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ZOOTAXA
Catalog of the Indo-Malayan/Australasian stingless bees (Hymenoptera: Apidae: Meliponini)
CLAUS RASMUSSEN
Magnolia PressAuckland, New Zealand
1935
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ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
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Accepted by E. Almeida: 16 Oct 2008; published: 17 Nov. 2008 3
Catalog of the Indo-Malayan/Australasian stingless bees (Hymenoptera: Apidae: Meliponini)
CLAUS RASMUSSENDepartment of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, USA. E-mail: [email protected]
Table of contents
Abstract ............................................................................................................................................................................... 4Catalog format ..................................................................................................................................................................... 4Taxonomy and classification ............................................................................................................................................... 5Distribution and biology ..................................................................................................................................................... 6Acknowledgments ............................................................................................................................................................... 7Catalog of the Indo-Malayan/Australasian stingless bees .................................................................................................. 8
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This catalog of the published literature on stingless bees from the Indo-Malayan/Australasian region was compiled from520 references and deals with 129 proposed names for species-group taxa of which 89 are considered currently valid.Information is summarized on organisms associated with stingless bee colonies as well as the 225 plants from which thebees have been reported to forage. Lectotypes are designated for Trigona ornata described by Rayment and the followingspecies described by Friese: Trigona anamitica, T. australis, T. borneënsis, T. flaviventris, T. keyensis, T. luteiventris, T.pygmaea, T. reepeni, T. sericea, and T. versicolor.
Stingless bees (Hymenoptera: Apidae: Meliponini) are important pollinators of native plants in tropical andsubtropical parts of the world (Heard 1999a). They form large perennial colonies with complex social organi-zation (Michener 2007a), have a diverse nesting biology ranging from underground to exposed colonies(Schwarz 1948; Wille & Michener 1973; Camargo & Pedro 2003b; Roubik 2006; Rasmussen & Camargo2008), and are abundantly encountered within their native range (Roubik 2006). In addition to a well-corrobo-rated phylogeny, an up-to-date, comprehensive taxonomic and biological catalog is fundamental to any com-parative evolutionary, ecological, and behavioral research on any group of organisms. A catalog provides anindex to previous studies in taxonomy, behavioral research, and pollination ecology, thus consolidating theexisting knowledge in an accessible format. The extensive Neotropical fauna was recently cataloged byCamargo & Pedro (2007: 391 spp.) and a systematic revision of the Afrotropical stingless bees included anearly complete bibliography (Eardley 2004: 26 spp. incl. Madagascar). The present catalog treats the remain-ing Indo-Malayan/Australasian stingless bees (129 proposed species-group names, 89 currently acceptedvalid species) and completes efforts to catalog the global stingless bee fauna. Together with recent molecularphylogenies of the stingless bees (Rasmussen & Cameron 2007; Rasmussen 2008), it is expected that thesetools will spark renewed scientific interest in this diverse group of bees.
Catalog format
The catalog is sorted alphabetically by genus-group and currently valid species-group names; subspecies arenot recognized in the catalog. Genus-group headings are in boldface type. Following the accepted species-group names are listed synonyms, including validly proposed junior synonyms and nomina nuda. Referencesare sorted in chronological order and include citation, page numbers from publications, type-specimen infor-mation from both type-specimen labels, and the original publication where the species was first described, keywords, distribution, and floral records. Incertae sedis and nomina nuda are listed at the end of the catalog,except when the identity is known for a nomen nudum. Page numbers are not listed for brief works or if theparticular species is treated throughout the publication. Publication dates were determined by the publishersimprint date on the wrapper of the fascicle, when available, or information contained within the issue. Whenthe imprint year of the issue differed from the stated year of the volume, the latter are placed in parentheses.Asterisks (*) following page numbers indicate where the species discussion is located in major works. Keywords in parenthesis after a reference indicate the kind of information provided and associated organismsrecorded. Misidentifications are corrected in the key word section whenever necessary or known, and typo-graphical errors in spelling of taxa names in publications are noted followed by an exclamation mark. Distri-butions are listed alphabetically with country names in upper case type and type localities indicated as in theoriginal publication. Distributional records are not listed comprehensively, but rather are limited to records
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cited in either taxonomic works or faunal inventories. Spellings of floral records were corrected to conformwith the International Plant Name Index (www.ipni.org). Appendix 1 lists all plant species (including family)upon which stingless bees are known to forage in the region. Plant synonymies were not updated, but ratherleft as cited in the original publication.
Stingless bee type specimens are encountered in museums worldwide, as indicated in the catalog. Acro-nyms for institutions are abbreviated following Evenhuis (2007) and Tadauchi et al. (1998) for Sakagami-related collections (Appendix 3). Curators who assisted with loans and important information on types arealso listed following each institution. Of particular importance are the collections in the Natural HistoryMuseum (London), with 52 primary types, Museum für Naturkunde der Humboldt-Universität (Berlin), with15, and National Museum of Natural History (Washington, D.C.), with 10. As for the most prolific describersof new taxa, Theodore D.A. Cockerell described 38 new taxa, Heinrich Friese 17, Frederick Smith 14, HerbertF. Schwarz and Peter Cameron 11 each, and Shôichi F. Sakagami 10. The whereabouts of three Sakagami pri-mary types remain unknown (i.e., Trigona gressitti, T. pagdeniformis, and T. lieftincki) as these have not beenfound in the collections indicated by the late Sakagami nor in his private collection (Tadauchi et al. 1998). Itmay be that the types are in those collections by Tadauchi et al. (1998), but that they are not labeled as types.The type of Trigona kusutkana Dover is missing from the Raffles Museum of Biodiversity Research where itwas originally deposited. All of the types of Smith have been accounted for, although as stressed by Baker(1993), several of the specimens labeled as types in the Natural History Museum (London) may not be truetypes and putative type specimens from London needs to be examined and compared with specimens fromUniversity Museum of Natural History (Oxford) when in doubt. A number of species described before thestandardization of taxonomy with the International Code of Zoological Nomenclature (latest edition is 1999)are represented by syntypes (multiple primary types). In several cases a syntype series consists of multiplebiological and morphological species. Lectotypes are here designated for several species proposed by Frieseas the identity of his true types is often confusing (see Rasmussen & Ascher 2008).
Literature has been updated until mid-2007. References cited include those found in Zoological Record(Zoological Society of London 1864–present) and all references cited within these records, totaling 520papers. A number of popular travel accounts are listed in the nearly complete stingless bees bibliography bySchwarz (1948). These were all consulted, but only included if they provided either the scientific name orsome description of the natural history of the bees, which was infrequently the case.
Taxonomy and classification
I have followed the classification proposed by Moure (1961) and Rasmussen & Cameron (2007) in not treat-ing some of the Old World stingless bees as subgenera of Trigona Jurine (Table 1). The phylogenetic groupsof Rasmussen & Cameron (2007) are fully congruent with the systematic arrangement proposed by Moure(1961). Therefore, all supraspecific groups are treated at the generic level as in the New World stingless beecatalog (Camargo & Pedro 2007). Other studies propose a simpler classification to avoid a profusion of for-mal names, which is equally valid when based on monophyletic lineages. This dual approach to classifyinggenera (chiefly Moure 1961; Michener 1990, 2007a), however, complicates literature searches with specifictaxa appearing under several different generic names. The genus-group name or synonymy is listed for eachtaxon in Appendix 2.
Most synonymies here are based on the published examination of types by Schwarz (1937; 1939a) andMoure (1961). However, a number of species were only tentatively synonymized by them, and re-examinationof primary types was necessary for confirming and establishing their synonymies. In those cases, I indicatedthe types I examined. Just like the inconsistent usage of stingless bee generic combinations, some taxa havebeen treated as subspecies (e.g., Schwarz 1939a; Sakagami et al. 1990), species (e.g., Klakasikorn et al.
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2005), or synonyms. I have therefore refrained from stating whether I provide a new combination (comb. nov.)or a new status (stat. nov.), as all taxa have been given full species status or presented in the current combina-tion in the references provided. I did, however, recognize all previously proposed subspecies and varieties(mainly from the works of Schwarz) at the specific level when considered a valid taxon. This also reflects thefact that about 80% of the New World taxa originally described by Schwarz as subspecies in light of addi-tional biological information are now regarded fully valid species (Camargo & Pedro 2007).
Identification of stingless bees from the Indo-Malay/Australasian region is best done with specializedkeys, although the fauna has not yet undergone the taxonomic scrutiny comparable to that of many NewWorld taxa (e.g., Camargo & Moure 1994; Camargo & Moure 1996; Pedro & Camargo 2003b). WhileMichener (2007a) included genus-group keys, the synonymy of a larger part of the regional fauna under thepolyphyletic subgenus Heterotrigona, sensu Michener, makes that key less useful. The only other genus-group key currently available is that of Moure (1961). However, as the fauna is not very extensive, species-group keys may be preferred (Schwarz 1939a; Sakagami et al. 1990), although additional literature may needto be consulted for poorly characterized genera like Tetragonula. For these genera, regional keys can be useful(Sakagami 1978; Dollin et al. 1997). No workable key exists for Austroplebeia (although see Rayment 1935:735), Platytrigona (although see Michener 1990: 127), and some of the varieties here considered as valid spe-cies.
Distribution and biology
The distribution of stingless bees in the Indo-Malay/Australasian region stretches from India to the SolomonIslands and from China (Yunnan, Hainan, Taiwan: Wu 2000) to Australia (New South Wales). Schwarz (1937)suggested that the greater abundance of stingless bees in Thailand and Malaysia, including all of Borneo, isdue to the abundance of resin-secreting trees (Dipterocarpaceae) and humid tropical climate, although mostlikely an array of factors are responsible for this concentration of species.
Natural history observations summarized from the present catalog are as follow:Colony pests: Achroea grisella (Lepidoptera: Tineidae), Ceria cf. ornatifrons (Diptera: Syrphidae) (larvae
feeding from provision), and Protaetia mandarinea (Coleoptera: Scarabaeidae). Insects and other organisms inhabiting nests or inquilines: Brachypeplus auritus, B. basalis, B. meyricki,
Predators recorded from flowers and around nests: Amblyseius (Acarina: Phytoseiidae) (of inquilines?),Ardeola grayii (Ciconiiformes: Ardeidae), Argiope keyserlingi, A. sp. (Araneae: Araneidae), Bembix flavipes,B. musca (Hymenoptera, Crabronidae), Cypsiurus balasiensis (Apodidae, Apodiformes), Dicrurus macrocer-cus, D. paradiseus (Dicruridae: Passeriformes), Dorniphora trigonae (Diptera: Phoridae) (of brood?), Helarc-tos malayanus (Carnivora: Ursidae), Hermetia illucens (Diptera: Stratiomyidae) (of brood), Meropsleschenaulti (Coraciiformes: Meropidae), Nephila maculata (Araneae: Nephilidae), Oecophylla smaragdina,O. sp. (Hymenoptera: Formicidae), Pahabengkakia piliceps (Hemiptera: Reduviidae), Phoridae (Diptera),small birds, Solenopsis geminata (Hymenoptera: Formicidae), Syrphidae (Diptera), and Thomisus spectabilis(Araneae: Thomisidae). A bee was also reported “clinging to the leg of a large green cicada”, Dundubia spic-ulata (Hemiptera: Cicadidae), although the details of that interaction was not provided.
