Zootaxa, Catalog of the Indo-Malayan/Australasian ...

TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

ZOOTAXA 1935

Catalog of the Indo-Malayan/Australasian stingless bees (Hymenoptera: Apidae: Meliponini) CLAUS RASMUSSEN

Magnolia Press Auckland, New Zealand


Claus Rasmussen Catalog of the Indo-Malayan/Australasian stingless bees (Hymenoptera: Apidae: Meliponini) (Zootaxa 1935) 80 pp.; 30 cm. 17 Nov. 2008 ISBN 978-1-86977-295-6 (paperback) ISBN 978-1-86977-296-3 (Online edition)

FIRST PUBLISHED IN 2008 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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ISSN 1175-5334

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2 · Zootaxa 1935 © 2008 Magnolia Press

RASMUSSEN

TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Zootaxa 1935: 1–80 (2008) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2008 · Magnolia Press

ISSN 1175-5334 (online edition)

ZOOTAXA

Catalog of the Indo-Malayan/Australasian stingless bees (Hymenoptera: Apidae: Meliponini) CLAUS RASMUSSEN Department of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, USA. E-mail: [email protected]

Table of contents Abstract ............................................................................................................................................................................... 4 Catalog format ..................................................................................................................................................................... 4 Taxonomy and classification ............................................................................................................................................... 5 Distribution and biology ..................................................................................................................................................... 6 Acknowledgments ............................................................................................................................................................... 7 Catalog of the Indo-Malayan/Australasian stingless bees .................................................................................................. 8 Austroplebeia ............................................................................................................................................................... 8 Geniotrigona .............................................................................................................................................................. 11 Heterotrigona ............................................................................................................................................................. 13 Homotrigona .............................................................................................................................................................. 15 Lepidotrigona ............................................................................................................................................................. 17 Lisotrigona ................................................................................................................................................................. 21 Lophotrigona .............................................................................................................................................................. 23 Odontotrigona ............................................................................................................................................................ 24 Papuatrigona ............................................................................................................................................................. 24 Pariotrigona ............................................................................................................................................................... 25 Platytrigona ............................................................................................................................................................... 26 Sundatrigona .............................................................................................................................................................. 27 Tetragonilla ................................................................................................................................................................ 28 Tetragonula ................................................................................................................................................................ 31 Tetrigona .................................................................................................................................................................... 50 Meliponini indet ......................................................................................................................................................... 53 Incertae sedis ............................................................................................................................................................ 53 Nomina nuda .............................................................................................................................................................. 54 Appendix 1. Plants visited by stingless bees ..................................................................................................................... 55 Appendix 2. Stingless bee taxon names ............................................................................................................................ 59 Appendix 3. ...................................................................................................................................................................... 61 References ......................................................................................................................................................................... 62

Accepted by E. Almeida: 16 Oct 2008; published: 17 Nov. 2008

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Abstract This catalog of the published literature on stingless bees from the Indo-Malayan/Australasian region was compiled from 520 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 the bees have been reported to forage. Lectotypes are designated for Trigona ornata described by Rayment and the following species described by Friese: Trigona anamitica, T. australis, T. borneënsis, T. flaviventris, T. keyensis, T. luteiventris, T. pygmaea, T. reepeni, T. sericea, and T. versicolor. Key words: Austroplebeia, Geniotrigona, Heterotrigona, Homotrigona, Lepidotrigona, Lisotrigona, Lophotrigona, Odontotrigona, Papuatrigona, Pariotrigona, Platytrigona, Sundatrigona, Tetragonilla, Tetragonula, Tetrigona, Trigona, bibliography

Stingless bees (Hymenoptera: Apidae: Meliponini) are important pollinators of native plants in tropical and subtropical parts of the world (Heard 1999a). They form large perennial colonies with complex social organization (Michener 2007a), have a diverse nesting biology ranging from underground to exposed colonies (Schwarz 1948; Wille & Michener 1973; Camargo & Pedro 2003b; Roubik 2006; Rasmussen & Camargo 2008), and are abundantly encountered within their native range (Roubik 2006). In addition to a well-corroborated phylogeny, an up-to-date, comprehensive taxonomic and biological catalog is fundamental to any comparative evolutionary, ecological, and behavioral research on any group of organisms. A catalog provides an index to previous studies in taxonomy, behavioral research, and pollination ecology, thus consolidating the existing knowledge in an accessible format. The extensive Neotropical fauna was recently cataloged by Camargo & Pedro (2007: 391 spp.) and a systematic revision of the Afrotropical stingless bees included a nearly complete bibliography (Eardley 2004: 26 spp. incl. Madagascar). The present catalog treats the remaining Indo-Malayan/Australasian stingless bees (129 proposed species-group names, 89 currently accepted valid species) and completes efforts to catalog the global stingless bee fauna. Together with recent molecular phylogenies of the stingless bees (Rasmussen & Cameron 2007; Rasmussen 2008), it is expected that these tools 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 are not recognized in the catalog. Genus-group headings are in boldface type. Following the accepted speciesgroup names are listed synonyms, including validly proposed junior synonyms and nomina nuda. References are sorted in chronological order and include citation, page numbers from publications, type-specimen information from both type-specimen labels, and the original publication where the species was first described, key words, 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 the particular species is treated throughout the publication. Publication dates were determined by the publishers imprint date on the wrapper of the fascicle, when available, or information contained within the issue. When the 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. Key words in parenthesis after a reference indicate the kind of information provided and associated organisms recorded. Misidentifications are corrected in the key word section whenever necessary or known, and typographical errors in spelling of taxa names in publications are noted followed by an exclamation mark. Distributions are listed alphabetically with country names in upper case type and type localities indicated as in the original 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 conform with 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 rather left as cited in the original publication. Stingless bee type specimens are encountered in museums worldwide, as indicated in the catalog. Acronyms for institutions are abbreviated following Evenhuis (2007) and Tadauchi et al. (1998) for Sakagamirelated collections (Appendix 3). Curators who assisted with loans and important information on types are also listed following each institution. Of particular importance are the collections in the Natural History Museum (London), with 52 primary types, Museum für Naturkunde der Humboldt-Universität (Berlin), with 15, and National Museum of Natural History (Washington, D.C.), with 10. As for the most prolific describers of new taxa, Theodore D.A. Cockerell described 38 new taxa, Heinrich Friese 17, Frederick Smith 14, Herbert F. Schwarz and Peter Cameron 11 each, and Shôichi F. Sakagami 10. The whereabouts of three Sakagami primary types remain unknown (i.e., Trigona gressitti, T. pagdeniformis, and T. lieftincki) as these have not been found in the collections indicated by the late Sakagami nor in his private collection (Tadauchi et al. 1998). It may 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 it was 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 true types and putative type specimens from London needs to be examined and compared with specimens from University Museum of Natural History (Oxford) when in doubt. A number of species described before the standardization 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 multiple biological and morphological species. Lectotypes are here designated for several species proposed by Friese as 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 520 papers. A number of popular travel accounts are listed in the nearly complete stingless bees bibliography by Schwarz (1948). These were all consulted, but only included if they provided either the scientific name or some 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 treating some of the Old World stingless bees as subgenera of Trigona Jurine (Table 1). The phylogenetic groups of 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 bee catalog (Camargo & Pedro 2007). Other studies propose a simpler classification to avoid a profusion of formal names, which is equally valid when based on monophyletic lineages. This dual approach to classifying genera (chiefly Moure 1961; Michener 1990, 2007a), however, complicates literature searches with specific taxa appearing under several different generic names. The genus-group name or synonymy is listed for each taxon in Appendix 2. Most synonymies here are based on the published examination of types by Schwarz (1937; 1939a) and Moure (1961). However, a number of species were only tentatively synonymized by them, and re-examination of primary types was necessary for confirming and establishing their synonymies. In those cases, I indicated the types I examined. Just like the inconsistent usage of stingless bee generic combinations, some taxa have been treated as subspecies (e.g., Schwarz 1939a; Sakagami et al. 1990), species (e.g., Klakasikorn et al.

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES

Zootaxa 1935 © 2008 Magnolia Press ·

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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 combination 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 the fact that about 80% of the New World taxa originally described by Schwarz as subspecies in light of additional 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 specialized keys, although the fauna has not yet undergone the taxonomic scrutiny comparable to that of many New World taxa (e.g., Camargo & Moure 1994; Camargo & Moure 1996; Pedro & Camargo 2003b). While Michener (2007a) included genus-group keys, the synonymy of a larger part of the regional fauna under the polyphyletic subgenus Heterotrigona, sensu Michener, makes that key less useful. The only other genusgroup key currently available is that of Moure (1961). However, as the fauna is not very extensive, speciesgroup keys may be preferred (Schwarz 1939a; Sakagami et al. 1990), although additional literature may need to 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 species.

Distribution and biology The distribution of stingless bees in the Indo-Malay/Australasian region stretches from India to the Solomon Islands 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, is due to the abundance of resin-secreting trees (Dipterocarpaceae) and humid tropical climate, although most likely 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, B. planus (Coleoptera: Nitidulidae), Cerophagus trigona (Acari: Acaridae), Homalota trigonae (Coleoptera: Staphylinidae), Hypoaspis hoffmannae (Acari: Laelapidae), Neocypholaelaps phooni (Acari: Ameroseiidae), and Tribolium myrmecophilum (Coleoptera: Tenebrionidae). 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 macrocercus, D. paradiseus (Dicruridae: Passeriformes), Dorniphora trigonae (Diptera: Phoridae) (of brood?), Helarctos malayanus (Carnivora: Ursidae), Hermetia illucens (Diptera: Stratiomyidae) (of brood), Merops leschenaulti (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 spiculata (Hemiptera: Cicadidae), although the details of that interaction was not provided. Nest sites: aerial Crematogaster nests (Hymenoptera: Formicidae), bamboo stems and interspaces, beetle burrows, cavities among roots, coco nuts, cracks, crevices, holes in limestone, hollow pillars, hollow trunks (including Durio zibethinus, Bombacaceae; Eugenia aromatica, Myrtaceae; Ficus, Moraceae; Nephelium lappaceum, Sapindaceae; Shorea laevis, Dipterocarpaceae), house posts, a human skull, and old tree stumps.


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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. TABLE 1. Different classificatory schemes used in the literature. Genus: This work and Moure (1961)

Type species

Genus: Michener (2007)

Austroplebeia Moure, 1961

Trigona cassiae

Austroplebeia

Geniotrigona Moure, 1961

Trigona thoracica

Trigona (Heterotrigona)

Heterotrigona Schwarz, 1939

Trigona itama

T. (Heterotrigona)

Homotrigona Moure, 1961

Trigona fimbriata

T. (Homotrigona)

Lepidotrigona Schwarz, 1939

Trigona nitidiventris

T. (Lepidotrigona)

Lisotrigona Moure, 1961

Melipona cacciae

Lisotrigona

Lophotrigona Moure, 1961

Trigona canifrons

T. (Heterotrigona)

Odontotrigona Moure, 1961

Trigona haematoptera

T. (Heterotrigona)

Papuatrigona Michener & Sakagami, 1990

Trigona genalis

T. (Papuatrigona)

Pariotrigona Moure, 1961

Trigona pendleburyi

Pariotrigona

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)

Lastly, floral records are frequently reported and show that the stingless bees are truly polylectic with 69 families and 179 genera of plants recorded as hosts in the Indo-Malayan/Australasian region. This includes plants visited for nectar without performing pollination. The diversity of plants includes both cycads (Cycadaceae) 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 to Frank Koch for hosting me during a visit to the Museum für Naturkunde, David Furth and Brian Harris for hosting me multiple times at National Museum of Natural History, and Holger H. Dathe and Stephan Blank for hosting a brief visit to the DEI. David Notton kindly provided much information and loan of types from the Natural History Museum. J.M.F. Camargo gave me unlimited access to his comprehensive stingless bee library and collection which was helpful in forming the foundation for the present catalog, Camargo and Silvia Pedro also provided useful comments on this project. John S. Ascher, Sydney A. Cameron, Jim B. Whitfield, Chris Dietrich, Nils Cordes, and two anonymous reviewers made helpful comments on the manuscript and Ying Wang helped with translation of Chinese. Visits to institutions were supported by a Francis M. and Harlie M. Clark Research Support Grant and a National Science Foundation grant (DEB 0446325) to S. A. Cameron who also encouraged this study.



