea, Aglae coerulescens, Eufriesea combinata, Eufriesea concava, Eufriesea convexa, Eufriesea cordata, Euglossa corusca, Eufriesea cosmodora, Euglossa cotylisca, Euglossa crassipunctata, Euglossa crininota, Euglossa cupreicolor, see bursigera, Euglossa cupreiventris, see igniventris, Euglossa cyanaspis, Euglossa cyanea, Euglossa cyanescens, see smaragdina, Exaerete cyanochlora, Euglossa cyanura, Euglossa cybelia, Euglossa danielis, see auriceps, Eufriesea Dasystilbe deceptrix, Euglossa decorata, Euglossa dentata, Exaerete dentilabris, Eufriesea despecta, Euglossa difficilis, see polyzona, Eulaema dimidiata, see meriana, Eulaema dissimula, Euglossa distinguenda, Eufriesea dodsoni, Euglossa dressleri, Eufriesea dressleri, Euglossa


duckei, Eufriesea eburneocincta, Eufriesea elegans, Eufriesea erythrochlora, Euglossa erythrophana, Euglossa Eufriesea Euglossa Euglossella Eulaema Exaerete excellens, Eufriesea faceta, see nigrohirta, Eufriesea fallax, Eufriesea fasciata, see cingulata, Eulaema felipei, Eulaema fimbriata, Euglossa flammea, Euglossa flavescens, Eulaema flaviventris, Eufriesea formosa, Eufriesea fragrocara, Eufriesea frontalis, Exaerete fulvohirta, see magrettii, Eufriesea fuscatra, Eufriesea fuscifrons, Euglossa gaianii, see hemichlora, Euglossa gibbosa, Euglossa Glossura Glossurella Glossuropoda gorgonensis, Euglossa granti, Euglossa guaykuru, see trochanterica, Exaerete hansoni, Euglossa heideri, Eufriesea helvola, Eulaema hemichlora, Euglossa heterosticta, Euglossa hugonis, see rugilabris, Euglossa hyacinthina, Euglossa ignita, Euglossa igniventris, Euglossa imperialis, Euglossa inermis, see mussitans, Eufriesea inflata, Euglossa intersecta, Euglossa iopoecila, Euglossa ioprosopa, Euglossa iopyrrha, Euglossa jacquelynae, Euglossa jamaicensis, Euglossa juremae, Euglossa kimimari, Eufriesea kimseyae, Exaerete laevicincta, Euglossa laniventris, Eufriesea laurensi, Euglossa


lazulina, Euglossa lepeletieri, Exaerete leucopyga, Eulaema leucotricha, Euglossa limbata, Eufriesea liopoda, Euglossa longipennis, Eufriesea lucida, see frontalis, Exaerete lucida, Eufriesea lucifera, Eufriesea lugubris, Euglossa luteola, Eulaema macroglossa, Eufriesea macrorhyncha, Euglossa maculilabris, Euglossa magnipes, Euglossa magrettii, Eufriesea mandibularis, Euglossa manni, see superba, Eufriesea marcii, Eulaema mariana, Eufriesea marianae, Euglossa maroniensis, see polyzona, Eulaema melanotricha, Euglossa meliponoides, Euglossa melissiflora, Eufriesea meriana, Eulaema mexicana, Eufriesea micans, Euglossa micheneri, Eufriesea milenae, Euglossa mimetica, Eulaema mixta, Euglossa mocsaryi, Eulaema modestior, Euglossa moronei, Euglossa mourei, Euglossa mussitans, Eufriesea nanomelanotricha, Euglossa napensis, Eulaema natesi, Euglossa nigrescens, seenigrescens, Eufriesea nigriceps, see nigrita, Eulaema nigrifacies, Eulaema nigrita, see nigrescens, Eufriesea nigrita, Eulaema nigrohirta, Eufriesea nigropilosa, Euglossa nigrosignata, Euglossa nitida, see dentata, Exaerete niveofasciata, Eulaema nordestina, see auriceps, Eufriesea obrima, Euglossa obtusa, Euglossa hemichlora, Euglossa occidentalis, Euglossa oleolucens, Euglossa


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opulenta, Eufriesea orellana, Euglossa ornata, Eufriesea paisa, Euglossa pallescens, see meriana, Eulaema pallida, Eufriesea panamensis, see nigrifacies, Eulaema parapolyzona, Eulaema parvula, Euglossa perfulgens, Euglossa perpulchra, Euglossa peruviana, Eulaema perviridis, Euglossa pictipennis, Euglossa piliventris, Euglossa platymera, Euglossa pleosticta, Euglossa polita, Euglossa polychroma, Eulaema polyzona, Eulaema prasina, Euglossa pretiosa, Eufriesea pseudocingulata, see cingulata, Eulaema pulcherrima, see mariana, Eufriesea pulchra, Eufriesea purpurata, Eufriesea purpurea, Euglossa pyrrhopyga, Eufriesea quadrifasciata, see meriana, Eulaema raymondi, see nigrita, Eulaema retroviridis, Euglossa roderici, Euglossa romandii, see intersecta, Euglossa roubiki, Euglossa ruficauda, see decorata, Euglossa rufipes, Euglossa rufocauda, Eufriesea rugilabris, Euglossa rugosa, Eufriesea salsai, Exaerete samperi, Euglossa sapphirina, Euglossa sarapiquiensis, see nigrifacies, Eulaema schmidtiana, Eufriesea seabrai, Eulaema securigera, Euglossa semirufa, see speciosa, Eulaema


simillima, see coerulescens, Eufriesea singularis, Euglossa smaragdina, Eufriesea smaragdina, Exaerete solangeae, Euglossa sororia, Eulaema sovietica, Euglossa speciosa, Eulaema stellfeldi, Euglossa stenozona, see terminata, Eulaema stilbonota, Euglossa subcornuta, see dentata, Exaerete superba, see mariana, Eufriesea superba, Eufriesea surinamensis, Eufriesea tectora, see surinamensis, Eufriesea tenuifasciata, Eulaema terminata, Eulaema theresiae, Eufriesea tiputini, Euglossa townsendi, Euglossa tridentata, Euglossa trinotata, Euglossa trochanterica, Exaerete tropica, see surinamensis, Eufriesea truncata, Euglossa tucumana, see mariana, Eufriesea turbinifex, Euglossa ultima, Euglossa urarina, Euglossa variabilis, Euglossa velutina, Eufriesea venezolana, Eufriesea venusta, Eufriesea vidua, Eufriesea villosa, Euglossa villosiventris, Euglossa violacea, Eufriesea violaceifrons, see despecta, Euglossa violascens, Eufriesea violascens, Euglossa viridifrons, Euglossa viridis, Euglossa viridissima, Euglossa willei, see nigrita, Eulaema xantha, see vidua, Eufriesea