Nest sites: aerial Crematogaster nests (Hymenoptera: Formicidae), bamboo stems and interspaces, beetleburrows, cavities among roots, coco nuts, cracks, crevices, holes in limestone, hollow pillars, hollow trunks(including Durio zibethinus, Bombacaceae; Eugenia aromatica, Myrtaceae; Ficus, Moraceae; Nephelium lap-paceum, Sapindaceae; Shorea laevis, Dipterocarpaceae), house posts, a human skull, and old tree stumps.
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TABLE 1. Different classificatory schemes used in the literature.
Lastly, floral records are frequently reported and show that the stingless bees are truly polylectic with 69families and 179 genera of plants recorded as hosts in the Indo-Malayan/Australasian region. This includesplants visited for nectar without performing pollination. The diversity of plants includes both cycads (Cyca-daceae) and angiosperms, including Monocot (e.g., Arecaceae, Orchidaceae, and Poaceae) and Dicot families.
Acknowledgments
I am very grateful to the many curators who replied to my queries about their type holdings. In particular toFrank Koch for hosting me during a visit to the Museum für Naturkunde, David Furth and Brian Harris forhosting me multiple times at National Museum of Natural History, and Holger H. Dathe and Stephan Blankfor hosting a brief visit to the DEI. David Notton kindly provided much information and loan of types fromthe Natural History Museum. J.M.F. Camargo gave me unlimited access to his comprehensive stingless beelibrary and collection which was helpful in forming the foundation for the present catalog, Camargo and Sil-via Pedro also provided useful comments on this project. John S. Ascher, Sydney A. Cameron, Jim B. Whit-field, Chris Dietrich, Nils Cordes, and two anonymous reviewers made helpful comments on the manuscriptand Ying Wang helped with translation of Chinese. Visits to institutions were supported by a Francis M. andHarlie M. Clark Research Support Grant and a National Science Foundation grant (DEB 0446325) to S. A.Cameron who also encouraged this study.
Genus: This work and Moure (1961) Type species Genus: Michener (2007)
Platytrigona Moure, 1961 Trigona planifrons T. (Heterotrigona)
Sundatrigona Inoue & Sakagami, 1993 Trigona moorei T. (Heterotrigona)
Tetragonilla Moure, 1961 Trigona atripes T. (Heterotrigona)
Tetragonula Moure, 1961 Trigona iridipennis T. (Heterotrigona)
Tetrigona Moure, 1961 Trigona apicalis T. (Heterotrigona)
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Trigona (Geniotrigona) incisa Sakagami & Inoue 1989: 605-610*, 614, 615, 617, 618, 619: Holotype(RMNH); paratypes (RMNH, 6 workers; MBBJ, 1 worker; SFS, remaining). Rasmussen & Cameron(2007) found that this taxon is distinct from Geniotrigona and it is awaiting a new genus to be proposed(taxonomy); Type locality: INDONESIA "Modoinding Minahasa, N. Celebes, vi. 26-27, 1941, nativecollector, ded. F. Dupont" (3 workers, holotype, paratypes); "Todyamboe, 900m, C. Celebes" (3 workers);"Wuasa, Kab. Poso, Sulteng, Sulawesi" (9 workers); "Lorei Lindu Nat. Park, C. Sulawesi" (3 workers);Tadauchi et al. 1998: 242-243 (type depositories); Rasmussen & Cameron 2007: (systematics).
Trigona lacteifasciata Cameron 1902: 131: Holotype (BMNH 17b.1132). Probably a distinct taxon of tho-
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Trigona thoracica Smith 1857: 50: Type (BMNH 17b.1181) (taxonomy); Type locality: SINGAPORE "Sin-gapore" (worker); Smith 1871: 395 (distribution); SINGAPORE; Dalla Torre 1896: 584 (citation); Bing-ham 1897: 561, 564 (distribution, key to species, taxonomy); MYANMAR, Tenasserim; SINGAPORE;Bingham 1905: 58 (citation, distribution); Friese 1909("1908"): 358 (comparative note); Cockerell1919c: 79 (distribution); Cockerell 1923: 240 (key to species); Dover 1929: 62 (citation, distribution);MALAYSIA, Kuala Tahan; Pahang; SINGAPORE; Friese 1933a: 46 (comparative note); Schwarz 1937:317, 318; Schwarz 1939a: 85, 90, 92, 104-106* (distribution, key to species, taxonomy); Floral record:Citrus limon; Tephrosia candida; Schwarz 1948: 83, 104, 114, 118; Moure 1961: 212, 213 (systematicposition); Yoshikawa et al. 1969: 172, 173 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang;Tapah; Touaran; Sakagami 1975: 56* (distribution); SINGAPORE; THAILAND; Wille 1979b: 247(morphology); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floralrecord: Xerospermum intermedium; Sakagami et al. 1983b: 2, 4, 13-15, 16, 20 (honey, nest (in Shorealaevis: Dipterocarpaceae)); Sakagami et al. 1985: (distribution, key to species); Baker & Delfinado-Baker 1985: 231 (inquiline (Neocypholaelaps phooni: Ameroseiidae: Acari)); Camargo 1988: 370 (dis-tribution, taxonomy); SINGAPORE; Sakagami & Inoue 1989: (morphology); Michener 1990: 105, 126(illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species);INDONESIA, Sumatra; MALAYSIA, Borneo; MYANMAR; THAILAND; Salmah et al. 1990: (distribu-tion, nest (hollow tree trunks; holes in limestone)); INDONESIA, Sumatera Barat; Inoue et al. 1990b:(population); Inoue 1990: 582 (nest (huge tree cavities), population); Johnson & Appanah 1990: (preda-tor (Oecophylla: Formicidae: Hymenoptera)); Michener & Roubik 1993: 260 (comparative note, mor-phology); Cunha 1994: (systematics); Yamane et al. 1995: 294 (synonymy); Roubik 1996: 61(distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433 (pollination); Floral record:Santiria laevigata; Michener 1997: 25 (type species); Nagamitsu & Inoue 1998: (foraging, morphol-ogy); Heard & Dollin 1998: (citation); Heard & Dollin 1998: 11 (pollination); Floral record: Averrhoacarambola; Liow 1999: 106, 127 (distribution); Nagamitsu et al. 1999: (floral records (not by species),foraging); Heard 1999a: 191, 192, 195 (citation); Michener 2000: 801-802; Chinajariyawong & Sai-boon 2000: (distribution, nest (as thorasica!)); THAILAND, Songkhla; Liow et al. 2001: 184; Nag-amitsu & Inoue 2002: 305 (foraging); Eltz et al. 2003: 309 (citation); Eltz 2004: 322 (abundance);Nagamitsu & Inoue 2005: 76, 81, 83, 86, plate 9 (distribution, foraging); Klakasikorn et al. 2005: 3, 6(floral record); Floral record: Dipterocarpus alatus; Hopea; Roubik 2006: 129, 130, 131 (nest); Slaa et
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Melipona castanea Bingham 1903: vi: Type (BMNH 17b.1128) (taxonomy); Type locality: THAILAND"Bukit Besar, Nawngchik. 1500 to 2500 feet" (worker); Bingham 1905: 59, plate A10 (citation); Cocker-
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Melipona lutea Bingham 1897: 559, 560, 564: Type (BMNH 17b.1122) (distribution, illustration, key to spe-cies, taxonomy; notice this is no longer a senior homonym of Trigona mocsaryi lutea Friese 1903, seeCamargo & Pedro 2007: 472); Type locality: MYANMAR "Northern Tenasserim; Karennee" (worker);Cockerell 1929b: 140 (comparative note); Moure 1961: 201 (systematic position); Wu 2000: 387, 390,plate (key to species, morphology).
Homotrigona indet. Roubik 2006: 130 (nest).
Lepidotrigona
Lepidotrigona arcifera (Cockerell, 1929)
Trigona arcifera Cockerell 1929c: 591-592: Holotype (BMNH 17b.1081). The ventralis species group (tax-onomy); Type locality: INDIA "Testa bridge, Himalayas, India, 10.1.97 (Sladen)" (worker); Schwarz1937: 295; Schwarz 1939a: 85, 88, 134, 135 (distribution, key to species, taxonomy); Nogueira-Neto1949: 121 (bee-keeping); Moure 1961: 199 (possible synonymy); Sakagami 1975: 67, 71 (synonymy);Sakagami et al. 1990: 126 (synonymy); Wu 2000: 387 (synonymy).
Lepidotrigona doipaensis (Schwarz, 1939)
Trigona (Lepidotrigona) ventralis variety doipaensis Schwarz 1939a: 85, 94, 136*: Holotype (USNM 53563,worker): examined, "Doi Pa / Mai Deng / Siam 750m / 12-29-32", "HughSmith / Coll", "Type No./ 53563/ U.S.N.M.", "Holotype", "Trigona ventralis / var. doipaensis / H.F. Schwarz". The ventralis species group(distribution, key to species, taxonomy); Type locality: THAILAND "SIAM.-Doi Pa, Mai Deng, 750meters, Dec. 29, 1932 (H. M. Smith)" (worker); Moure 1961: 199 (systematic position); Sakagami 1975:67, 69, 71 (distribution, variation); THAILAND, N; Sakagami & Yamane 1984: 35; Sakagami et al.1990: 126 (synonymy); Wu 2000: 387 (synonymy); Klakasikorn et al. 2005: 3, 6 (floral record); Floralrecord: Ficus religiosa; Homalium grandiflorum.