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Catalog of the indo-malayan/australasian stingless bees Austroplebeia Austroplebeia australis (Friese, 1898) Trigona australis Friese 1898: 428, 430, 431: Lectotype (ZMHB, worker): here designated, "C. Austr. / v. Müller.93", "Trigona / australis / det. Friese 1898", "n.sp."; paralectotypes (SMNS (unknown number), ZMHB (1)) (comparative note, key to species, taxonomy); Type locality: AUSTRALIA "C.-Australien (v. Müller 1893)" (many workers); "Burnett-River (Queensland, Simon 1891-92)" (several workers); Cockerell 1905: 221 (comparative notes); Friese 1909: 272, 273 (citation, key to species); AUSTRALIA, Queensland, Mackay; Cockerell 1930b: 150 (comparative note); Rayment 1932f: 107 (taxonomy); Rayment 1935: 735 (key to species); Schwarz 1948: 117; Moure 1961: 196 (morphology, systematic position); Michener 1961: 5-11, 12, 13, 14, 15, 22, 23, 27, 38, 39, 40, 42, 44 (aggressiveness, nest); Nogueira-Neto 1970: 266, 314; Wille & Michener 1973: 10, 18, 42, 53, 64, 82, 91, 94, 95, 96, 97, 98, 99, 102-103, 267, 268 (nest); Sakagami et al. 1973: 180, 181 (comparative note); Michener 1974: 341, 342, 343 (nest); McKenzie 1975: (common name (kuta), honey); Sakagami 1982: 375 (citation); Wagner & Dollin 1982a: (distribution); Wille 1983: 48, 50 (nest); Wagner & Briscoe 1983: (genetics); Graur 1985: 194 (comparative note); Yong 1986: 628 (comparative note); Goebel 1986: (bee-keeping); Yong et al. 1987: 467 (biochemistry); Milborrow et al. 1987: (wax); Heard 1987: (nest); Michener 1990: 134, 137 (illustration, morphology, systematic position); Cardale 1993: 318 (distribution, floral record, list); AUSTRALIA, NE coastal; Queensland; New South Wales; SE coastal; Murray-Darling basin; Floral record: Alphitonia; Angophora; Callistemon; Eucalyptus; Melaleuca; Rapistrum; Swainsona; Wahlenbergia; Heard & Exley 1994: 97 (comparative note); Drumond et al. 1995: 43 (citation, comparative behavior); Heard 1996: (bee-keeping, nest); Dollin 1996a: (characteristics, nest); Dollin 1996b: (nest); Nogueira-Neto 1997: 85, 204, 334; Dollin & Heard 1997: (domestication); Dollin & Heard 1997: 4 (bee-keeping); Dollin 1997: (characteristics); Drumond et al. 1999: (oviposition); Crane 1999: 89, 90; Dollin & Heard 1999: (bee-keeping); Michener 2000: 782-783; Drumond et al. 2000: (reproduction); Heard & Dollin 2000: (bee-keeping); Green et al. 2001: 91 (microsatellite); Serrão 2001: (morphology (proventricular structure)); Palmer et al. 2002: 2112 (citation); Koedam et al. 2002: 231, 232 (comparative note, wax); Green & Oldroyd 2002: 197 (citation); Dollin 2002a: (bee-keeping); Dollin 2002b: (bee-keeping); Costa et al. 2003: (systematics); Heard 2003: (citation); Tóth et al. 2004: 6 (citation); Heiling & Herberstein 2004: (predator (Thomisus spectabilis: Thomisidae: Araneae)); Slaa et al. 2006: 301 (pollination); Michener 2007a: 806-807; Temaru et al. 2007: 281 (honey properties); Klumpp 2007: 5, 6, 10, 11, 15-18, 29, 47, 51, 56, 57, 76, 77, 82. Austroplebeia cassiae (Cockerell, 1910) Trigona cassiæ Cockerell 1910: 247: Holotype (BMNH 17b.1137, worker): examined, "Type" (red border), "B.M. TYPE / HYM. / 14B.1137", "Trigona / cassiae / Ckll TYPE", "Turner Coll. / 1912-111", "Mackay / 12.99 / Cassia", "900"; paratype in "Rayment collection" (not located) (comparative notes, floral record, taxonomy); Type locality: AUSTRALIA "Mackay, Queensland, Dec. 1899 (Turner, 900)" (worker); male, Nov. 1891 (Turner, 318); Floral record: Cassia; Hockings 1884: (common name (kootchar), honey, morphology, nest (hollow trees) (as kootchar only; identification uncertain)); Cockerell 1922a: 279 (distribution); Cockerell 1929d: 299, 301 (common name (kootchar), distribution, nest (tree hollows), parasite? (Ceria cf. ornatifrons: Syrphidae: Diptera)); AUSTRALIA, Brisbane; Cockerell 1930b: 150 (common name (kootchar), comparative note, floral record, nest); Floral record: Cassia; Eucalyptus;


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Rayment 1930: 59 (comparative note); Rayment 1932a: 185 (common name (koochee)); Rayment 1932d: (honey, pollen, wax); Rayment 1932b: (nest); Rayment 1932c: (behavioral comparison); Rayment 1932e; Rayment 1932f: 105, 106 (comparative note); Rayment 1935: 735-736 (comparative note, key to species, taxonomy); Schwarz 1939a: 103 (synonymy); Schwarz 1948: 4, 23, 24, 29, 31, 32, 37, 38, 41, 62, 67, 68, 75-77, 80, 84, 91, 97, 107, 117, 134, 166; Moure et al. 1958: 484 (citation); Moure 1961: 195, 196, 197 (systematic position); Michener 1961: 5 (synonymy); Wille 1961: 117, 121, 125, 129; Darchen & Louis 1961: 21 (nest); Michener 1965: 230 (citation); Darchen 1969: 462 (illustration, physiology); Wille & Michener 1973: 10 (synonymy); Camargo 1988: 372 (distribution); AUSTRALIA, Queensland; Michener 1990: 96, 101, 105, 133, 134 (illustration, morphology, systematic position); Sakagami et al. 1990: 131 (see cacciae); Michener & Roubik 1993: 259, 262 (comparative note, morphology); Cardale 1993: 318-319 (distribution, floral record, list); AUSTRALIA, NE coastal; Queensland; Brisbane; Caloundra; Mackay; Floral record: Cassia; Eucalyptus; Cunha 1994: (systematics); Michener 1997: 10 (type species); Michener 2000: 791; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Michener 2007a: 815. Austroplebeia cincta (Mocsáry in Friese, 1898) Trigona cincta Friese 1898: 431: Holotype (HNHM, worker): examined, "Friedrich- / Wilh.-hafen", "N. Guinea / Biró 96" (taxonomy); Type locality: PAPUA NEW GUINEA "Neu-Guinea (Friedrich Wilhelmshafen)" (1 worker); Friese 1909("1908"): 354, 356 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Cyclopen Gebirge; Moaif; Moso; Sentani; Tawarin; Timena; Friese 1909: 272, 273 (citation, distribution, key to species); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Cyclopen Gebirge; Moaif; Moso; Sentani: Tawarin; Timena; Cockerell 1910: 247 (misidentification, uncertain identity, possibly percincta); AUSTRALIA, Hermannsburg, Finke River (probable misidentification); Friese 1912: 94 (distribution); PAPUA NEW GUINEA, Sepik-Bivak; Strand 1915: 139 (taxonomy, uncertain identity); Friese 1915: 4 (distribution); AUSTRALIA, Derby (probably misidentification); Kimberley (probably misidentification); INDONESIA (Irian Jaya)/PAPUA NEW GUINEA, Bewani Fluss; Cyclop Gebirge; Finschhafen; Friedrich Wilhelmshafen; Hussin; Jasa; Moaif; Moso; Njao; Sentani; Sepik-Bivak; Simbang; Stephansort; Tawarin; Timena; Zoutbron; Floral record: Eucalyptus "staunus"; Friese 1917: 9 (distribution); AUSTRALIA, Kimberley (probable misidentification); Cockerell 1929e: 242 (distribution, taxonomy); PAPUA NEW GUINEA, Mt. Lamington; Schwarz 1932: 245; Rayment 1932f: 106 (taxonomy); Rayment 1935: 735 (key to species); Schwarz 1948: 117; Moure 1961: 197 (morphology, systematic position); Michener 1961: 10, 11-15, 38, 42, 44 (aggressiveness, nest); INDONESIA, Irian Jaya/ PAPUA NEW GUINEA, Bubia near Lae; Kapagere, Central dist.; Kerema, Gulf dist.; Port Moresby; Michener 1965: 230 (citation); Wille & Michener 1973: 10, 18, 42, 53, 64, 91, 94, 96, 97, 98, 99, 267 (nest); Sakagami et al. 1973: 180, 181 (comparative note); Michener 1974: 341 (nest); Wille 1983: 48, 50 (nest); Camargo 1988: 372 (distribution); PAPUA NEW GUINEA, Astrolabe Bay; Huon Golf; Simbag, Stephansort; Michener 1990: 97, 110, 133, 134 (systematic position); Michener & Roubik 1993: 262 (comparative note, morphology); Cardale 1993: 319 (distribution (identity uncertain)); AUSTRALIA, N (probable misidentification); Drumond et al. 1995: 43 (citation, comparative behavior); Michener 2000: 780; Michener 2001: 232 (morphology); Hohmann 2003: (communication, nest); Michener 2007a: 805. Austroplebeia cockerelli (Rayment, 1930) Trigona cockerelli Rayment 1930: 57-59: Holotype (ANIC, worker): examined, "TYPE", "Borroloola, N.T. / 25.8.1911 / G. F. Hill", "Trigona / cockerelli Raym / [below] Euryglossa strigata Ckll", "AUST. NAT. /