Lepidotrigona flavibasis (Cockerell, 1929)
Trigona flavibasis Cockerell 1929c: 592: Holotype (AMNH). The ventralis species group (taxonomy); Typelocality: THAILAND "Doi Sutep, Siam, Feb. 9 (Alice Mackie)" (worker); Schwarz 1939a: 85, 94, 134-135*, 136 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199 (systematicposition); Yoshikawa et al. 1969: 174 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang; Sakag-ami 1975: 67, 70, 71, 74 (distribution, variation); CAMBODIA; MALAYSIA; THAILAND; Sakagami& Yamane 1984: 38, 47; Sakagami et al. 1990: 126 (synonymy); Wu 2000: 387 (synonymy); Green etal. 2001: 91 (microsatellite); Chinh et al. 2005: 366, 368-371 (nest (as ventralis in part)); VIETNAM,Cucphuong National Forest, Red river delta; Chinh & Sommeijer 2005: (reproduction (as ventralis in
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Trigona (Lepidotrigona) ventralis variety hoosana Schwarz 1939a: 85, 94, 135-136*: Holotype (AMNH)(distribution, key to species, taxonomy); Type locality: TAIWAN "FORMOSA.-Taihorin, (H. Sauter)"(workers, March 10 and May 10); Moure 1961: 199 (systematic position); Sakagami 1975: 67 (varia-tion); Sakagami & Yamane 1984: 34-48; Sakagami et al. 1990: 126, 131 (distribution); Wu 2000: 387(synonymy); Chinh et al. 2005: 371.
Trigona ventralis hoozana Strand 1913b: 66: Syntypes (DEI, 9 workers). According to S. Blank (DEI)labeled "II. Hoozan Gormosa H. Sauter 10." (for February 1910); "Trigona ventralis Sm. Strand det.",some with "F. Sm." instead of "Sm.". Strand proposed the name conditionally (ICZN 11.5.1 and 15.1), butas it was proposed as a geograhical restricted (=subspecific) form before 1961, it is considered valid(45.6.1). The ventralis species group (taxonomy); Type locality: TAIWAN "Hoozan, Februar" ('a row' ofworkers); Sakagami et al. 1985: (key to species); Sakagami & Yamane 1987: (oviposition); Salmah etal. 1990: 155 (altitude); Bravo 1992: 876 (citation, altitude); Yamane et al. 1995: 294 (comparative note);Sung et al. 2004: (morphology); Nagamitsu & Inoue 2005: 76 (distribution, foraging); Chinh & Som-meijer 2005: 500 (citation); Sung et al. 2006c: (distribution, nest); Sung et al. 2006a; Sung et al. 2006b:167 (pollination); Floral record: Mangifera indica.
Lepidotrigona javanica (Gribodo, 1891)
Trigona javanica Gribodo 1891: 109: Holotype (MSNG, worker). The terminata species group (taxonomy);Type locality: INDONESIA "Giava" (=Java) (1 worker); Dalla Torre 1896: 579 (citation); Friese1914a: 13 (distribution); INDONESIA, Nusa Kambangan; Friese 1914b: 58 (nest); Cockerell 1929c: 591(comparative note); Schwarz 1937: 295, 296; Schwarz 1939a: 85, 90, 94, 140* (distribution, key to spe-cies, taxonomy); Moure 1961: 199 (systematic position); Wille & Michener 1973: 14, 24, 48 (nest);Sakagami 1975: 65 (comparative note).
Lepidotrigona latebalteata (Cameron, 1902)
Trigona latebalteata Cameron 1902: 130-131: Syntypes (BMNH 17b.1084, 2 workers). The terminata spe-cies group (taxonomy); Type locality: MALAYSIA "Kuching, Sarawak" (worker); Smith 1857: 50 (asventralis, in part); MALAYSIA, "Malacca (Mt. Ophir)"; Cameron 1908: 193 (key to species (as latibal-teata!)); Schwarz 1937: 282, 285, 288, 294, 295-297*, 326 (key to species, taxonomy); Schwarz 1939a:85, 94, 135, 137, 139-140* (distribution, key to species, taxonomy); Schwarz 1948: 118 (morphology);Moure 1961: 199 (systematic position); Sakagami 1975: 65 (comparative note).
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Trigona terminata Smith 1878: 169: Type (BMNH 17b.1100). The terminata species group (taxonomy);Type locality: MYANMAR no precise locality, presumably "Maulmain, Tenasserim Provinces"(worker); Dalla Torre 1896: 584 (citation); Bingham 1897: 563 (comparative note, taxonomy); Cocker-ell 1929c: 591 (comparative note); Schwarz 1937: 282, 284, 288, 295, 296, 326; Schwarz 1939a: 85, 90,94, 95, 100, 127, 128, 133, 135, 136-139* (distribution, key to species, taxonomy); Schwarz 1948: 118;
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Pagden 1957: 17 (comparative note); Sakagami 1960: 146 (distribution, variation); CAMBODIA, Snoul;Stung Treng; Moure 1961: 199 (systematic position); Wille 1961: 117, 121; Sakagami & Yoshikawa1961: 440 (distribution); LAOS, Thakhek; THAILAND, Mae Fack; Yoshikawa et al. 1969: 168, 169, 174(distribution, nest); MALAYSIA, George Town, Penang; Wille & Michener 1973: 14, 25, 49, 60 (nest);Sakagami 1975: 65-66*, 72, 73, 74 (distribution, variation); LAOS; THAILAND; VIETNAM; Sakag-ami 1978: 239 (comparative note); Sakagami et al. 1983b: 2, 15-19, 20 (nest); Sakagami & Yamane1984: 40, 47; Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts));Sakagami & Yamane 1987: 24, 25 (comparative note); Camargo 1988: 371 (distribution, taxonomy);INDONESIA, Java; MALAYSIA, Penang; Roubik 1989: 181; Michener 1990: 95, 105, 120, 122 (illus-tration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); CAMBO-DIA; INDONESIA, Java; Sumatra; MALAYSIA; MYANMAR; THAILAND; Salmah et al. 1990:(distribution, nest (hollow tree trunks)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population);Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Michener & Roubik1993: 260 (comparative note, morphology); Yong 1993: (biochemistry); Yamane et al. 1995: 294 (synon-ymy); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 434, 436(pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Liow1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener2000: 800; Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Wu 2000:386, 388-390 (key to species, morphology); Eltz et al. 2001b: 343, 345 (foraging); MALAYSIA, Borneo,Sabah; Green et al. 2001: 91 (microsatellite); Eltz et al. 2002a: 30, 31, 32 (distribution, foraging, nest,population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Floral record: Rhizophora apiculata;Klein et al. 2002: 1010 (floral record); Floral record: Coffea canephora; San & Osawa 2002: (popula-tion); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Kleinet al. 2003b: 957 (floral record); Floral record: Coffea canephora; Klein et al. 2003a: 841 (floral record);Floral record: Coffea canephora; Eltz 2004: 322 (abundance); Samejima et al. 2004: (population);MALAYSIA, Sarawak, Miri, Upper Baram area; Wattanachaiyingcharoen et al. 2004: (diversity, nest);Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Chinh et al. 2005: 368, 371 (citation);Klakasikorn et al. 2005: 3, 6 (floral record); Floral record: Ficus religiosa; Irvingia malayana; Slaa et al.2006: 297, 298 (citation); Tangmitcharoen et al. 2006: (pollination); Floral record: Tectona grandis;Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum;Rasmussen & Cameron 2007: (systematics); Michener 2007a: 824; Wattanachaiyingcharoen &Jongjitvimol 2007: 73 (comparative note).
Lepidotrigona trochanterica (Cockerell, 1920)
Trigona trochanterica Cockerell 1920a: 115: Holotype (BMNH 17b.1102) (taxonomy); Type locality:MALAYSIA "Sandakan, Borneo (Baker)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA,Sandakan; Schwarz 1937: 285, 287, 292-293*, 295, 296, 327 (key to species, taxonomy); Schwarz1939a: 85, 94, 133-134* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199(systematic position); Sakagami 1975: 65 (citation); Michener 1990: 122 (systematic position); Sakag-ami et al. 1990: (distribution, key to species, variation); INDONESIA, Sumatra; Salmah et al. 1990: (dis-tribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener 2000: 802;Nagamitsu & Inoue 2005: 76 (distribution, foraging); Michener 2007a: 827.
Lepidotrigona ventralis (Smith, 1857)
Trigona ventralis Smith 1857: 50: Type (BMNH 17b.1186); Mt. Ophir specimen is latebalteata. The ventra-
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"Trigona (Lepidotrigona) ventralis" nigribasis forma nov. Sakagami 1975: 71: Nomen nudum. The name isunavailable for taxonomic purposes, as it must be deemed to be infrasubspecific (ICZN, art. 15.2 and45.6.3.) (variation); VIETNAM.
Melipona tunneli Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 13.1.1). The statement that it "is differentfrom the other two in appearance as it possesses somewhat yellow spots in the abdomen", however, couldmake it a nomen dubium, although most likely then a junior synonymon of arcifera (common name (ngapsiwor), honey, nest).
Lisotrigona
Lisotrigona cacciae (Nurse, 1907)
Melipona cacciæ Nurse 1907: 619: Lectotype (BMNH 17b.1103, worker); Type locality: INDIA "Hoshang-
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abad, Central Provinces" (workers); Moure 1961: 194, 195 (lectotype designation (see Engel 2000), mor-phology, systematic position); Sakagami et al. 1985: (key to species (as cassiae!)); Michener 1990: 131(systematic position); Sakagami et al. 1990: 131 (distribution (as cassiae!)); Cunha 1994: (systematics);Michener 1997: 33 (type species); Michener 2000: 793; Engel 2000: (key to species, taxonomy);MALAYSIA, Sandakan, Borneo; THAILAND, Pat Meeung mountains; Chantabari; "Yunzalin"; Jobiraj& Narendran 2004: 40, 43 (comparative note, key to species); Michener 2007b: (morphology);Michener 2007a: 817.