9


INS. COLL.", paratype (ANIC (1)) (illustration, taxonomy); Type locality: AUSTRALIA "Borroloola, North Australia (Gerald Hill, 25th September, 1911)" (worker); Rayment 1932c: (behavioral comparison); Rayment 1932f: 106 (comparative note); Rayment 1932f: 107 (taxonomy); Rayment 1935: 735 (key to species); Moure 1961: 197 (possible synonymy); Michener 1961: 38 (comparative note); Michener 1965: 230 (citation); Adams & Lawson 1988: (pollination); Cardale 1993: 319 (distribution, floral record, list); AUSTRALIA, N Gulf; Northern Territory; Floral record: Cassia; Eucalyptus; Dollin 1997: (characteristics); Klumpp 2007: 5. Austroplebeia essingtoni (Cockerell, 1905) Trigona essingtoni Cockerell 1905: 220-221: Holotype (BMNH 17b.1138, worker): examined (head missing), "Syntype" (blue border), "B.M. TYPE / HYM. / 14B.1138", "Pt. Essing / ton / 42-", "Trigona / essingtoni Ckll / TYPE"; paratype (BMNH, labeled cotype (1)) (taxonomy); Type locality: AUSTRALIA "Port Essington, N. Australia, with no. 42.1" (2 workers); Cockerell 1930b: 150 (taxonomy); AUSTRALIA, Port Essington; Rayment 1930: 59 (comparative note); Rayment 1932f: 106 (taxonomy); Rayment 1935: 735 (key to species); Rayment 1953: (distribution); Moure 1961: 196-197 (morphology, systematic position, variation); Michener 1965: 230 (citation); Evans & Matthews 1973: (predator (Bembix flavipes: Crabronidae: Hymenoptera)); Kistner 1982: 162 (citation, predator (Bembix flavipes: Crabronidae: Hymenoptera)); Evans et al. 1982: 78 (predator (Bembix flavipes: Crabronidae: Hymenoptera)); Dollin & Dollin 1988: (type locality); Michener 1990: 102, 130, 133 (illustration, morphology, systematic position); Cardale 1993: 319-320 (distribution, list); AUSTRALIA, N coastal; NW coastal; Northern Territory; Western Australia; Welch 1995: (rock art); Dollin 1997: (characteristics); Crane 1999: 89; Klumpp 2007: 5. Austroplebeia ornata (Rayment, 1932) Trigona cockerelli ornata subsp. nov. Rayment 1932f: 107: Lectotype (MVMA, worker): here designated, "4", "TYPE", "Trigona / cockerelli Raym. / ornata / C. York Q.", "SYNTYPE / T-11575 / Trigona / cockerelli / ornata", "MUS. VIC. / ENTO 2008-8 L"; Paralectotypes (MVMA (3)) (taxonomy (as manuscript name)); Type locality: AUSTRALIA "Cape York, Queensland (Donald Thomson)" (workers); Rayment 1935: 735 (key to species, taxonomy (described as new species again)); Moure 1961: 197 (possible synonymy); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution, list); AUSTRALIA, NE coastal; Queensland; Dollin 1997: (characteristics); Klumpp 2007: 5. Austroplebeia percinta (Cockerell, 1929) Trigona cincta subsp. percinta nov. Cockerell 1929e: 242: Syntypes (ZMHB, 4 workers): examined, labeled by Friese as Trigona cincta? (comparative note (corrected identification of Cockerell 1910)); Type locality: AUSTRALIA "Hermannsberg, Finke River" (unknown); Cockerell 1930b: 150 (taxonomy); Rayment 1932f: 106 (taxonomy); Rayment 1935: 735 (key to species); Schwarz 1948: 92 (citation); Moure 1961: 197 (systematic position); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution, list); AUSTRALIA, Lake Eyre basin, Northern Territory; Dollin 1996b: (nest); Dollin 1997: (characteristics); Green et al. 2001: 91 (microsatellite); Klumpp 2007: 5. Austroplebeia symei (Rayment, 1932) Trigona symei Rayment 1932f: 106: Holotype (ANIC, worker): examined, "TYPE", "Trigona / symesi Raym


RASMUSSEN


/ NT. QUEENS", "AUST. NAT. / INS. COLL." (taxonomy (as manuscript name)); Type locality: AUSTRALIA "North Queensland" (worker); Rayment 1935: 735, 737-738 (key to species, taxonomy (described as new species again)); MacPherson & Rupp 1936: (pollination); Floral record: Dendrobium speciosum; Moure 1961: 197 (possible synonymy); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution, floral record, list); AUSTRALIA, Queensland; Floral record: Dendrobium; Dollin 1997: (characteristics); Drumond et al. 1999: (oviposition); Camargo et al. 2000: 9 (comparative notes, taxonomy); Drumond et al. 2000: 333 (citation); Heard & Dollin 2000: (bee-keeping); Green et al. 2001: 91 (microsatellite); Palmer et al. 2002: (maternity); Dollin 2002a: (bee-keeping); Costa et al. 2003: (systematics); Tóth et al. 2004: 6 (citation); Rasmussen & Cameron 2007: 29, 31, 33, 34 (systematics); Klumpp 2007: 5. Austroplebeia websteri (Rayment, 1932) Trigona websteri Rayment 1932f: 105: Holotype (ANIC, worker): examined, "TYPE", "Trigona / websteri Raym. / WYNDHAM, W.A. / [below] 25/1/1931", "AUST. NAT. / INS. COLL."; paratype (ANIC (1)) (taxonomy); Type locality: AUSTRALIA "Wyndham, North-western Australia (U.N. Webster, M.D., 25th January, 1931)" (worker); Rayment 1932f: 107 (taxonomy); Rayment 1935: 735 (key to species); Moure 1961: 197 (possible synonymy); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution); AUSTRALIA, N coastal; Western Australia; Dollin 1997: (characteristics); Klumpp 2007: 5. Austroplebeia indet. Michener 1965: 230; Camargo & Wittmann 1989: 226 (biogeography); Dollin et al. 1997: (comparative note); Camargo et al. 2000: 22 (comparative notes, taxonomy); Heard & Dollin 2000: (bee-keeping); Maynard et al. 2004: 319 (citation); Roubik 2006: 126, 128 (citation, nest); Cortopassi-Laurino et al. 2006: (bee-keeping); Slaa et al. 2006: 305 (citation).

Geniotrigona Geniotrigona incisa (Sakagami & Inoue, 1989) 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, native collector, 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). Geniotrigona lacteifasciata (Cameron, 1902) Trigona borneënsis Friese 1933a: 46: Lectotype (ZMHB, worker): here designated, "Borneo / Sanggau / 247-32", "Trigona / borneensis / 1925 Friese det. Fr."; paralectotypes (DEI (4 (2 on 1 pin), ZMHB (2)); possible additional types (AMNH, USNM) (taxonomy); Type locality: INDONESIA "Sanggau (Borneo) 24. Juli 1932" (several workers); Schwarz 1937: 317, 318, 328, plate 3, 7; Schwarz 1939a: 105, 106 (synonymy); Moure 1961: 213 (morphology, systematic position); Camargo 1988: 370 (comparative notes, synonymy); Sakagami et al. 1990: 127 (synonymy). Trigona lacteifasciata Cameron 1902: 131: Holotype (BMNH 17b.1132). Probably a distinct taxon of tho-



11


racica from the island of Borneo (taxonomy); Type locality: MALAYSIA "Borneo" (unknown); Cameron 1908: 193 (key to species); Cockerell 1919c: 78 (comparative note); Cockerell 1923: 242 (key to species); Schwarz 1937: 283, 291, 317-319*, 326, plate 3, 7 (key to species, taxonomy); Schwarz 1939a: 105, 106 (synonymy); Schwarz 1948: 91; Moure 1961: 213 (systematic position); Sakagami 1975: 56 (distribution, variation); SINGAPORE; THAILAND; Camargo 1988: 370 (distribution, taxonomy); MALAYSIA, Sandakan; Sakagami et al. 1990: 127 (synonymy). Geniotrigona thoracica (Smith, 1857) Trigona ambusta Cockerell 1918: 387: Holotype (BMNH 17b.1131) (distribution, key to species); Type locality: MALAYSIA/SINGAPORE "Sandakan and Singapore" (unknown); Cockerell 1919c: 78 (taxonomy (described in previous paper)); SINGAPORE; Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Cockerell 1929b: 139 (distribution); MALAYSIA, Kuala Lumpur; Cockerell 1930a: 157 (distribution, nest (hollow tree), variation); CAMBODIA, Koh Po; Pattani; Cockerell 1934: 62, fig. 7 (comparative note, illustration (mandible)); Schwarz 1937: 317, 318, 327; Schwarz 1939a: 105 (synonymy); Moure 1961: 213 (synonymy); Camargo 1988: 370 (synonymy); Sakagami et al. 1990: 127 (synonymy). Trigona thoracica Smith 1857: 50: Type (BMNH 17b.1181) (taxonomy); Type locality: SINGAPORE "Singapore" (worker); Smith 1871: 395 (distribution); SINGAPORE; Dalla Torre 1896: 584 (citation); Bingham 1897: 561, 564 (distribution, key to species, taxonomy); MYANMAR, Tenasserim; SINGAPORE; Bingham 1905: 58 (citation, distribution); Friese 1909("1908"): 358 (comparative note); Cockerell 1919c: 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 (systematic position); 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; Floral record: Xerospermum intermedium; Sakagami et al. 1983b: 2, 4, 13-15, 16, 20 (honey, nest (in Shorea laevis: Dipterocarpaceae)); Sakagami et al. 1985: (distribution, key to species); Baker & DelfinadoBaker 1985: 231 (inquiline (Neocypholaelaps phooni: Ameroseiidae: Acari)); Camargo 1988: 370 (distribution, 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: (distribution, 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: (predator (Oecophylla: Formicidae: Hymenoptera)); Michener & Roubik 1993: 260 (comparative note, morphology); 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, morphology); Heard & Dollin 1998: (citation); Heard & Dollin 1998: 11 (pollination); Floral record: Averrhoa carambola; 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 & Saiboon 2000: (distribution, nest (as thorasica!)); THAILAND, Songkhla; Liow et al. 2001: 184; Nagamitsu & 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


RASMUSSEN


al. 2006: 297 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics); Michener 2007a: 826. Geniotrigona indet. Roubik 2006: 130 (nest).