Trigona scintillans Cockerell 1920a: 116: Holotype (BMNH 17b.1115) (taxonomy); Type locality:MALAYSIA "Sandakan, Borneo (Baker)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA,Sandakan; Cockerell 1929c: 590 (distribution, ecology ("great nuisance in camp setting on one's handsand face")); CAMBODIA, Angkor; Pah Meeung Mt.; Pahtoop Mt.; Schwarz 1937: 289, 290, 307*, 327,plate 7 (key to species, taxonomy); Schwarz 1939a: 85, 86, 94, 130*, 132 (distribution, key to species,taxonomy); Schwarz 1948: 118; Moure 1961: 193, 195 (morphology, systematic position); Sakagami &Yoshikawa 1961: 440 (distribution); THAILAND, Bangkok; Sakagami 1975: 51-52* (distribution, mor-phology); LAOS; THAILAND; VIETNAM; Appanah 1982: (pollination); MALAYSIA, Negri Sembi-ran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution,key to species); Michener 1990: 96, 100, 105, 131 (illustration, morphology, systematic position); Saka-gami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra; LAOS, MALAY-SIA; THAILAND; VIETNAM; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat;Inoue et al. 1990b: (population); Oldroyd et al. 1992: 5 (pollination); THAILAND, Chanthaburi; Floralrecord: Archontophoenix alexandrae; Michener & Roubik 1993: 257, 258 (comparative note, morphol-ogy); Nagamitsu & Inoue 1997: 436 (pollination); Floral record: Santiria laevigata; Nagamitsu et al.1999: (floral records (not by species), foraging); Engel 2000: (synonymy); Rajitparinya 2000: (distribu-tion (as scintillan! in part)); Rajitparinya 2000: (distribution, nest); Samejima et al. 2004: (population);MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (distribution, foraging);Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAI-LAND, Kanchanaburi, Thong Pha Phum; Michener 2007b: 130 (synonymy).
Lisotrigona carpenteri Engel, 2000
Lisotrigona carpenteri Engel 2000: 232-235, 236, figs. 1-3: Holotype (IEBR); 3 paratypes (AMNH, IEBR)(illustration, key to species, taxonomy); Type locality: VIETNAM "VIETNAM: Nghê An Province: KheBo, 19 03N 104 43E, 25-28 April 1998, 123 m, James M. Carpenter" (worker); VIETNAM, Ha Thing, SEHuong Son; Jobiraj & Narendran 2004: 39 (key to species); Chinh et al. 2005: 363, 365-367, 370(nest); VIETNAM, Cucphuong National Forest, Red river delta; Michener 2007b: 130 (comparativenote).
Lisotrigona furva Engel, 2000
Lisotrigona furva Engel 2000: 235-236: Holotype (SEMC); 5 paratypes (AMNH, SEMC) (floral record, keyto species, taxonomy); Type locality: THAILAND "THAILAND: Nakhon Ratchasima Prov.: ThaChang, 10 February 1993, S. Boongird and C. Michener" (worker); THAILAND, Chiang Mai, W Fang;Floral record: Buddleja asiatica; Callistemon; Jobiraj & Narendran 2004: 40 (key to species);Michener 2007b: (morphology).
Lisotrigona mohandasi Jobiraj & Narendran, 2004
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Trigona canifrons Smith 1857: 51: Syntype (OUMNH, 1 worker); additional putative type (BMNH17b.1183) (taxonomy); Type locality: MALAYSIA "Borneo (Sarawak)" (worker); Smith 1871: 395 (dis-tribution); MALAYSIA, Borneo; Dalla Torre 1896: 576 (citation); Bingham 1897: 560, 562 (distribu-tion, key to species, taxonomy); INDONESIA, Sumatra; MALAYSIA, Borneo; MYANMAR, Tenasserim;Cockerell 1905: 220 (see laeviceps); AUSTRALIA, Adelaide River (misidentification); SRI LANKA(misidentification, *iridipennis group); Bingham 1905: 58 (citation, distribution); Cameron 1908: 192,194 (comparative notes, key to species); Schulz 1909: (see laeviceps); Cockerell 1911: 176 (comparativenotes, taxonomy, misidentification, uncertain identity); Ihering 1912: 3 (citation); Friese 1914b: 53, 54,57 (identification, previous misidentification); Cockerell 1918: 385 (comparative notes); Cockerell1919b: 244 (comparative note); Prell 1919: (nest (in human skull)); Cockerell 1922a: 279 (misidentifica-tion); Cockerell 1923: 241, 242 (key to species); MALAYSIA, Sarawak; Cockerell 1929e: 242 (compar-ative note); Dover 1929: 62 (citation, distribution); SINGAPORE; Schwarz 1932: 246; Rayment 1932a:184 (citation); Rayment 1932f: 106 (taxonomy); Friese 1933a: 46 (distribution); INDONESIA, Meliau;Rayment 1935: 735, 740 (distribution, key to species, taxonomy); Schwarz 1937: 282, 289, 307-308*,320, 326, 327, plate 2, 5 (key to species, taxonomy); Floral record: Mallotus; Schwarz 1939a: 85, 88, 89,90, 92, 98, 101-102*, 111, 112 (distribution, key to species, taxonomy); Floral record: Mimosa pudica;Schwarz 1948: 18, 23, 117, 165; Moure 1961: 205, 206 (systematic position); Michener 1965: 231 (cita-tion); Yoshikawa et al. 1969: 173 (distribution); MALAYSIA, Kuala Lumpur; Michener 1974: 217; Sak-agami 1975: 55-56* (distribution, variation); MALAYSIA; THAILAND; Sakagami 1978: 239(comparative note); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve;Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Sakagami& Khoo 1987: 207 (comparative note); Uji 1987: (pollination); INDONESIA, E Kalimantan; Floralrecord: Nephelium lappaceum; Camargo 1988: 370 (synonymy); Roubik 1989: 105, 193; Michener1990: 88, 97, 105, 120, 126, 127 (systematic position); Sakagami et al. 1990: (distribution, key to spe-cies); INDONESIA, Sumatra; MALAYSIA, Borneo; MYANMAR; THAILAND; Salmah et al. 1990:(distribution, nest (hollow trunks of Ficus: Moraceae)); INDONESIA, Sumatera Barat; Inoue et al.1990b: (population); Roubik 1990b: 253 (defense); Inoue 1990: 582 (nest (huge tree cavities), popula-tion); Johnson & Appanah 1990: 586 (predator (Oecophylla: Formicidae: Hymenoptera) (as carni-frons!)); Roubik 1992: 345 (citation); Michener & Roubik 1993: 260, 264 (comparative note,
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Trigona (Hypotrigona) pendleburyi variety klossi Schwarz 1939a: 85, 94, 132*: Holotype (BMNH, no num-ber) (distribution, key to species, taxonomy); Type locality: MALAYSIA "MALAYA. State of Selangor:Bukit Kutu, 200 feet, Sept. 21, 1932, (H. M. Pendlebury)" (workers, holotype); "State of Pahang: KualaTeku, 500 feet, Dec. 8, 1921 (H. M. Pendlebury)" (workers); "N. BORNEO.-Bettotan, nr. Sandakan, (C.B. Kloss and H. M. Pendlebury)" (workers, July 28, 1927, and Aug. 15, 1927); Schwarz 1948: 118;Moure 1961: 193-194, 195 (morphology, systematic position); Yoshikawa et al. 1969: 174 (distribution);MALAYSIA, Fraser's Hill, Selagor-Pahang; Kiew 1985: (luring (unwashed shorts)); Michener 1990: 131(systematic position); Michener 2001: 235 (synonymy); San & Osawa 2002: (population); MALAYSIA,Negri Sembiran, Pasoh Forest Reserve; Inson & Malaipan 2006: (distribution, foraging, nest); THAI-LAND, Kanchanaburi, Thong Pha Phum.
Trigona (Hypotrigona) pendleburyi Schwarz 1939a: 85, 86, 94, 130-132*: Holotype (BMNH, no number);paratypes (BMNH (5)) (distribution, key to species, taxonomy); Type locality: MALAYSIA "CameronHighlands, Rhododendron Hill, 5000 feet (H. M. Pendlebury)" (workers, holotype, June 19,1923, andJune 22, 1923); MALAYA.-State of Pahang: Cameron Highlands, G. Tarbakar, 4500 feet, June 9, 1923 (H.M. Pendlebury); Cameron Highlands, Bukit Lendong, 5000 feet, May 21, 1931 (H. M. Pendlebury);Lubok Tamang, 3500 feet, March 5, 1924, and 4000 feet, June 10, 1923 (H. M. Pendlebury); Fraser's Hill,4600 feet, Pine Tree Hill, Oct. 25, 1932 (H. M. Pendlebury); Schwarz 1948: 118; Moure 1961: 192, 193-194, 195 (morphology, systematic position); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran,Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, keyto species); Michener 1990: 96, 100, 105, 129, 131 (illustration, morphology, systematic position); Saka-gami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; MALAYSIA; Salmah et al. 1990:(distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener & Rou-bik 1993: 257, 258, fig.15 (comparative note, morphology); Cunha 1994: (systematics); Roubik 1996:61, 62 (distribution, nest); BRUNEI, Kuala Belalong; Michener 1997: 46 (type species); Michener 2000:796; Engel 2000: (comparative note); Chinajariyawong & Saiboon 2000: (distribution, nest); THAI-LAND, Songkhla; Michener 2001: (distribution, morphology (male characters)); BRUNEI; INDONE-SIA, Sumatra; MALAYSIA, Sabah; THAILAND; Eltz et al. 2002a: 30 (distribution, foraging, nest,population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Eltz et al. 2003: (nest, tree preference);Jobiraj & Narendran 2004: 39; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Momose &Hamid 2005: 67, 71; Roubik 2006: 127 (nest); Inson & Malaipan 2006: (distribution, foraging, nest);THAILAND, Kanchanaburi, Thong Pha Phum; Michener 2007a: 821.
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Trigona hobbyi Schwarz 1937: 283, 288, 298-300*, 328: Holotype (BMNH 17b.1118). The hobbyi speciesgroup (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, 4000 feet, moss for-est, Oct. 18, 1932" (1 worker); Schwarz 1939a: 85, 129, 130* (distribution, key to species, taxonomy);Moure 1961: 205 (systematic position); Sakagami et al. 1985: (distribution); Michener 1990: 127 (sys-tematic position); Sakagami et al. 1990: 131 (distribution); Michener & Roubik 1993: 260 (comparativenote, morphology); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Samejima et al. 2004:(population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (distribu-tion, foraging); Rasmussen & Cameron 2007: (systematics).