Heterotrigona Heterotrigona bakeri (Cockerell, 1919) Trigona bakeri Cockerell 1919c: 78, 79: Holotype (USNM 29468, worker): examined, "Island of / Penang / Baker", "9069", "Type No. / 29468" (head glued to label), "Trigona / bakeri / Ckll TYPE" (comparative notes, key to species); Type locality: MALAYSIA "Penang Island (Baker 9068)" (worker); Cockerell 1930a: 157 (distribution, floral record, variation); CAMBODIA, Tung Nui; Satul; Floral record: Melastoma malabaricum; Schwarz 1937: 323, 324; Schwarz 1939a: 96 (synonymy); Moure 1961: 203 (morphology, systematic position); Camargo 1988: 370 (distribution, taxonomy); MALAYSIA, Penang; Sakagami et al. 1990: 126 (synonymy); Rasmussen & Cameron 2007: (systematics). Heterotrigona erythrogastra (Cameron, 1902) Trigona erythrogastra Cameron 1902: 129-130: Holotype (BMNH 17b.1130); 1 paratype (BMNH?) (comparative notes, taxonomy); Type locality: MALAYSIA "Sarawak (R. Shelford)" (worker); Cameron 1908: 193, 194 (key to species); Cockerell 1920a: 115 (comparative note); Cockerell 1923: 242 (key to species); Schwarz 1937: 291, 319-320*, 323, 326, 327, plate 3, 6 (key to species, taxonomy); Floral record: Melastoma; Schwarz 1939a: 84, 85, 86, 90, 96, 100-101* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 203 (systematic position); Baltazar 1966: 408 (distribution); PHILIPPINES, Palawan; Jander & Jander 1970: 362, 363, 367 (orientation); Sakagami 1975: 55* (distribution, variation); MALAYSIA; Wille 1979b: 265 (comparative note); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Appanah et al. 1986: (foraging, nest); Floral record: Calophyllum; Elaeis; Elaeocarpus; Sindora; Xanthophyllum; Camargo 1988: 370 (distribution, taxonomy); MALAYSIA, Borneo, Sandakan; Sakagami & Inoue 1989: 617, 619 (comparative note); Michener 1990: 120, 126, 127 (systematic position); Sakagami et al. 1990: 131, 135 (distribution, key to species); Michener & Roubik 1993: 260 (comparative note, morphology); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Bruijn & Sommeijer 1997: 44 (citation (as erythrogaster!)); Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Heard 1999a: 187 (citation (as erythrogaster!)); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76, 78, 79, 81, 83, plate 6 (distribution, foraging); Rasmussen & Cameron 2007: (systematics). Trigona luteiventris Friese 1909("1908"): 354, 358, fig. 15-5: Lectotype (ZMHB, worker): here designated, "Malakka / Perak / 1912", "Trigona / luteiventris / 1904 Friese det. Fr / n Fr.", "Type" (red label), "Coll. / Friese"; paralectotype (ZMHB (1)); possible additional types (AMNH) (taxonomy); Type locality: MALAYSIA/PHILIPPINES "Perak (Malaka)"; "Palawan" (workers); Cockerell 1919c: 77 (key to species); PHILIPPINES, Palawan; MALAYSIA, Perak; Cockerell 1920b: 229 (distribution); PHILIPPINES, Palawan; Schwarz 1932: 245; Schwarz 1937: 319, 320, 327, plate 3, 6; Schwarz 1939a: 86, 100, 101 (synonymy); Moure 1961: 203 (systematic position); Baltazar 1966: 408 (synonymy). Trigona sandacana Cockerell 1919b: 242-243: Holotype (BMNH 17b.1129) (comparative notes, taxonomy);



13


Type locality: MALAYSIA "Sandakan, Borneo (Baker, 9593)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 319, 327; Schwarz 1939a: 100, 101 (synonymy); Moure 1961: 203 (synonymy); Camargo 1988: 370 (synonymy (as sandakana!)). Heterotrigona itama (Cockerell, 1918) Trigona breviceps Cockerell 1919b: 244: Holotype (BMNH 17b.1123, worker): examined, "Type" (red border), "Sandakan / Borneo / Baker", "Trigona / breviceps / Ckll TYPE", "B.M. TYPE / HYM. / 17B.1123", "9591" (comparative notes, taxonomy); Type locality: MALAYSIA "Sandakan, Borneo (Baker, 9591)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 292, 322-325*, 327, plate 3, 6 (key to species, taxonomy); Schwarz 1939a: 84, 96, 97, 98 (synonymy); Moure 1961: 203 (systematic position); Sakagami et al. 1990: 126 (synonymy). Trigona itama Cockerell 1918: 387: Holotype (USNM 29471, worker): examined (distribution, key to species); Type locality: SINGAPORE "Singapore" (unknown); Cockerell 1919b: 241-242, 244 (taxonomy, variation); MALAYSIA, Penang; Cockerell 1919c: 78, 79 (comparative notes, key to species); Cockerell 1919a: 121 (distribution, taxonomy, variation); INDONESIA, Pelaboean Ratoe, Java; SINGAPORE; Cockerell 1920a: 115 (comparative note); Cockerell 1925a: 492 (variation); MALAYSIA, Borneo, Sandakan; Cockerell 1927: 541 (distribution); MALAYSIA, Selangor, Pahang, near Genting, Sempat, Hill stream; Schwarz 1937: 283, 292, 319, 322-325*, 327, plate 3, 6 (key to species, taxonomy); Floral record: Melastoma; Schwarz 1939a: 84, 85, 86, 90, 92, 95, 96-100*, 127, 133 (distribution, key to species, taxonomy); Floral record: Mimosa pudica; Tephrosia candida; Sandhouse 1943: 558 (citation); Schwarz 1948: 118, 119; Sakagami 1959: 119 (distribution); SINGAPORE; Moure 1961: 201, 203 (morphology, systematic position); Wille 1961: 117; Yoshikawa et al. 1969: 168, 172, 173, 174 (distribution, nest); MALAYSIA, Cameron Highland, Pahang; Fraser's Hill, Selagor-Pahang; George Town, Penang; Sabah, Jesselton; Sabah, Touaran; Sarawak, Kuching; Wille & Michener 1973: 14, 48 (nest); Sakagami 1975: 55* (distribution, variation); MALAYSIA; THAILAND; Wille 1979b: 265 (comparative note); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1983b: 2-4, 13, 19, 20 (nest); Sakagami et al. 1985: (distribution, key to species); Inoue et al. 1985a: (aggressiveness, foraging, nest); Kiew 1985: (luring (unwashed shorts)); Inoue et al. 1985b: 45, 47, 48 (foraging); Baker & Delfinado-Baker 1985: 231 (inquiline (Neocypholaelaps phooni: Ameroseiidae: Acari)); Atmowidjojo & Adisoemarto 1986: 227, 229 (pollination); Floral record: Indigofera; Sakagami & Inoue 1987: 625 (comparative note); Uji 1987: (pollination); INDONESIA, E Kalimantan; Floral record: Nephelium lappaceum; Camargo 1988: 370 (distribution, taxonomy); MALAYSIA, Kuala Lampur; SINGAPORE; Sakagami & Inoue 1989: 610, 613, 614, 616, 617 (morphology); Roubik 1989: 105, 133, 137, 156, 158; Michener 1990: 105, 120, 122, 125, 126, 127 (illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; Java; MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest (hollow trunks of clove (Eugenia aromatica: Myrtaceae); coconut; crevices)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest (tree cavities and others)); Michener & Roubik 1993: 260 (comparative note, morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cunha 1994: (systematics); Tezuka & Maeta 1995: (foraging, pollen robbing); Roubik 1995: 151 (antropogenic nest); Camargo & Moure 1996: 109 (taxonomy (type series)); Salmah et al. 1996: (reproduction); Roubik 1996: 61, 62 (distribution); BRUNEI, Kuala Belalong; Michener 1997: 29 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Heard 1999a: 196 (citation); Michener 2000: 801802; Chinajariyawong & Saiboon 2000: (distribution, nest); THAILAND, Songkhla; Amano et al. 2000: (citation); Liow et al. 2001: 184; Costa et al. 2003: (systematics); Eltz 2004: 322 (abundance);


RASMUSSEN


Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Chinh et al. 2005: 368 (citation); Klakasikorn et al. 2005: 6 (citation); Roubik 2006: 131 (nest); Rasmussen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Averrhoa carambola; Cassia fistula; Michener 2007a: 826-827; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note). Heterotrigona indet. Roubik 2006: 127, 132 (nest).

Homotrigona Homotrigona aliceae (Cockerell, 1929) Trigona aliceæ Cockerell 1929b: 139, 140: Holotype (BMNH 17b.1116) (comparative note, taxonomy); Type locality: THAILAND "Siam: On trail in jungle between Pahtoop mountain and Nan, Jan. 9, 1928 (Alice Mackie)" (worker); Schwarz 1937: 305, 306; Schwarz 1939a: 85, 92, 104* (distribution, key to species, taxonomy); Schwarz 1948: 118; Schwarz 1951: 63 (distribution); CAMBODIA, Angkor; Moure 1961: 201 (systematic position); Sakagami & Yoshikawa 1961: 434-435 (distribution, taxonomy); THAILAND, Doi Inthanon; VIETNAM, Kompong Cham; Kamphangpet; Sakagami 1963: (morphology (as fimbriata)); Sakagami 1975: 52, 54, 55 (distribution, variation); INDONESIA, Sumatra; THAILAND; Sakagami et al. 1990: 127 (synonymy); Klakasikorn et al. 2005: 6 (citation). Trigona kusutkana Dover 1929: 62-64: Holotype (not in ZRC); paratype (not in ZRC) (taxonomy, variation); Type locality: MALAYSIA "Kedah Peak, 3200 ft., December, 1915" (holotype worker, paratype worker); Schwarz 1939a: 104* (distribution, key to species, taxonomy); Moure 1961: 201 (systematic position); Sakagami 1975: 54 (synonymy); Sakagami et al. 1990: 127 (synonymy). Homotrigona anamitica (Friese, 1909) Trigona anamitica Friese 1909("1908"): 358-359, fig 15-4: Lectotype (ZMHB, worker): here designated, "Süd-Annam / Xom-Gom / Februar / H. Fruhstorfer", "Trigona / anamitica / 1904 Friese det. / Fr."; paralectotype (ZMHB (1)) (comparative note, taxonomy); Type locality: VIETNAM "Xom-Gom, SüdAnnam, Februar 1901 (Fruhstorfer leg.)" (3 workers); Schwarz 1937: 304, 307; Schwarz 1939a: 85, 92, 103-104* (distribution, key to species, taxonomy); Moure 1961: 201 (systematic position); Sakagami 1975: 52, 54 (distribution, variation); MALAYSIA, Borneo; VIETNAM; Camargo 1988: 371 (synonymy); Sakagami et al. 1990: 127 (synonymy). Trigona melanotricha Cockerell 1918: 386, 387: Holotype (BMNH 17b.1095) (key to species, comparative notes, key to species, taxonomy); Type locality: MALAYSIA "Sandakan, Borneo (Baker 9222)" (worker); Cockerell 1919b: 243 (comparative notes); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 289, 305, 306, 320, 327 (key to species); Schwarz 1939a: 103, 104 (synonymy); Moure 1961: 201 (systematic position); Sakagami 1975: 52 (synonymy); Camargo 1988: 370371 (distribution, synonymy, taxonomy); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1990: 127 (synonymy); Leonhardt et al. 2007: (foraging); Floral record: Ardisia elliptica. Homotrigona fimbriata (Smith, 1857) 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-



15


ell 1918: 386 (comparative notes); Cockerell 1919b: 243 (comparative notes); Cockerell 1929b: 139 (comparative note); Schwarz 1939a: 102 (synonymy); Moure 1961: 201 (synonymy). Trigona fimbriata Smith 1857: 52: Type (BMNH 17b.1182) (taxonomy); Type locality: SINGAPORE "Singapore" (worker); Smith 1871: 396 (distribution); SINGAPORE; Gribodo 1893: 263 (comparative note); Dalla Torre 1896: 578 (citation); Cockerell 1919c: 79 (distribution); Cockerell 1923: 241 (key to species); Schwarz 1937: 283, 289, 304-307*, 320, 326, 327, plate 2, 4 (key to species, taxonomy); Schwarz 1939a: 84, 85, 90, 92, 102-103*, 104, 106, 132 (distribution, key to species, taxonomy); Schwarz 1948: 73, 94; Moure 1961: 200, 201, 215 (systematic position); Sakagami & Yoshikawa 1961: 435 (comparative note); Sakagami 1963: (see aliceae); Sakagami 1975: 52-55* (distribution, morphology, variation); LAOS; MALAYSIA; SINGAPORE; THAILAND; VIETNAM; Sakagami et al. 1983b: 5 (nest); Sakagami et al. 1985: (distribution, key to species); Sakagami & Khoo 1987: 207 (comparative note); Camargo 1988: 371 (synonymy); Roubik 1989: 133, 135, 417; Michener 1990: 105, 120, 125, 126 (systematic position); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra; MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Roubik 1993: 541 (illustration, nest); Michener & Roubik 1993: 260 (comparative note, morphology); Cunha 1994: (systematics); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: Santiria laevigata; Michener 1997: 29 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 127; Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 802; Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Green et al. 2001: 91 (microsatellite); Eltz et al. 2002a: 30 (distribution, foraging, nest, population); MALAYSIA, Danum Valley; Sepilok; Deramakot; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); 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, 80, 82, 83, 84, 85, plate 9 (distribution, foraging); Klakasikorn et al. 2005: 3, 6 (floral record); Floral record: Tectona grandis; Roubik 2006: 131 (nest); Jongjitvimol et al. 2006: (foraging); Jongjitvimol & Wattanachaiyingcharoen 2006: (foraging); THAILAND, Phitsanulok; Floral record: Butea monosperma; Cassia bakeriana; Erythrina stricta; Fernandoa adenophylla; Ipomoea aquatica; Lagerstroemia calyculata; Lagerstroemia macrocarpa; Lagerstroemia tomentosa; Merremia vitifolia; Mimosa pigra; Mimosa pudica; Ruellia tuberose; Senna siamea; Thunbergia laurifolia; Tridax procumbens; Zingiber officinale; Rasmussen & Cameron 2007: (systematics); Raffudin & Crozier 2007: 544, 545, 546 (behavior); Michener 2007a: 826-827; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note). Trigona flavistigma Cameron 1902: 130: Holotype (BMNH 17b.1097) (taxonomy); Type locality: MALAYSIA "Kuching, Sarawak" (male); Cameron 1908: 193 (key to species); Cockerell 1923: 241 (key to species); Cockerell 1929b: 139 (comparative note); Schwarz 1937: 304, 305, 306, 307, 326; Schwarz 1939a: 102, 103 (synonymy); Moure 1961: 201 (systematic position); Sakagami 1975: 52 (synonymy); Sakagami et al. 1990: 127 (synonymy). Trigona versicolor Friese 1909("1908"): 358, 359, fig. 15-1: Lectotype (ZMHB, worker): here designated, "Tandjong / SO.-Borneo / coll.Speyer", "Trigona / versicolor / 1904 Friese det. / n. Fr.", "Type" (red label) (comparative note, taxonomy); Type locality: MALAYSIA "Malaka (Perak); "SO-Borneo (Tandjong)" (2 workers); Schwarz 1937: 304, 306, 327, plate 4; Schwarz 1939a: 102, 103, 104 (synonymy); Schwarz 1948: 118; Moure 1961: 201 (synonymy); Sakagami 1975: 52 (synonymy); Sakagami et al. 1990: 127 (synonymy).


RASMUSSEN


Homotrigona lutea (Bingham, 1897) Trigona ferrea Cockerell 1929b: 139-140: Holotype (BMNH 17b.1117) (comparative note, taxonomy); Type locality: THAILAND "Siam: Mekami River, Feb. 3, 1928 (Cockerell)" (worker); Schwarz 1937: 305, 306; Schwarz 1939a: 85, 92, 103 (distribution, key to species, taxonomy); Moure 1961: 201 (synonymy); Sakagami 1975: 52, 54, 55 (distribution, variation); MALAYSIA; THAILAND; Sakagami et al. 1990: 127 (synonymy); Klakasikorn et al. 2005: 6 (citation). Melipona lutea Bingham 1897: 559, 560, 564: Type (BMNH 17b.1122) (distribution, illustration, key to species, taxonomy; notice this is no longer a senior homonym of Trigona mocsaryi lutea Friese 1903, see Camargo & 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 (taxonomy); Type locality: INDIA "Testa bridge, Himalayas, India, 10.1.97 (Sladen)" (worker); Schwarz 1937: 295; Schwarz 1939a: 85, 88, 134, 135 (distribution, key to species, taxonomy); Nogueira-Neto 1949: 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, 750 meters, 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); Floral record: Ficus religiosa; Homalium grandiflorum. Lepidotrigona flavibasis (Cockerell, 1929) Trigona flavibasis Cockerell 1929c: 592: Holotype (AMNH). The ventralis species group (taxonomy); Type locality: THAILAND "Doi Sutep, Siam, Feb. 9 (Alice Mackie)" (worker); Schwarz 1939a: 85, 94, 134135*, 136 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199 (systematic position); Yoshikawa et al. 1969: 174 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang; Sakagami 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 et al. 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



17


part)); Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Jongjitvimol & Wattanachaiyingcharoen 2007: 29 (citation (as flavibis!)). Lepidotrigona hoozana (Strand, 1913), comb. nov. 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 (variation); 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), but as 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' of workers); Sakagami et al. 1985: (key to species); Sakagami & Yamane 1987: (oviposition); Salmah et al. 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 & Sommeijer 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); Friese 1914a: 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 species, 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 species group (taxonomy); Type locality: MALAYSIA "Kuching, Sarawak" (worker); Smith 1857: 50 (as ventralis, in part); MALAYSIA, "Malacca (Mt. Ophir)"; Cameron 1908: 193 (key to species (as latibalteata!)); 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). Lepidotrigona latipes (Friese, 1900) Trigona latipes Friese 1900: 384: Holotype (ZMHB, worker): examined, "India / Singapore / 1890", "Trigona / latipes / 1909 Friese det. / Fr.", "Type" (red label), "Coll. / Friese". The nitidiventris species group (taxonomy); Type locality: SINGAPORE "Malacca (Singapore)" (1 worker); Schwarz 1937: 293; Schwarz 1939a: 85, 94, 134* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199 (systematic position); Sakagami 1975: 65 (citation).


RASMUSSEN


Lepidotrigona nitidiventris (Smith, 1857) Trigona fulvopilosella Cameron 1908: 193, 194: Syntypes (BMNH 17b.1101, 2 workers) (key to species, taxonomy); Type locality: MALAYSIA "Kuching, May and October (John Hewitt)" (worker); Schwarz 1937: 285, 287, 293-294*, 295, 296, 326 (key to species, taxonomy); Schwarz 1939a: 133 (synonymy); Moure 1961: 199 (possible synonymy); Sakagami 1975: 65 (citation). Trigona nitidiventris Smith 1857: 50-51: Type (OUMNH (=Wilson Saunders collection)). The nitidiventris species group (taxonomy); Type locality: MALAYSIA "Malacca (Mount Ophir)" (worker); Smith 1871: 395 (distribution); MALAYSIA, Malacca; Dalla Torre 1896: 582 (citation); Bingham 1905: 58 (citation, distribution); Cockerell 1920a: 115 (comparative note); Cockerell 1923: 241, 242 (key to species); MALAYSIA, Mt. Ophir; Schwarz 1937: 292, 293, 296, 327 (key to species); Schwarz 1939a: 85, 94, 132, 133*, 134 (distribution, key to species, taxonomy); Sandhouse 1943: 543 (citation); Schwarz 1948: 118; Moure 1961: 197, 199 (systematic position); Baltazar 1966: 407 (distribution); PHILIPPINES, Palawan; Sakagami 1975: 64-65* (distribution, variation); MALAYSIA; THAILAND; Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Sakagami & Yamane 1987: 25 (comparative note); Uji 1987: (pollination); INDONESIA, E Kalimantan; Floral record: Nephelium lappaceum; Michener 1990: 121, 122 (systematic position); Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Yong 1993: (biochemistry); Cunha 1994: (systematics); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 434, 436 (pollination); Floral record: Santiria laevigata; Michener 1997: 32 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 802; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Michener 2007a: 827; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note). Lepidotrigona palavanica (Cockerell, 1915) Trigona palavanica Cockerell 1915: 2-3: Holotype (BMNH 17b.1120). The nitidiventris species group (comparative notes, taxonomy); Type locality: PHILIPPINES "P. Princesa, Palawan (Baker coll 3839)" (worker); Cockerell 1919c: 77 (key to species); PHILIPPINES, Palawan, Puerto Princesa; Cockerell 1920b: 229 (distribution); PHILIPPINES, Palawan; Schwarz 1939a: 85, 86, 133 (distribution, key to species, taxonomy); Moure 1961: 199 (systematic position); Baltazar 1966: 407 (synonymy). Lepidotrigona terminata (Smith, 1878) Trigona fulvomarginata Cockerell 1919c: 78: Holotype (BMNH 17b.1083) (key to species); Type locality: MALAYSIA "Penang" (worker); Cockerell 1929c: 591 (comparative note); CAMBODIA, Angkor; near Nam Geeung; Nan; Pah Meesung Mt.; Pahtoop Mt.; Cockerell 1934: 62, fig. 9 (illustration (mandible)); Schwarz 1937: 295, 296; Schwarz 1939a: 136, 137 (synonymy); Moure 1961: 199 (synonymy). 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); Cockerell 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;



19


Pagden 1957: 17 (comparative note); Sakagami 1960: 146 (distribution, variation); CAMBODIA, Snoul; Stung Treng; Moure 1961: 199 (systematic position); Wille 1961: 117, 121; Sakagami & Yoshikawa 1961: 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; Sakagami 1978: 239 (comparative note); Sakagami et al. 1983b: 2, 15-19, 20 (nest); Sakagami & Yamane 1984: 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 (illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; 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 & Roubik 1993: 260 (comparative note, morphology); Yong 1993: (biochemistry); Yamane et al. 1995: 294 (synonymy); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 434, 436 (pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 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: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Klein et 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); Schwarz 1939a: 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); Sakagami et al. 1990: (distribution, key to species, variation); INDONESIA, Sumatra; Salmah et al. 1990: (distribution, 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-