Platytrigona keyensis (Friese, 1901)
Trigona keyensis Friese 1901: 271: Lectotype (ZMHB, worker): here designated, "AsiaArch. / Key Ins. /1900 / Kühn", "Trigona / keyensis / 1900 Friese det. / Fr.", "Type" (red label), "Coll. / Friese"; paralecto-type (AMNH (1)). The planifrons species group (manuscript name (latigena in litt.), taxonomy); Typelocality: INDONESIA "Key-Eilanden (Amboina, Nederland India) durch H. Kühn" (6 workers); Friese1909("1908"): 354, 356, fig. 15-7 (distribution); INDONESIA, Irian Jaya, Moaif; Timena; Friese 1909:272, 273, 277 (key to species, taxonomy); INDONESIA, Irian Jaya, Moaif; Timena; Strand 1911: 87 (dis-tribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Holl. N.G.; Pauwi; Samberi; Teba; Friese1912: 94 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, D. Neu Guinea; Pauwi; Sam-
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Trigona lamingtonia Cockerell 1929e: 243: Holotype (AMS, worker): examined, "Mt. Lamington / NorthernDivision / Papua v.1927 / C.T. McNamara", "Trigona / lamingtonia / Ckll TYPE", "HOLOTYPE", "Aus-tralian Museum K 238299". The planifrons species group (taxonomy); Type locality: PAPUA NEWGUINEA "Mt. Lamington, May, 1927 (C.T.McNamara)" (workers); Michener 1965: 231 (citation).
Platytrigona planifrons (Smith, 1865)
Trigona planifrons Smith 1865: 93-94: Holotype (OUMNH, worker): According to Baker (1993) the speci-men is labeled " 'N' [New Guinea (Allen); white disc] and 'Trigona planifrons 'Smith' ". The BMNH"type" (17b.1180) indicated by Moure (1961), is not a true type (Baker, 1993). The planifrons speciesgroup (taxonomy); Type locality: INDONESIA (IRIAN JAYA) "New Guinea" (worker); Smith 1871:396 (distribution); "NEW GUINEA"; Dalla Torre 1896: 582 (citation); Friese 1898: 427, 428 (citation,key to species); MALAYSIA, Malacca (misidentification); "NEW GUINEA"; Friese 1909("1908"): 354,356, fig. 15-8 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai;Jamur Gebiet; Manikion; Manokwari; Merauke; Moaif; Tawarin; Friese 1909: 272, 273, 274 (distribu-tion, key to species); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai; JanourGebiet; Manokwari; Merauke, Moaif; Tawarin; Friese 1915: 4 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai; Manikion; Manokwari; Merauke; Moaif; Sattelberg;Sekofro; Simbang; Tawarin; Tjahe Fluss; Zoutbron; Cockerell 1923: 241 (key to species); Cockerell1929e: 242 (comparative note, distribution); PAPUA NEW GUINEA, Fly River; Mt. Lamington;Schwarz 1948: 83, 109, 118; Moure 1961: 203, 204 (systematic position); Michener 1965: 231 (cita-tion); Camargo 1988: 372 (distribution); PAPUA NEW GUINEA, N. Guinea, Simbang, Huon Golf;Michener 1990: 88, 105, 120, 127, 153 (illustration, morphology, systematic position); Michener &Roubik 1993: 260 (comparative note, morphology); Cunha 1994: (systematics); Michener 1997: 48(type species); Michener 2000: 800-801; Hohmann 2003: (communication, nest); Michener 2007a: 824,826.
Sundatrigona
Sundatrigona lieftincki (Sakagami & Inoue, 1987)
Trigona (Trigonella) lieftincki Sakagami & Inoue 1987: 610, 611, 613-615, 617-624: Holotype (RMNH, notlocated, nor in SEHU); Paratypes (RMNH, 3 workers, 2 males) (morphology, taxonomy); Type locality:INDONESIA "N.E. Sumatra, Tongkoh, probably Mt. Talamau (Pasaman), VI 1941, v.d. M. Mohr leg." (5workers, 4 males, holotype worker, paratypes); "Koto Atas, Solok, Sumatera Barat" (6 workers); Sakag-ami & Inoue 1989: 615 (comparative note); Michener 1990: 127 (systematic position); Sakagami et al.1990: (distribution, key to species); INDONESIA, Sumatra; Salmah et al. 1990: (distribution, nest);INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (population); Tadauchi
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Trigona collina Smith 1857: 51-52: Type (OUMNH (=Wilson Saunders collection)) (taxonomy); Type local-ity: MALAYSIA "Malacca (Mount Ophir)" (worker); Smith 1871: 396 (distribution); MALAYSIA, Mal-acca; Dalla Torre 1896: 577 (citation); Bingham 1897: 560, 562 (distribution, key to species, taxonomy);MALAYSIA, Malacca; MYANMAR, S Tenasserim; Cameron 1901: 36 (distribution); THAILAND,Patalung; Waterhouse 1903: 133-134, plate 4 (see apicalis); Cameron 1908: 192 (key to species); Cock-erell 1918: 387 (comparative notes); Cockerell 1920a: 116 (comparative note); Cockerell 1923: 241, 242(key to species); Dover 1929: 62 (see apicalis); Schwarz 1937: 283, 288, 297-298*, 300, 301, 302, 326,327, plate 2, 5 (key to species, taxonomy); Schwarz 1939a: 85, 90, 94, 95, 123-125*, 126 (distribution,key to species, taxonomy); Floral record: Mimosa pudica; Schwarz 1948: 23, 117, 165; Schwarz 1951:63 (distribution); CAMBODIA, Angkor; Pagden 1957: 16-17 (flight); Sakagami 1960: 146 (distribu-tion); CAMBODIA, Stung Treng; Moure 1961: 211-212 (morphology, systematic position); Sakagami &Yoshikawa 1961: 437-439 (distribution, nest (only fig. 28)); CAMBODIA, Angkor Wat; Poipet; THAI-LAND, Chieng Mei; Doi Suthep; Li; Mae Hoi; Sara Buri; Yoshikawa et al. 1969: 168, 169, 173, 174 (dis-tribution, nest); MALAYSIA, Fraser's Hill, Selagor-Pahang; George Town, Penang; Nogueira-Neto 1970:291; Wille & Michener 1973: 13, 23, 47, 58, 70, 87 (nest); Sakagami 1975: 59-60*, 61 (distribution,variation); LAOS; MALAYSIA; THAILAND; VIETNAM; Sakagami 1978: 167-181, 231, 236, 237, 239(comparative note); Sakagami 1982: 389 (citation); Sakagami et al. 1983b: 5 (nest); Sakagami et al.
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Trigona bismarkiana Tadauchi et al. 1998: 245: Nomen nudum. The name is unavailable for taxonomic pur-poses, as no description or definition of the taxon was provided (ICZN, art. 15.1) (type depositories).
Trigona carbonaria Smith 1854: 414: Holotype (BMNH 17b.1136, worker) (taxonomy); Type locality:AUSTRALIA "Australia" (worker); Pascoe 1863: 171 (hive pest (Brachypeplus auritus: Nitidulidae:Coleoptera)); Smith 1863b: (nest); Smith 1863c: (nest); Smith 1863a: (inquiline (Brachypeplus auritus:Nitidulidae: Coleoptera)); Packard 1867: 365-366 (nest); Hockings 1884: (common name (karbi; keelar;kootchar), honey, morphology, nest (hollow trees), colony pest (Achroea grisella: Tineidae: Lepidoptera;Protaetia mandarinea: Scarabaeidae: Coleoptera), predator (small birds)); Dalla Torre 1896: 576 (cita-tion); Friese 1898: 428, 429, 430 (comparative note, key to species); Friese 1905: 118 (citation); Friese
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Trigona dapitanensis Cockerell 1925a: 492: Holotype (BMNH 17b.1135). Here considered a valid and largerform of laeviceps from the Philippines (taxonomy); Type locality: PHILIPPINES "Dapitan, Mindanao(Baker, 23119)" (worker); Schwarz 1939a: 86, 112, 113 (synonymy); Moure 1961: 208 (morphology,systematic position).
Meliponorytes devictus Cockerell 1921: 514-515: Holotype (BMNH In. 20702) (illustration, taxonomy);Type locality: MYANMAR "bead of clear pale Burmese amber from the Hukong Valley, received fromMr. R.C.J. Swinhoe" (worker); Cockerell 1922b: (age of fossil); Kerr & Maule 1964: (systematics, tax-onomy); Kerr & Esch 1965: (comparative note); Kerr 1969: 166 (systematics); Kerr & Cunha 1976:(systematics); Zeuner & Manning 1976: 222, 223, 224 (synonymy); Wille 1977: (systematics); Moure& Camargo 1978: 564-565 (morphology); Sakagami 1978: 216, 218-219 (synonymy); Wille 1979b: 243(comparative note); Grimaldi et al. 1995: 261, 262; Wu 2000: 392 (synonymy); Zherikhin & Ross2000: 6, 13-14 (synonymy).
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Trigona (Tetragona) sarawakensis variety drescheri Schwarz 1939a: 85, 93, 106-107*: Holotype (AMNH);paratype (AMNH (1) (distribution, key to species, taxonomy); Type locality: INDONESIA "M. JAVA.-South Banjoemas, Koebangkangkoeng, 25 meters, July 1935 (F. C. Drescher)" (1 worker, holotype); "E.JAVA.-Mt. Goemitir, Nov. 3 (R. van der Veen)" (1 worker, paratype); Schwarz 1948: 118; Sakagami1959: 120 (citation); Moure 1961: 210 (morphology, systematic position); Yoshikawa et al. 1969: 174(distribution); MALAYSIA, Tapah; Sakagami 1978: 210 (synonymy); Sakagami et al. 1983b: 10-12, 20(nest); Sakagami & Inoue 1985: 174, 176, 177, 178, 179, 180, 181, 182-184*, 186, 187-188 (key to spe-cies, nest, taxonomy); Sakagami et al. 1985: (distribution, key to species); Sakagami & Inoue 1987: 626(comparative note); Starr & Sakagami 1987: 97 (citation); Roubik 1989: 105; Sakagami et al. 1989:266 (comparative note); Sakagami et al. 1990: (distribution, key to species); INDONESIA, Java;Sumatra; MALAYSIA, Borneo; Sakagami & Inoue 1990: 201 (mixed brood arrangement); Salmah et al.1990: (distribution, nest (beetle burrows; crevices)); INDONESIA, Sumatera Barat; Inoue et al. 1990b:(population); Roubik 1996: 61, 62 (distribution, nest, predator (Helarctos malayanus: Ursidae: Car-nivora)); BRUNEI, Kuala Belalong; Michener & Boongird 2004: 144, 145 (comparative note); Nag-amitsu & Inoue 2005: 77 (distribution, foraging).
Tetragonula fuscobalteata (Cameron, 1908)
Trigona atomella Cockerell 1919b: 243-244: Holotype (USNM 29467, worker): examined, "Island of / Pen-ang / Baker", "Type No. / 29467 / U.S.N.M.", "Trigona / atomella / Ckll TYPE"); 2 paratypes (USNM)(comparative notes, taxonomy); Type locality: MALAYSIA "Island of Penang (Baker, 9585)" (worker);Cockerell 1920a: 116 (comparative note); Cockerell 1929c: 590 (distribution, ecology ("found clingingto the leg of a large green cicada (Platylomia spiculata (=Dundubia spiculata): Cicadidae: Hemiptera)"));CAMBODIA, Pahtoop Mt.; Cockerell 1930a: 157 (distribution, ecology (attracted to sweat)); CAMBO-DIA, Chun Het; Trang; Yasumatsu 1935: 94 (common name (mitsubachi), distribution); PALAUISLANDS; Esaki 1936: 44, 46-37 (common name (galadngikl; chaladngikl; mitsubachi), nest); PALAUISLANDS; Schwarz 1937: 290, 310, 311*, 316, 328 (taxonomy); Schwarz 1939a: 107, 108 (synonymy);Schwarz 1939b: 152 (synonymy); Cockerell 1939a: 61, 62, 64 (distribution ("presumably introduced byman")); PALAU ISLANDS, Melekeiok on Babelthuap; Ngeremlengui, Ogiwal; Schwarz 1948: 69, 117;Krombein 1950: 133 (synonymy); Moure 1961: 209 (morphology, systematic position); Sakagami1978: 223, 224 (synonymy); Camargo 1988: 371 (synonymy, taxonomy); Sakagami et al. 1990: 128(synonymy).
Trigona brevis Friese 1925: 41: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 12.1) (comparative note).
Trigona erythrostoma Cameron 1908: 193-194: Syntypes (BMNH 17b.1113, 2 workers) (key to species, tax-onomy); Type locality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker or queen); Cockerell1918: 386 (comparative notes); Cockerell 1920a: 116 (comparative note); Schwarz 1937: 290, 310*, 326(key to species, taxonomy); Schwarz 1939a: 112 (synonymy); Moure 1961: 195, 209 (synonymy).
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Trigona iridipennis Smith 1854: 413-414: Holotype (BMNH 17b.1114, worker): examined, "Type" (orangeborder), "iridipennis / Type Sm.", "B.M.TYPE / HYM. / 17B.1114", "Trigona / iridipennis / TYPE.Smith.", "Ceylon" (reverse side "53 / 23"). Citations for iridipennis are included in a broad sense, as spe-cies limits for this taxon is uncertain. To the same group belongs also T. bengalensis, T. praeterita, and T.ruficornis, as well as several undescribed species (see Rasmussen & Cameron (2007)) (taxonomy); Typelocality: SRI LANKA "Ceylon" (worker); Tennent 1859: 282 (list); Tennent 1861: 454 (list); Castets1893: 468, 481-488 (nest); Dalla Torre 1896: 579 (citation); Bingham 1897: 560, 563-564 (distribution,key to species, taxonomy); MYANMAR, Tenasserim; SRI LANKA; Friese 1904: 140 (see laeviceps);Friese 1909: 273 (comparative note); Strand 1913a: 149-150 (distribution (as laeviceps)); SRI LANKA;Friese 1914a: 12-13 (see laeviceps); Friese 1914a: 12 (synonymy (as canifrons)); Friese 1914a: 12 (syn-onymy (as laeviceps)); Friese 1914b: 49, 57 (citation (as laeviceps)); Friese 1914b: 43-58 (citation, nest);Friese 1914b: 53-54, 57 (citation, synonymy (as canifrons)); Cockerell 1919b: 242 (comparative notes);Cockerell 1919c: 79 (comparative notes, key to species); Cockerell 1919a: 121 (see laeviceps); Cocker-ell 1929c: 590, 592 (distribution, floral record, taxonomy, variation); CAMBODIA, Angkor (misidentifi-cation); Kum Puang Creek (misidentification); Nan (misidentification); Pahtoop Mt. (misidentification);INDIA, Bombay; Mangalore; PHILIPPINES, Palawan, Puerto Princesa (misidentification); SRI LANKA;THAILAND, Bangkok (misidentification); Floral record: Helianthus annuus; Cockerell 1930b: 149 (dis-tribution, floral record); MYANMAR, Rangoon; Floral record: Helianthus annuus; Friese 1933a: 46 (seelaeviceps); George 1933: (physiology (as irridipennis!)); George 1934: (brood, distribution, morphology,nest (crevice of wall), physiology (as irridipennis!)); INDIA, Coimbatore; Schwarz 1937: 283, 290, 308-310*, 314, 316, 326, 328, plate 3, 4 (key to species, taxonomy); Schwarz 1939a: 85, 86, 88, 89, 90, 93,96, 102, 108, 109, 110, 111-116*, 117, 118, 119, 120 (distribution, key to species, taxonomy (most of thespecimens examined by Schwarz correspond to laeviceps, see laeviceps)); Schwarz 1939b: 151 (compar-ative note); Schwarz 1945: 139, 141 (citation); Schwarz 1948: 4, 23, 31, 33, 39, 48, 50, 60, 62, 67, 68,71, 83, 84, 92, 109, 118, 141, 165, 233; Nogueira-Neto 1949: 121 (bee-keeping (as iridipenis!)); Krom-bein 1950: 133, 135 (comparative note); Krombein 1951: 278, 295 (morphology); Lindauer 1956: 546-554, 555 (communication, nest); Pagden 1957: 16 (flight); Lindauer & Kerr 1958: 407, 410 (communi-cation); Sakagami 1959: 120, 121 (comparative note); Portugal-Araújo & Kerr 1959: 226 (comparativenote); Sakagami 1960: 146 (see laeviceps); Lindauer & Kerr 1960: 31, 32, 34 (communication); SRILANKA; Lindauer & Kerr 1960: 31, 32, 34; Kerr 1960: 386 (citation); Moure 1961: 206, 207, 208(morphology, systematic position); Michener 1961: 3, 10, 28-33, 35, 38, 39, 42 (see laeviceps); Sakag-ami & Yoshikawa 1961: 434, 435, 436 (comparative note); Darchen & Louis 1961: 18, 21, 25-27 (bio-geography, communication, nest); Kerr et al. 1963: 81 (citation); Kerr & Maule 1964: (comparativenote); Schmid 1964: 588 (citation); Michener 1965: 231 (citation); Kerr & Esch 1965: (comparativenote); Baltazar 1966: 407 (see laeviceps); Phadke 1968: (honey, uncertain identity (unidentified));INDIA, Mysore, Madhya Pradesh; Kerr 1969: 166 (systematics); Darchen 1969: 482 (flight);Yoshikawa et al. 1969: 168, 169, 173 (see laeviceps); Jander & Jander 1970: 361 (citation); Sakagami1971: 345 (citation); Wille & Michener 1973: 14, 24, 30, 48, 59, 71, 94, 96, 98, 100, 102-103, 257 (nest);Michener 1974: 57, 153, 154, 343 (colony, nest); Sakagami 1975: 59 (comparative note); Kerr &Cunha 1976: (systematics); Zeuner & Manning 1976: 222-224, plate 2 (morphology); Wille 1977: (sys-tematics); Kshirsagar & Chauhan 1977: (oviposition); INDIA, Karnataka; Moure & Camargo 1978:565 (synonymy); Sakagami 1978: 165, 166-194, 196, 197, 200, 201, 202, 203, 205, 206, 208, 211, 212,
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Trigona læviceps Smith 1857: 51: Holotype (OUMNH). Moure (1961) indicated that the BMNH type speci-men (17b.1184) came from Mt. Ophir, which was not the type locality and the specimen cannot be consid-ered a true type. Baker (1993) located three specimens in OUMNH and labeled one as the holotype.Unfortunately that specimen is identic to fuscobalteata according to Baker (1993), but this issue will beresolved in a separate paper with C.D. Michener. Citations below for laeviceps are in a broad sense, asspecies limits are uncertain and may include both valdezi and testaceitarsis. These taxa have previouslybeen proposed as junior synonymons of laeviceps, but are slightly larger (taxonomy); Type locality: SIN-GAPORE "Singapore" (worker); Smith 1858: (propolis, wax); Smith 1859: 135 (distribution); INDO-NESIA, Aru; INDIA (misidentification); SINGAPORE; Smith 1865: 93 (distribution); "NEWGUINEA"; Parish 1866: (common name (pwai-ngyet), illustration, nest, propolis); Smith 1871: 395 (dis-tribution); SINGAPORE; Anderson 1890: 24 (citation); Dalla Torre 1896: 580 (citation (as leviceps!));Bingham 1897: 560, 564 (distribution, key to species, taxonomy, uncertain identity (as praeterita));MYANMAR, Rangoon; SRI LANKA, Trincomali; Bingham 1897: 560, 563 (distribution, key to species,nest (often build in crevices in the brickwork of the walls of houses), taxonomy); MYANMAR, Tenasse-rim; SINGAPORE; Friese 1898: 427 (key to species); Friese 1904: 140 (floral record, uncertain identity(as iridipennis)); INDONESIA, Java; Floral record: Canna flavescens; Cockerell 1905: 220 (comparativenotes, synonymy, uncertain identity (as canifrons)); AUSTRALIA, Adelaide River (misidentification);SRI LANKA (misidentification, *iridipennis group); Bingham 1905: 59 (citation, distribution, nest(house-post in village)); Brown 1906: 686 (distribution); PHILIPPINES; Schulz 1907: (nest, predator(Oecophylla smaragdina: Formicidae: Hymenoptera)); Friese 1909("1908"): 354, 355 (distribution,uncertain identity (as praeterita)); INDONESIA, Irian Jaya, Etna Bai; Merauke; Friese 1909("1908"):354, 356 (distribution); INDONESIA, Irian Jaya, Manikion; Wendesi; Friese 1909: 272, 275 (key to spe-cies, nest (in structural wood)); INDONESIA, Ost-Java; Sumatra; Schulz 1909: (nest (as canifrons inpart)); Friese 1912: 94 (distribution); PAPUA NEW GUINEA, Sepik-Bivak; Ihering 1912: 3 (citation (aslaevicops!)); Strand 1913a: 149-150 (see iridipennis); Friese 1914a: 12-13 (distribution, floral record,uncertain identity (as iridipennis)); INDONESIA, Bandung; Batavia; Buitenzorg; Nongkodjadjar; NusaKambangan; Salatiga; Samarang; Floral record: Canna flavescens; Friese 1915: 4 (distribution, uncertainidentity (as praeterita)); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Etna Bai; Jasa; Merauke;Njao; Zoutbron; Cockerell 1918: 385 (comparative notes); Cockerell 1919c: 77, 79 (distribution, previ-ous misidentifications); Cockerell 1919a: 121 (comparative note, uncertain identity (as iridipennis));INDONESIA, Java; Cockerell 1922a: 279 (distribution); Cockerell 1922b: (comparative note); Cocker-ell 1923: 241 (key to species); INDONESIA, Aru; Cockerell 1925a: 492 (comparative note); Cockerell1925b: (misidentification (was a Diptera)); Cockerell 1926b; Salt 1929: 460 (citation); Cockerell 1929c:590, 592 (comparative note); Cockerell 1929e: 242, 243 (comparative note); Cockerell 1929d: 300 (com-parative note); Cockerell 1930b: 150 (comparative note (previously as canifrons)); Schwarz 1932: 245,246; Rayment 1932f: 106 (taxonomy); Friese 1933a: 46 (distribution, uncertain identity (as iridipennis));INDONESIA, Meliau; Rayment 1935: 735, 739 (key to species, taxonomy); Schwarz 1937: 307, 308,314, 315; Schwarz 1939a: 86, 88, 111, 112 (distribution, key to species, taxonomy (most of the specimensexamined by Schwarz and identified as iridipennis are actually laeviceps, see iridipennis)); Floral record:Typha angustifolia; Schwarz 1945: 141 (citation); Schwarz 1948: 23, 84, 141; Nogueira-Neto 1949: 121(synonymy); Sakagami 1959: 120 (comparative note); Sakagami 1960: 146 (comparative note, distribu-