RASMUSSEN


lis species group (taxonomy); Type locality: MALAYSIA "Borneo (Sarawak)"; "Malacca (Mt. Ophir)" (worker); Smith 1871: 395 (distribution); MALAYSIA, Borneo; Malacca; Gribodo 1891: 109 (comparative note); Dalla Torre 1896: 584 (citation); Bingham 1897: 560, 562-563 (distribution, key to species, taxonomy); MALAYSIA, Borneo, Malacca; MYANMAR, Tenasserim; Strand 1913b: 66 (morphology); Cockerell 1915: 3 (comparative notes); Cockerell 1919c: 79 (comparative note); Cockerell 1923: 241 (key to species); Cockerell 1929c: 591 (comparative note); Friese 1933b: 147 (comparative note); Schwarz 1937: 282, 283, 284, 287, 294-295*, 296, 326, plate 3, 7 (key to species, taxonomy); Schwarz 1939a: 85, 94, 134*, 135, 136, 139 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199 (systematic position); Sakagami 1975: 66, 67-75* (distribution, morphology, variation); CAMBODIA; LAOS; MALAYSIA; THAILAND; VIETNAM; Sakagami 1978: 239 (comparative note); Sakagami 1982: 400, 412 (reproduction); Sakagami & Yamane 1984: 38, 47; Sakagami et al. 1985: (distribution, key to species); Sakagami & Yamane 1987: 17, 25 (comparative note); Michener 1990: 122 (systematic position); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDIA; INDONESIA, Java; Sumatra; MALAYSIA; MYANMAR; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener & Roubik 1993: 260 (comparative note, morphology); Yong 1993: (biochemistry); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 434, 435, 436 (pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 110, 127 (distribution (as ventra! in part)); Nagamitsu et al. 1999: (floral records (not by species), foraging); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Wu 2000: 386, 387-388 (key to species, morphology); Liow et al. 2001: 184; Costa et al. 2003: (systematics); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Velthuis et al. 2005: 171 (citation); Nagamitsu & Inoue 2005: 76, 82, 83, 84, 85, plate 9 (distribution, foraging); Chinh et al. 2005: 363, 364, 366, 368-370, 371 (nest (see flavibasis)); VIETNAM, Cucphuong National Forest, Red river delta; Chinh & Sommeijer 2005: (see flavibais); Roubik 2005: 96 (distribution, food niche); Klakasikorn et al. 2005: 6 (citation (as ventalis!)); SantosFilho et al. 2006: 208, 217 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Sung et al. 2006c: (see hoosana); Rasmussen & Cameron 2007: (systematics); Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note). Lepidotrigona indet. Drumond et al. 1996a: 399 (behavioral comparison); Roubik 2006: 127 (nest). "Trigona (Lepidotrigona) ventralis" nigribasis forma nov. Sakagami 1975: 71: Nomen nudum. The name is unavailable for taxonomic purposes, as it must be deemed to be infrasubspecific (ICZN, art. 15.2 and 45.6.3.) (variation); VIETNAM. Melipona tunneli Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as no description or definition of the taxon was provided (ICZN, art. 13.1.1). The statement that it "is different from the other two in appearance as it possesses somewhat yellow spots in the abdomen", however, could make it a nomen dubium, although most likely then a junior synonymon of arcifera (common name (ngap siwor), honey, nest).

Lisotrigona Lisotrigona cacciae (Nurse, 1907) Melipona cacciæ Nurse 1907: 619: Lectotype (BMNH 17b.1103, worker); Type locality: INDIA "Hoshang-



21


abad, Central Provinces" (workers); Moure 1961: 194, 195 (lectotype designation (see Engel 2000), morphology, 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 hands and 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, morphology); LAOS; THAILAND; VIETNAM; Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, 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); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra; LAOS, MALAYSIA; THAILAND; VIETNAM; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Oldroyd et al. 1992: 5 (pollination); THAILAND, Chanthaburi; Floral record: Archontophoenix alexandrae; Michener & Roubik 1993: 257, 258 (comparative note, morphology); Nagamitsu & Inoue 1997: 436 (pollination); Floral record: Santiria laevigata; Nagamitsu et al. 1999: (floral records (not by species), foraging); Engel 2000: (synonymy); Rajitparinya 2000: (distribution (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); THAILAND, 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: Khe Bo, 19 03N 104 43E, 25-28 April 1998, 123 m, James M. Carpenter" (worker); VIETNAM, Ha Thing, SE Huong 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 (comparative note). Lisotrigona furva Engel, 2000 Lisotrigona furva Engel 2000: 235-236: Holotype (SEMC); 5 paratypes (AMNH, SEMC) (floral record, key to species, taxonomy); Type locality: THAILAND "THAILAND: Nakhon Ratchasima Prov.: Tha Chang, 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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Lisotrigona mohandasi Jobiraj & Narendran 2004: 39, 40-43, fig. 1-4: Holotype ("Calicut", worker); paratypes ("Calicut" (2)) (floral record, key to species, taxonomy); Type locality: INDIA "India, Kerala Forest Research Institute (Peechi), K. Mohandas, 3-IV-2000" (3 workers); Floral record: Tectona grandis. Lisotrigona indet. Engel 2000; Jobiraj & Narendran 2004: 39; Momose & Hamid 2005; Roubik 2006: 127, 128 (nest).

Lophotrigona Lophotrigona canifrons (Smith, 1857) Trigona busara Cockerell 1918: 387: Holotype (USNM 29496, worker): examined, "Singapore / Coll. Baker", "9072", Type No. / 29496 / U.S.N.M.", "Trigona / busara / Ckll TYPE" (distribution, key to species); Type locality: MALAYSIA "Sandakan" (unknown); Cockerell 1919c: 78, 79 (comparative notes, key to species, taxonomy); SINGAPORE; Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Cockerell 1925a: 492 (comparative note); Schwarz 1937: 307, 308, 327; Schwarz 1939a: 98, 101 (synonymy); Moure 1961: 206 (synonymy); Camargo 1988: 370 (synonymy). Trigona canifrons Smith 1857: 51: Syntype (OUMNH, 1 worker); additional putative type (BMNH 17b.1183) (taxonomy); Type locality: MALAYSIA "Borneo (Sarawak)" (worker); Smith 1871: 395 (distribution); MALAYSIA, Borneo; Dalla Torre 1896: 576 (citation); Bingham 1897: 560, 562 (distribution, 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 (comparative notes, taxonomy, misidentification, uncertain identity); Ihering 1912: 3 (citation); Friese 1914b: 53, 54, 57 (identification, previous misidentification); Cockerell 1918: 385 (comparative notes); Cockerell 1919b: 244 (comparative note); Prell 1919: (nest (in human skull)); Cockerell 1922a: 279 (misidentification); Cockerell 1923: 241, 242 (key to species); MALAYSIA, Sarawak; Cockerell 1929e: 242 (comparative 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 (citation); Yoshikawa et al. 1969: 173 (distribution); MALAYSIA, Kuala Lumpur; Michener 1974: 217; Sakagami 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; Floral record: Nephelium lappaceum; Camargo 1988: 370 (synonymy); Roubik 1989: 105, 193; Michener 1990: 88, 97, 105, 120, 126, 127 (systematic position); Sakagami et al. 1990: (distribution, key to species); 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), population); Johnson & Appanah 1990: 586 (predator (Oecophylla: Formicidae: Hymenoptera) (as carnifrons!)); Roubik 1992: 345 (citation); Michener & Roubik 1993: 260, 264 (comparative note,



23


morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cunha 1994: (systematics); Yamane et al. 1995: 294 (synonymy); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Dollin et al. 1997: 861 (comparative note); Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: Santiria laevigata; Michener 1997: 34 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Momose et al. 1998: (pollination); Floral record: Dryobalanops aromatica; Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 780, 800-801; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: 309 (citation); Biesmeijer & Slaa 2004: 148 (citation, foraging); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Roubik 2006: 127, 131 (nest); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics); Michener 2007a: 804, 824826.

Odontotrigona Odontotrigona haematoptera (Cockerell, 1919) Trigona haematoptera variety dulitae Schwarz 1937: 282, 283, 286, 291, 320-321*, 328, plate 2, 6: Holotype (BMNH 17b.1125); paratypes (AMNH, BMNH (8)) (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Foot of Mt. Dulit, junction of rivers Tinjar and Lejok" (2 workers Aug. 2, 1932; 10 workers Aug 3, 1932); "Mt. Kalulong, 1800 ft." (1 worker); Schwarz 1939a: 85, 90, 102 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 214 (synonymy). Trigona hæmatoptera Cockerell 1919b: 243: Holotype (BMNH 17b.1126) (comparative notes, taxonomy); Type locality: MALAYSIA "Sandakan, Borneo (Baker, 9592)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 282, 383, 286, 291, 320*, 321, 327 (key to species, taxonomy); Schwarz 1939a: 85, 90, 92, 102*, 130 (distribution, key to species, taxonomy); Schwarz 1948: 73, 118; Moure 1961: 213, 214 (systematic position); Sakagami et al. 1985: (distribution); Camargo 1988: 371 (distribution); MALAYSIA, Borneo, Sandakan; Michener 1990: 127 (systematic position); Sakagami et al. 1990: 131 (distribution); Cunha 1994: (systematics); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Michener 1997: 42 (type species); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 801; Eltz et al. 2002a: 30 (distribution, foraging, nest, population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Eltz et al. 2003: (nest, tree preference); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Rasmussen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Ardisia elliptica; Dillenia excelsa; Mallotus; Michener 2007a: 826.

Papuatrigona Papuatrigona genalis (Friese, 1909) Trigona genalis Friese 1909("1908"): 354, 356, 357, fig. 15-2: Lectotype (ZMHB, worker): examined, designated Michener 1990, "Manikion / 16.28.II.03", "Trigona / genalis / 1904 Friese det. / n. Fr.", "Type" (red label), "Coll. Friese", "Zool. Mus. / Berlin"; paralectotype (ZMHB (3)) (distribution, taxonomy); Type locality: INDONESIA (IRIAN JAYA) "Manikion, 14.-28. Februar 1903 (Neu-Guinea)" (several work-


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ers); Friese 1909: 272, 273, 276-277 (citation, key to species); Schwarz 1932: 245; Schwarz 1948: 118; Michener 1961: 23, 42 (aggressiveness, nest); PAPUA NEW GUINEA, Gulf District, Murua River near Kerema; Michener 1965: 231 (citation); Wille & Michener 1973: 14, 24, 48, 59, 70 (nest); Michener 1990: 95, 97, 105, 122, 127, 153-157 (distribution, illustration, lectotype designation, morphology, systematic position); INDONESIA, Irian Jaya, Bodem; Bodem Sarmi area; Klamono oil fields; Waris S of Hollandia; PAPUA NEW GUINEA, Dreikikir; Marua river near Kerema; Northeast New Guinea; One; Sepik; Michener & Roubik 1993: 260 (comparative note, morphology); Snelling 1996: 131 (distribution); Michener 1997: 44 (type species); Michener 2000: 802-803; Michener 2007a: 807.

Pariotrigona Pariotrigona pendleburyi (Schwarz, 1939) Trigona (Hypotrigona) pendleburyi variety klossi Schwarz 1939a: 85, 94, 132*: Holotype (BMNH, no number) (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: Kuala Teku, 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); THAILAND, 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 "Cameron Highlands, Rhododendron Hill, 5000 feet (H. M. Pendlebury)" (workers, holotype, June 19,1923, and June 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, 193194, 195 (morphology, systematic position); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Michener 1990: 96, 100, 105, 129, 131 (illustration, morphology, systematic position); Sakagami 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 & Roubik 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); THAILAND, Songkhla; Michener 2001: (distribution, morphology (male characters)); BRUNEI; INDONESIA, 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.



25


Platytrigona Platytrigona atricornis (Smith, 1865) Trigona atricornis Smith 1865: 94: Holotype (OUMNH, worker): According to Baker (1993) the specimen is labeled " 'N' [New Guinea (Allen); white disc] and 'Trigona atricornis' 'Smith' ". The atricornis species group (taxonomy); Type locality: INDONESIA (IRIAN JAYA) "New Guinea" (worker); Smith 1871: 396 (distribution); "NEW GUINEA"; Gribodo 1893: 263 (comparative note); Dalla Torre 1896: 576 (citation); Friese 1898: 428, 429 (citation, key to species); "NEW GUINEA"; Friese 1898: 429 (comparative note); Friese 1909: 272, 273, 277-278 (citation, key to species); Friese 1915: 4 (distribution); INDONESIA, Irian Jaya, Jasa; Njao; Cockerell 1923: 240 (key to species); Schwarz 1948: 117; Michener 1965: 231 (citation); Michener 1990: 127, 153-156 (distribution, illustration, morphology, systematic position). Platytrigona flaviventris (Friese, 1909) Trigona flaviventris Friese 1909("1908"): 354, 356, 357-358, fig. 15-6: Lectotype (ZMHB, worker): here designated, "Cyclops / geb. / 13.IV.03", "Trigona / flaviventris / 1904 Friese det. / Fr.", "Type" (red label), "Coll. / Friese"; paralectotype (ZMHB (2)). The planifrons species group (taxonomy); Type locality: INDONESIA (IRIAN JAYA) "Cyclopen-Gebirge, 3.-14. April 1903 (New Guinea)" (4 workers); Friese 1909: 272, 275 (citation, key to species); INDONESIA, Irian Jaya, Cyclopen Gebirge; Cockerell 1919b: 243 (comparative notes); Cockerell 1929e: 243 (comparative note); Schwarz 1932: 245; Schwarz 1937: 289, 320 (key to species); Schwarz 1948: 118; Moure 1961: 204 (systematic position); Michener 1965: 231 (citation); Michener 1990: 120, 127, 153 (systematic position); Michener 2000: 800; Hohmann 2003: (communication, nest); Michener 2007a: 824. Platytrigona hobbyi (Schwarz, 1937) Trigona hobbyi Schwarz 1937: 283, 288, 298-300*, 328: Holotype (BMNH 17b.1118). The hobbyi species group (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, 4000 feet, moss forest, 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 (systematic position); Sakagami et al. 1990: 131 (distribution); Michener & Roubik 1993: 260 (comparative note, morphology); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (distribution, 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"; paralectotype (AMNH (1)). The planifrons species group (manuscript name (latigena in litt.), taxonomy); Type locality: INDONESIA "Key-Eilanden (Amboina, Nederland India) durch H. Kühn" (6 workers); Friese 1909("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 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Holl. N.G.; Pauwi; Samberi; Teba; Friese 1912: 94 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, D. Neu Guinea; Pauwi; Sam-


RASMUSSEN


beri; Teba; Strand 1912: (distribution); Friese 1915: 4 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Moaif; Pauwi; Samberi; Teba; Timena; Zoutbron; Schwarz 1948: 118; Michener 1965: 231 (citation); Michener 1990: 120, 127, 153 (systematic position); Michener & Roubik 1993: 260 (comparative note, morphology); Snelling 1996: 131 (distribution); Michener 2000: 800; Hohmann 2003: (communication, nest); Michener 2007a: 824. Platytrigona lamingtonia (Cockerell, 1929) Trigona lamingtonia Cockerell 1929e: 243: Holotype (AMS, worker): examined, "Mt. Lamington / Northern Division / Papua v.1927 / C.T. McNamara", "Trigona / lamingtonia / Ckll TYPE", "HOLOTYPE", "Australian Museum K 238299". The planifrons species group (taxonomy); Type locality: PAPUA NEW GUINEA "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 specimen 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 species group (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 (distribution, key to species); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai; Janour Gebiet; 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); Cockerell 1929e: 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 (citation); 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, not located, 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." (5 workers, 4 males, holotype worker, paratypes); "Koto Atas, Solok, Sumatera Barat" (6 workers); Sakagami & 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



27


et al. 1998: 244-245 (type depositories); Nagamitsu & Inoue 2005: 76 (distribution, foraging). Sundatrigona moorei (Schwarz, 1937) Trigona (Tetragona) matsumurai Sakagami 1959: 120-121: Holotype (SEHU, worker); 1 paratype (SEHU); Type locality: SINGAPORE "Singapore, Sept., 22, 1932, S. Matsumura leg." (2 workers); Sakagami 1975: 56, 57 (synonymy); Sakagami et al. 1990: 127 (synonymy); Tadauchi et al. 1998: 243 (type depositories). Trigona moorei Schwarz 1937: 283, 292, 321-322*, 328, plate 3, 4: Holotype (BMNH 17b.1124); 1 paratype (AMNH) (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, R. Lejok, near sweat and water, Oct. 5, 1932" (2 workers); Schwarz 1939a: 85, 90, 106* (distribution, key to species, taxonomy); Schwarz 1948: 83, 118; Sakagami 1959: 120-121 (morphology); Sakagami 1975: 56-58* (distribution, morphology, variation); MALAYSIA; Sakagami 1978: 239 (comparative note); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1983c: (reproduction); Sakagami et al. 1985: (distribution, key to species); Inoue et al. 1985a: (aggressiveness, foraging, nest (in aerial Crematogaster: Formicidae: Hymenoptera nests)); Inoue et al. 1985b: 45, 47, 48 (foraging); Sakagami & Inoue 1987: 610-617, 618, 619, 620, 621, 622, 623, 624 (morphology); Salmah et al. 1987: (development); Sakagami & Inoue 1989: 615, 617, 618 (morphology); Roubik 1989: 133, 196, 197, 218; Sakagami et al. 1989: (nest); Sakagami 1989: (nest); Michener 1990: 98, 126, 127, 153 (morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; MALAYSIA; Salmah et al. 1990: (distribution, nest (in aerial Crematogaster: Formicidae: Hymenoptera nests)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest (Crematogaster: Formicidae: Hymenoptera nests), population); Roubik 1990c: 663 (citation); Salmah et al. 1991: 199 (comparative note); Inoue & Sakagami 1993: 769 (citation); Salmah et al. 1996: 402, 404, 407, 409, 410 (comparative note); Drumond et al. 1996b: 40 (citation); Nogueira-Neto 1997: 381; Michener 1997: 56, 59 (type species, Trigonella as homonym); Nagamitsu & Inoue 1998: (foraging, morphology); Cobara & Dejan 1998: 491 (citation); Nagamitsu et al. 1999: (floral records (not by species), foraging); Heard 1999a: 187 (citation); Michener 2000: 801-802; Amano et al. 2000: (citation); Rasmussen 2004: 599 (citation); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76, 83 (distribution, foraging); Roubik 2006: 135 (nest); Rasmussen & Cameron 2007: (systematics); Michener 2007a: 826. Sundatrigona indet. Inoue & Sakagami 1993: 769 (citation); Inoue & Sakagami 1993: 769 (homonym (for Trigonella)); Roubik 2006: 127 (nest).

Tetragonilla Tetragonilla atripes (Smith, 1857) Trigona atripes Smith 1857: 50: Type (OUMNH (=Wilson Saunders collection)) (taxonomy); Type locality: MALAYSIA "Malacca (Mt. Ophir)" (worker); Smith 1871: 395 (distribution); MALAYSIA, Malacca; Dalla Torre 1896: 576 (citation); Bingham 1897: 560, 561 (distribution, key to species, taxonomy); MALAYSIA, Malacca; MYANMAR, Mergui, S Tenasserim; Friese 1901: 271 (comparative note); Bingham 1905: 59 (citation, distribution); Friese 1909: 277 (comparative note); Friese 1918: 520 (taxonomy); INDONESIA, Sumatra, Tandjong Slamat; Cockerell 1919c: 78, 79 (comparative note); Cockerell 1923:


RASMUSSEN


240 (key to species); Cockerell 1927: 541 (distribution); MALAYSIA, Selangor, Pahang, near Genting, Sempat, Hill stream; Schwarz 1939a: 85, 90, 93, 94, 95, 121-123*, 124 (distribution, key to species, taxonomy); Floral record: Citrus limon; Schwarz 1948: 117, 290; Moure 1961: 210, 211 (morphology, systematic position); Yoshikawa et al. 1969: 173 (distribution); MALAYSIA, Kuala Lumpur; George Town, Penang; Touaran, Sabah; Sakagami 1975: 59* (distribution); MALAYSIA; THAILAND; Sakagami 1978: 230 (comparative note); Wille 1979b: 273 (citation, illustration); Wille 1979a: (morphology (pollen press)); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Sakagami & Khoo 1987: 207, 208, 209 (comparative note); Uji 1987: (pollination); INDONESIA, E Kalimantan; Floral record: Nephelium lappaceum; Camargo 1988: 371 (distribution); MALAYSIA, Malacca; Roubik 1989: 133, 417; Michener 1990: 105, 126 (illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); MALAYSIA; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Yong 1991: (biochemistry); Michener & Roubik 1993: 260 (comparative note, morphology); Yong 1993: 54 (biochemistry); Cunha 1994: (systematics); Bruijn & Sommeijer 1997: 44 (citation); Michener 1997: 57 (type species); Liow 1999: 127; Michener 2000: 801; Chinajariyawong & Saiboon 2000: (distribution, nest); THAILAND, Songkhla; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics); Michener 2007a: 826. Tetragonilla collina (Smith, 1857) Trigona cambodiensis Cockerell 1926a: 224: Holotype (USNM 29470, worker): examined (nest, taxonomy); Type locality: CAMBODIA "Angkov (=Angkor) Wat, Cambodia, Jan. 11, 1926 (H.M. Smith)" (4 workers); Cockerell 1929b: 140 (distribution, nest); CAMBODIA, Angkor; Kum Puang Creek; Nan; Cockerell 1929c: 591 (distribution); CAMBODIA, Ban Maa Hia; Pah Meeung Mt.; Schwarz 1937: 297, 300; Schwarz 1939a: 124 (synonymy); Schwarz 1948: 117; Moure 1961: 211 (morphology, possible synonymy); Sakagami 1975: 59 (synonymy); Sakagami & Khoo 1987: 207 (comparative note). Trigona collina Smith 1857: 51-52: Type (OUMNH (=Wilson Saunders collection)) (taxonomy); Type locality: MALAYSIA "Malacca (Mount Ophir)" (worker); Smith 1871: 396 (distribution); MALAYSIA, Malacca; 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); Cockerell 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 (distribution); CAMBODIA, Stung Treng; Moure 1961: 211-212 (morphology, systematic position); Sakagami & Yoshikawa 1961: 437-439 (distribution, nest (only fig. 28)); CAMBODIA, Angkor Wat; Poipet; THAILAND, Chieng Mei; Doi Suthep; Li; Mae Hoi; Sara Buri; Yoshikawa et al. 1969: 168, 169, 173, 174 (distribution, 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.