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tion, uncertain identity (as iridipennis)); CAMBODIA, Stung Treng; Moure 1961: 208 (morphology, sys-tematic position); Michener 1961: 3, 10, 28-33, 35, 38, 39, 42 (nest, uncertain identity (as iridipennis));PAPUA NEW GUINEA, Bisianumu, Sogeri Plateau; Bubia, vicinity of Lae; Kapagere, near Rigo;Koitakinumu Estate, Sogeri Plateau; Port Moresby; Michener 1965: 231 (citation); Baltazar 1966: 407(distribution, uncertain identity (as iridipennis)); PHILIPPINES, Mindanao, Lanao, Zamboanga; Pala-wan; Yoshikawa et al. 1969: 168, 169, 173 (distribution, nest, uncertain identity (as iridipennis));MALAYSIA, Fraser's Hill, Selagor-Pahang; Kuching; Bukit-Matajam, Prov. Wellesley; Zeuner & Man-ning 1976: 222, 223, 224 (synonymy); Wille 1977: 45 (synonymy); Sakagami 1978: 166-194, 196, 197,198, 200-208*, 209, 210, 211, 212, 213, 215, 216, 218, 219, 220, 221, 222, 226, 228, 229, 230, 234, 235,236, 237, 238, 239, 240, 247, plate 5 (nest, taxonomy); LAOS; MALAYSIA; SINGAPORE; THAI-LAND; VIETNAM; Sakagami & Ito 1981: 368, 373 (morphology); Sakagami et al. 1983b: 2, 6, 8, 10,13, 20 (nest); Sakagami et al. 1983a: (see minangkabau); INDONESIA, Sumatera Barat at Lubuk; Saka-gami et al. 1983d: (see minangkabau); Sakagami et al. 1983c: (reproduction); Inoue et al. 1984b: (seeminangkabau); Sakagami & Yamane 1984: 37, 48; Inoue et al. 1984a: (see minangkabau); Salmah et al.1984: (see minangkabau); Sakagami & Inoue 1985: 175, 176, 177, 180, 181, 182, 184, 186 (comparativenote); Sakagami et al. 1985: (distribution, key to species); Utami 1986: (pollination, uncertain identity(as irridipenis!)); INDONESIA, West Java, Pandeglang; Floral record: Metroxylon sagu; Sakagami &Khoo 1987: 208, 209, 210, 211, 212 (morphology); Sakagami & Inoue 1987: 611, 624 (comparativenote); Starr & Sakagami 1987: 97 (citation); Salmah et al. 1987: 384 (synonymy); Uji 1987: (pollina-tion, uncertain identity (as irridipenis!)); INDONESIA, E Kalimantan; Floral record: Nephelium lappa-ceum; Camargo 1988: 372 (distribution, synonymy); MALAYSIA, Sandakan, Penang; Sakagami &Inoue 1989: 615 (comparative note); Roubik 1989: 181, 205; Sakagami et al. 1989: 272 (misidentifica-tion); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDIA (misidentification);INDONESIA, Java; Sumatra; MALAYSIA; MYANMAR; THAILAND; Sakagami & Inoue 1990: (ovi-position); Salmah et al. 1990: (distribution, nest (hollow pillars; cracks), also as laeviceps small); INDO-NESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Johnson& Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Engels & Imperatriz-Fonseca1990: (see minangkabau); Salmah et al. 1991: 196 (synonymy); Prasad & Hemanth 1992: (predator(Ardeola grayii: Ardeidae: Ciconiiformes) (as leviseps!)); Michener & Roubik 1993: 260 (comparativenote, morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cardale 1993: 323-324 (see sapi-ens); Dollin & Dollin 1994: (collecting males); Yamane et al. 1995: 276, 293 (comparative note);Drumond et al. 1995: 44 (citation, comparative behavior); Imperatriz-Fonseca & Zucchi 1995: 239(citation); Inoue et al. 1995: 98 (pollination); Floral record: Neuwiedia borneensis; Inoue et al. 1996b:643 (misidentification); Salmah et al. 1996: 402, 410 (synonymy); Roubik 1996: 61 (distribution); BRU-NEI, Kuala Belalong; Ikudome & Kusigemati 1996: 20 (taxonomy); Smiley et al. 1996: 198, 200(inquiline (Hypoaspis hoffmannae: Laelapidae: Acari), uncertain identity (as iridipennis)); MALAYSIA,Selangor; Okada et al. 1996: 178 (pollination); Floral record: Neuwiedia veratrifolia; Nogueira-Neto1997: 94 (citation); Dollin et al. 1997: (revision, synonymy, taxonomy); Nagamitsu & Inoue 1997: 435,436 (pollination); Floral record: Santiria laevigata; Dollin 1997: (compared to sapiens); Nagamitsu &Inoue 1998: (foraging, morphology); Liow 1999: 106, 116, 127 (nest); Nagamitsu et al. 1999: (floralrecords (not by species), foraging); Peters et al. 1999: 382 (citation); Crane 1999: 89; van Veen & Som-meijer 2000: 70, 74, 75 (citation); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribu-tion, nest); Wu 2000: 387, 391-392, plate (key to species, morphology); Zherikhin & Ross 2000: 6, 13-14 (amber); Chinajariyawong & Saiboon 2000: (distribution, nest); THAILAND, Songkhla; Liow et al.2001: 184, 190; Green et al. 2001: 91 (microsatellite); Peeters & Ito 2001: 610-611 (citation);Wijesekara 2001: 154 (uncertain distribution); Kocyan & Endress 2001: 848, 856, 861, 864 (pollina-tion); Floral record: Neuwiedia veratrifolia; Eltz et al. 2002a: 30 (distribution, foraging, nest, population);
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Trigona melina Gribodo 1893: 262-263, 264: Syntypes (MSNG, 3 workers; unknown depository, 2 workers):Two syntypes are labeled "Bandjarmas" (?=Bandjarmasin, near Liangtelan), while the third syntype carryno locality label (F. Penati, pers. com.) (taxonomy); Type locality: MALAYSIA "Liangtelan (Borneo)" (2
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Nomen nudum. The name is unavailable for taxonomic purposes, as it must be deemed to be infrasubspe-cific (ICZN, art. 15.2 and 45.6.3.) (taxonomy); Sakagami et al. 1990: (distribution, key to species, varia-tion); Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b:(population); Tadauchi et al. 1998: 244 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution,foraging).
Trigona (Tetragonula) minangkabau Sakagami & Inoue 1985: 175, 176, 177, 178, 179, 180, 181, 184-186*,187-188: Holotype (MZB, Hymn.0198, worker); paratypes (MZB (Hymn.0195-0197)) (key to species,nest, taxonomy); Type locality: INDONESIA "Lubuk Mintrun nr. Padang, Sumatera Barat, Indonesia, ix1981 in mass flight, S.F. Skagami" (1 worker, holotype); "Lubuk Mintrun" (many workers, 25 males,paratypes); Sakagami et al. 1983a: (nest, taxonomy (misidentification; as laeviceps)); INDONESIA,Sumatera Barat at Lubuk; Sakagami et al. 1983d: (oviposition behavior (misidentification; as laeviceps));Inoue et al. 1984b: (brood, nest, reproduction (as laeviceps)); Inoue et al. 1984a: (physiology, reproduc-tion (as laeviceps)); Salmah et al. 1984: (development (as laeviceps)); Sakagami et al. 1985: (distribu-tion, key to species); Inoue et al. 1985a: (aggressiveness, foraging, nest); Inoue et al. 1985b: (foraging);Sakagami & Inoue 1987: 626 (comparative note); Starr & Sakagami 1987: 97, 106 (citation); Salmahet al. 1987: 383, 388, 389, 390 (development); Roubik 1989: 86, 133, 137; Sakagami et al. 1989: 271,272 (comparative note); Bego et al. 1989: 48, 53 (foraging, greenhouse, introduction); Sakagami 1989:164; Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; Sakagami & Inoue1990: 201, 202, 203, 205, 210, 211, 212, 215 (comparative note); Salmah et al. 1990: (distribution, nest(hollow rambutan branches (Nephelium lappaceum: Sapindaceae); bamboo; interspaces)); INDONESIA,Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Suka et al. 1990:(nestmate recognition); Roubik 1990a: 115 (citation); Engels & Imperatriz-Fonseca 1990: 186, 207(citation (as laeviceps)); Inoue et al. 1990a: 229; Salmah et al. 1991: (reproduction); Kakutani et al.1993: (pollination); Floral record: Fragaria; Suka & Inoue 1993: (behavior); Inoue et al. 1993: (nest(hollow rambutan branches (Nephelium lappaceum: Sapindaceae))); INDONESIA, Sumatra; Hoshiba &Imai 1993: 469, 477, 481, 487 (chromosome); Yamane et al. 1995: 276, 282, 293 (comparative note);Roubik 1995: 151 (antropogenic nest); Inoue et al. 1995: 98 (pollination); Floral record: Neuwiediaborneensis; Inoue et al. 1996b: (development, nest, variation); Salmah et al. 1996: 402, 407, 408, 409,410 (comparative note); Inoue et al. 1996a: (population); Tadauchi et al. 1998: 243-244 (type deposito-ries); Inoue et al. 1999: 208, 217 (citation, comparative table); Ranger & O'Donnell 1999: 189 (compar-ative note); Heard 1999a: 187, 193 (citation); Breed et al. 1999: 772-773 (citation, foraging); van Veen& Sommeijer 2000: 70, 74, 75 (citation (as laeviceps)); Amano et al. 2000: (citation); Nagamitsu &Inoue 2002: 309 (citation); San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh For-est Reserve; Rocha et al. 2003: 317 (citation); Tóth et al. 2003: 366 (citation); Costa et al. 2004: 187(citation); Tóth et al. 2004: 5, 6 (citation); Tsuji et al. 2004: (predator (Oecophylla smaragdina: Formi-cidae: Hymenoptera)); Nagamitsu & Inoue 2005: 77, 78 (distribution, foraging); Hofstede & Som-meijer 2006: 388 (citation); Cepeda 2006: 176, 184 (citation); Cortopassi-Laurino et al. 2006: 278(citation); Slaa et al. 2006: 297, 305, 306 (citation); Leonhardt et al. 2007: 126 (citation).
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Melipona smithii Bingham 1897: 560, 563: Unnecessary replacement name for Trigona ruficornis Smith, necLamarck. Lamarck 1817 (and repeated in 1835) only listed Melipona favosa, M. amalthea, M. ruficrus, M.postica, and M. pallida. The replacement name therefore remains enigmatic, unless mistaken for M. rufi-crus Latreille. BMNH has a labeled type for this replacement name (17b.1129) (distribution, key to spe-cies, taxonomy); INDIA, Mainpuri, NW provinces; MYANMAR, Tenasserim; Bingham 1905: 59(citation, distribution); Schwarz 1939a: 88, 107, 108 (synonymy); Schwarz 1948: 14, 23, 56; Nogueira-
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Trigona wallacei Tadauchi et al. 1998: 245: Nomen nudum. The name is unavailable for taxonomic purposes,as no description or definition of the taxon was provided (ICZN, art. 15.1) (type depositories).
Trigona (Tetragonula) sirindhornae Michener & Boongird 2004: 143-146: Holotype (SEMC, worker);paratypes (AMNH (3), Insect Museum, Chakthong Building, Department of Agriculture, Bangkok, Thai-land (3), SEMK (11) (taxonomy); Type locality: THAILAND "Thailand, on the Isthmus of Kra: RanongProvince: Muang district, 9–13 November 2002" (11 workers); "Ngow Waterfall National Park" (7 work-
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Tetragonula indet. Anderson et al. 1982: (pollination); Floral record: Mangifera indica; Camargo 1988:372 (taxonomy); Drumond et al. 1996a: 396, 398, 399 (behavioral comparison); Dollin et al. 1997: (tax-onomy); Roubik 2006: 126, 127, 130, 132 (nest).
Tetrigona
Tetrigona apicalis (Smith, 1857)
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tion); Type locality: MALAYSIA "Perak, F.M.S., Batang Padang, Jor Camp, 1800 ft., June 4, 1923(Pendlebury)"; CAMBODIA, Patalung; MALAYSIA, Kualu Lumpur, Gombak valley; Schwarz 1937:302; Schwarz 1939a: 85, 94, 102, 127, 129-130* (distribution, key to species, taxonomy); Schwarz 1948:117; Moure 1961: 216 (morphology, systematic position); Sakagami 1975: 61, 62, 63, 64* (distribution,morphology, variation); MALAYSIA; THAILAND; Sakagami et al. 1985: (distribution, key to species);Michener 1990: 127 (systematic position); Sakagami et al. 1990: 131, 136 (distribution, key to species);San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu &Inoue 2005: 76 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation).
Tetrigona vidua (Lepeletier de Saint Fargeau, 1836)
Melipona vidua Lepeletier de Saint Fargeau 1836: 429: Syntype (MNHN, 1 worker): putative syntypeexamined, "Museum Paris / Bengale / Diard & Duvaucel 1815", "vidua", "Diard et Duvaucel", "type","TYPE", "M. Vidua / Lep. S. Farg / Bengale". I consider this a true type: Alfred Duvaucel (1793-1825)and Pierre-Médard Diard (1794-1863) arrived on their first expedition to Kolkata, India, in January 1818and spent years collecting in the Indo-Malayan region. The label date is wrong as they were still in Francein 1815, but did reside for long periods in India later (Claeys: 1954) and I consider the label provenancemerely the port of embarkation of the specimen, rather than the collecting locality. The type is similar tomelanoleuca and could be considered a senior synonymon of that. However, melanoleuca is a variabletaxon, and is here retained until further studies may document if in fact it is only a single taxon (taxon-omy); Type locality: INDONESIA/E TIMOR "Ile de Timor" (unknown); Spinola 1853: 94 (citation);Smith 1854: 413 (citation); Smith 1871: 395 (distribution); INDONESIA/EAST TIMOR, Timor; Castets1893: 481 (citation); Dalla Torre 1896: 585 (citation); Bingham 1897: 560, 561 (distribution, key to spe-cies, taxonomy, uncertain identity); INDIA (misidentification); INDONESIA; MALAYSIA; MYAN-MAR, Tenasserim; Haddon 1901: 87 (common name (manyi), bad omen for Ibans ("if a swarm of beessettled underneath a house that had recently been built it would be considered a bad sign"), uncertain iden-tity); Bingham 1905: 59 (citation, distribution); Friese 1914a: 13 (distribution (maybe itama?)); INDO-NESIA, Gunung Ungaran; Friese 1918: 519 (comparative note (maybe itama?)); Cockerell 1918: 387(comparative notes); Cockerell 1926a: 224 (comparative note); Salt 1929: 454 (citation); Schwarz 1937:299-300; Schwarz 1939a: 86, 88, 98, 129 (synonymy, taxonomic position); Schwarz 1948: 422; Moure1961: 217 (morphology, systematic position); Sakagami & Khoo 1987: 207 (comparative note); Cama-rgo 1988: 372 (distribution); INDONESIA, Sumatra; Wu 2000: 386, 390, plate (key to species, morphol-ogy).
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Trigona pallidicincta Cockerell 1918: 386: Holotype (BMNH 17b.1134, male) (key to species, comparativenotes, taxonomy); Type locality: SINGAPORE "Singapore (Baker)" (male).
Nomina nuda
Melipona pulla Illiger 1806: 158: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 12.1) (citation); INDONESIA, Sumatra;Melipona pulla Dalla Torre 1896: 582 (citation).
Melipona Khasiana Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 13.1.1) (common name (nagp hamang),honey, nest).
Melipona terrestris Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 13.1.1) (common name (nagp khyndew),honey, nest).
Trigona (Pacificotrigona) okazawai Tadauchi et al. 1998: 245: Nomen nudum (as Trigona (Pacificotrigona)okazawai irianensis). The name is unavailable for taxonomic purposes, as no description or definition ofthe taxon was provided (ICZN, art. 15.1).
Trigona (Pacificotrigona) okazawai irianensis Tadauchi et al. 1998: 245: Nomen nudum. The name is
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
TERMS OF USEThis pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.
sericea, see apicalissirindhornae, Tetragonula sirindhornae (Michener & Boongird, 2004) smithii, see ruficornissymei, Austroplebeia symei (Rayment, 1932) terminata, Lepidotrigona terminata (Smith, 1878) terrestris, nomen nudum testaceinerva, see melanocephalatestaceitarsis, Tetragonula testaceitarsis (Cameron, 1901) thoracica, Geniotrigona thoracica (Smith, 1857) trochanterica, Lepidotrigona trochanterica (Cockerell, 1920) tunneli, nomen nudum, see Lepidotrigona valdezi, Tetragonula valdezi (Cockerell, 1918) ventralis, Lepidotrigona ventralis (Smith, 1857) versicolor, see fimbriatavidua, Tetrigona vidua (Lepeletier de Saint Fargeau, 1836) wallacei, nomen nudum, see sapienswebsteri, Austroplebeia websteri (Rayment, 1932) wybenica, see clypeariszucchii, Tetragonula zucchii (Sakagami, 1978)
Appendix 3. Museum acronyms. Curators and sources of information concerning the collection are listed in parenthesis
AMNH USA, New York, New York, American Museum of Natural History (Jerome G.Rozen, John S. Ascher)
AMS Australia, New South Wales, Sydney, Australian Museum (David Britton, MaxBeatson)
ANIC Australia, Australian Capital Territory, Canberra City, CSIRO, AustralianNational Insect Collection (John La Salle, Nicole Fisher)
BMNH United Kingdom, London, The Natural History Museum (David Notton)BPBM USA, Hawaii, Honolulu, Bernice P. Bishop Museum (Shepherd P. Myers)Calicut India, Kerala, Calicut University, Zoology DepartmentDEI Germany, Müncheberg, Deutsches Entomologisches Institut im ZALF (Holger
H. Dathe, Stephan Blank)EKYU Japan, Fukuoka, Kyushu University, Entomological laboratory (Tadauchi et al.
1998, website)HNHM Hungary, Budapest, Hungarian Natural History Museum (Sándor Csõsz)IEBR Vietnam, Hanoi, Institute of Ecology and Biological ResourcesMNHN France, Paris, Muséum National d'Histoire Naturelle (Claire Villemant)MSNG (Collezione Gribodo)Italy, Genova, Museo Civico di Storia Naturale "Giacomo Doria" (Fabio Penati)MVMA Australia, Victoria, Abbotsford, Museum of Victoria (Ken Walker)MZB Indonesia, Bogor, Museum Zoologicum Bogoriense (Yayuk R. Suhardjono)OUMNH (=Wilson Saunders collection)United Kingdom, Oxford, University Museum of Natural History
(James Hogan)QM Australia, Queensland, South Brisbane, Queensland MuseumRMNH Netherlands, Leiden, Nationaal Natuurhistorische Museum (“Naturalis”) (Cor-
nelius van Achterberg)SEHU Japan, Sapporo, Hokkaido University Museum, Systematic entomology (Masa-
hiro Ohara)SEMC USA, Kansas, Lawrence, University of Kansas, Snow Entomological Museum
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SFS Shôichi F. Sakagami’s collection (see SEHU)SMNS Germany, Stuttgart, Staatliches Museum für NaturkundeUSNM USA, Washington D.C., National Museum of Natural History, (formerly, United
States National Museum) (David Furth, Brian Harris)ZMHB Germany, Berlin, Museum für Naturkunde der Humboldt-Universität (Frank
Koch)ZRC Singapore, National University of Singapore, Raffles Museum of Biodiversity
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