29


1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Sakagami & Khoo 1987: 207, 209, 210 (comparative note); Sakagami & Inoue 1987: 626 (comparative note); Roubik 1989: 133, 196, 243, 387; Sakagami et al. 1989: 269 (comparative note); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra; MALAYSIA; MYANMAR; THAILAND; Sakagami & Inoue 1990: 212; Salmah et al. 1990: (distribution, nest (subterranean among roots)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest (cavities among roots), population); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Yong 1991: (biochemistry); Michener & Roubik 1993: 260 (comparative note, morphology); Roubik 1996: 61, 62 (distribution, nest); BRUNEI, Kuala Belalong; Nogueira-Neto 1997: 383; Dollin et al. 1997: 877 (comparative note); Nagamitsu & Inoue 1997: 433, 434, 436 (pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Eltz et al. 2001a: (pollen foraging); Eltz et al. 2001b: (foraging); MALAYSIA, Borneo, Sabah; Floral record: Caesalpinia; Citrus; Clausena; Croton; Durio; Jasminium; Manihot esculenta; Mimosa pudica; Neonauclea; Passiflora; Sterculia; Symplocos; Terminalia; Asteraceae; Leguminosae; Green et al. 2001: 91 (microsatellite); Eltz et al. 2002b: (foraging (Rhizopus: Mucorales: Zygomycota)); Eltz et al. 2002a: (distribution, foraging, nest, population, predator (Helarctos malayanus: Ursidae: Carnivora)); MALAYSIA, Danum Valley; Sepilok; Deramakot; Nagamitsu & Inoue 2002: 305, 307, 308, 310, 312 (foraging); Tangmitcharoen et al. 2002: 213; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Franck et al. 2004: (genetics, microsatellite); Cameron et al. 2004: (genetics, reproduction); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Wattanachaiyingcharoen et al. 2004: (diversity, nest); Velthuis et al. 2005: 177 (citation); Nagamitsu & Inoue 2005: 76, 80, 83, 84, 86, 88, plate 9 (distribution, foraging); Klakasikorn et al. 2005: 2, 3, 6 (floral record); Floral record: Diospyros rhodocalyx; Dipterocarpus alatus; Ficus religiosa; Irvingia malayana; Jongjitvimol et al. 2005: (nest); THAILAND, Phitsanulok; Roubik 2006: 131, 137 (nest); Tangmitcharoen et al. 2006: (pollination); Floral record: Tectona grandis; Jongjitvimol et al. 2006: (foraging); Jongjitvimol & Wattanachaiyingcharoen 2006: (foraging); THAILAND, Phitsanulok; Floral record: Agave angustifolia; Alangium salviifolium; Butea monosperma; Caryota bacsonensis; Cassia bakeriana; Coccinia grandis; Costus speciosus; Croton roxburghii; Dalbergia lanceolaria; Delonix regia; Erythrina stricta; Fernandoa adenophylla; Ipomoea aquatica; Ixora grandifolia; Lagerstroemia calyculata; Lagerstroemia macrocarpa; Lagerstroemia tomentosa; Merremia vitifolia; Millettia brandisiana; Mimosa pigra; Mimosa pudica; Paederia linearis; Ruellia tuberose; Senna siamea; Tectona grandis; Thunbergia laurifolia; Torenia fournieri; Tridax procumbens; Zingiber officinale; Rasmussen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Archidendron jiringa; Jongjitvimol & Wattanachaiyingcharoen 2007: (nest); Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (predator (Argiope: Araneidae: Araneae; Cypsiurus balasiensis: Apodiformes: Apodidae; Dicrurus macrocercus; Dicrurus paradiseus: Passeriformes: Dicruridae; Merops leschenaulti: Meropidae: Coraciiformes; Nephila maculata: Nephilidae: Araneae; Oecophylla smaragdina: Formicidae: Hymenoptera; Pahabengkakia piliceps: Reduviidae: Hemiptera; Solenopsis geminata: Formicidae: Hymenoptera)). Tetragonilla fuscibasis (Cockerell, 1920) Trigona fuscibasis Cockerell 1920a: 115-116: Holotype (BMNH 17b.1082) (taxonomy); Type locality: MALAYSIA "Sandakan, Borneo (Baker, 9964)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Cockerell 1929b: 140 (comparative note); Schwarz 1937: 288, 298, 301, 327 (key to species); Schwarz 1939a: 85, 94, 125*, 126 (distribution, key to species, taxonomy); Schwarz


RASMUSSEN


1948: 117; Moure 1961: 212 (systematic position); Yoshikawa et al. 1969: 173 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang; Sakagami 1975: 59, 60 (comparative note); Sakagami et al. 1985: (distribution, key to species); Sakagami et al. 1990: (distribution, key to species, morphology); INDONESIA, Sumatra; MALAYSIA; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Johnson & Appanah 1990: 586 (predator (Oecophylla: Formicidae: Hymenoptera)); Yong 1991: (biochemistry); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics). Tetragonilla rufibasalis (Cockerell, 1918) Trigona rufibasalis Cockerell 1918: 387: Holotype (BMNH 17b.1111) (key to species, comparative notes, taxonomy); Type locality: MALAYSIA "Sandakan, Borneo (Baker 9225)" (worker); Cockerell 1920a: 116 (variation); MALAYSIA, Borneo, Sandakan; Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 283, 290, 311*, 327 (key to species, taxonomy); Schwarz 1939a: 85, 93, 125-126* (distribution, key to species, taxonomy); Moure 1961: 212 (morphology, systematic position); Sakagami 1975: 59, 60 (comparative note); Sakagami et al. 1985: (distribution, key to species); Camargo 1988: 371 (distribution); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1990: 131, 136 (distribution); Yong 1991: (biochemistry); Michener & Roubik 1993: 260 (comparative note, morphology (as rufibasis!)); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433, 436 (pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Nagamitsu & Inoue 2002: 305, 307, 308 (foraging); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 77, 80, 83, 86 (distribution, foraging); Rasmussen & Cameron 2007: (systematics). Tetragonilla indet. Camargo 1988: 371, 373 (distribution); INDIA; MYANMAR.

Tetragonula Tetragonula bengalensis (Cameron, 1897) Trigona bengalensis Cameron 1897: 143-144: Probably OUMNH (nest (in old tree stumps), taxonomy); Type locality: INDIA "Barrackpore (Rothney)" (worker); Schwarz 1937: 308; Schwarz 1939a: 111 (synonymy); Moure 1961: 208 (possible synonymy); Sakagami 1978: 216, 247 (synonymy); Sakagami et al. 1983a: 106 (comparative note); Sakagami & Inoue 1985: 177 (comparative note); Sakagami et al. 1985: (distribution); Sakagami & Inoue 1987: 626 (comparative note); Sakagami et al. 1990: 130, 131 (comparative note, distribution); Wu 2000: 392 (synonymy); Nagamitsu & Inoue 2005: 77 (distribution, foraging). Tetragonula biroi (Friese, 1898) Trigona birói Friese 1898: 428, 429: Syntypes (ZMHB, 2 workers): examined, "N. Guinea / Biró 96", "Friedrich- / Wilh.-hafen", "Trigona / biroi / det. Friese 1897" (1 worker); "Philippinen / Schadenberg", "Trigona / biroi / det. Friese 1897 / n.sp.", "Coll. / Friese" (1 worker) (key to species, taxonomy); Type locality: PHILIPPINES "Philippinen (Schadenburg)" (few workers); "N-Guinea (Berlinhafen und



31


Friedrich Wilhelmshafen, 1896 Biró" (3 workers); Cockerell 1905: 220 (synonymy); Ashmead 1905: 957 (citation, distribution); PHILIPPINES; Brown 1906: 686 (distribution); PHILIPPINES; Friese 1909: 272, 273, 274 (key to species, taxonomy); PAPUA NEW GUINEA, Erima; Simbang; Strand 1911: 87 (synonymy); Cockerell 1912: 225 (comparative notes); Cockerell 1919c: 77, 79 (comparative notes, floral record, key to species); PHILIPPINES, Bataan, Lamao; Luzon, Los Banos; Floral record: Nipa fruticans; Cockerell 1925c: 53, 55 (distribution); PHILIPPINES, Bangui, Ilocos norte; Limay; Cockerell & LeVeque 1925: 175 (distribution, floral record); PHILIPPINES, Loquilocon; Samar; Wright (former Paranas); Floral record: Citrus; Melicope triphylla; Mimosa pudica; Hedicke 1926: 423 (distribution); PHILIPPINES, Luzon; Mindanao; Siargao; Williams 1928: 91-92 (nest); Cockerell 1929d: 301 (comparative note (as birsi!)); Cockerell 1930a: 157 (distribution, floral record, uncertain identity); CAMBODIA, Tung Nui (misidentification); Satul (misidentification); Floral record: Melastoma malabaricum; Schwarz 1939a: 85, 86, 90, 101, 119-120* (distribution, key to species, taxonomy); Floral record: Citrus; Melicope triphylla; Mimosa pudica; Nipa fruticans; Moure 1961: 209 (systematic position); Michener 1961: 28 (comparative note); Michener 1965: 231 (citation); Baltazar 1966: 406-407 (distribution); PHILIPPINES, Luzon, Bataan; Ilocos Norte; Laguna; Masbate, Samar; Nueva Vizcaya; Siargao; Mindanao, Lanao; Starr & Sakagami 1987: 97 (citation); Camargo 1988: 371 (distribution); PHILIPPINES, Luzon, Mt. Makiling; Cervancia & Barile 1993: (foraging, nest); Floral record: Asystasia gangetica; Callistemon citrinus; Dillenia suffruticosa; Jatropha podagrica; Mussaenda; Nerine; Nicotiana tabacum; Plumbago auriculata; Roubik 1995: 151 (antropogenic nest); Barile & Cervancia 1995: (ecology, flight, floral record, foraging, nest); Floral record: Antigonon leptopus; Asystasia gangetica; Callistemon citrinus; Coccinia; Cocos nucifera; Dillenia suffruticosa; Euphorbia pulcherrima; Galphimia glauca; Hamelia patens; Impatiens balsamina; Ixora; Jatropha hastata; Jatropha podagrica; Leucaena leucocephala; Mussaenda; Nerine; Nicotiana tabacum; Plumbago auriculata; Dollin et al. 1997: 877 (comparative note, distribution); PHILIPPINES; New Guinea; Oshy & Arikumar 2000: 1651 (pollination); Floral record: Bambusa vulgaris; Koshy et al. 2001: (see iridipennis); Cortopassi-Laurino et al. 2006: 303 (citation (as biro!)); Slaa et al. 2006: 303 (citation (as biro!)); Temaru et al. 2007: 282, 284 (honey properties). Trigona bismarkiana 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). Tetragonula carbonaria (Smith, 1854) Trigona angophoræ Cockerell 1912: 225: Holotype (BMNH 17b.1139) (comparative notes, floral record, taxonomy); Type locality: AUSTRALIA "Sydney, New South Wales, Dec. 1, 1910 (Froggatt, 118)" (worker); Floral record: Angophora; Cockerell 1929d: 301 (comparative note, new status (as subspecies of carbonaria)); Cockerell 1930b: 150 (distribution, floral record, taxonomy); AUSTRALIA, Sydney; Floral record: Angophora; Rayment 1932f: 106 (taxonomy); Rayment 1935: 735, 740 (key to species, taxonomy); Rayment 1939: (distribution); Schwarz 1948: 91; Rayment 1953: (distribution); Moure 1961: 209 (morphology, systematic position); Michener 1965: 231 (citation); Camargo 1988: 372 (taxonomy); Dollin et al. 1997: (revision, synonymy, taxonomy). 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 (citation); Friese 1898: 428, 429, 430 (comparative note, key to species); Friese 1905: 118 (citation); Friese


RASMUSSEN


1909: 272, 273-274 (key to species, taxonomy); AUSTRALIA, Mackay; Sydney; INDONESIA, KeyInseln (misidentification); MICRONESIA, Karolinen (misidentification); PAPUA NEW GUINEA, Sattelberg (misidentification); Simbang (misidentification); Cockerell 1910: 247 (comparative notes); Lea 1910: 119, 207 (inquiline (Brachypeplus auritus; Brachypeplus basalis: Nitidulidae: Coleoptera)); Strand 1911: 87 (distribution); Cockerell 1911: 176 (comparative notes, taxonomy); Friese 1912: 94 (distribution); Strand 1912: (distribution); Cockerell 1912: 225 (comparative notes); Lea 1912: 38-39 (inquiline (Homalota trigonae: Staphylinidae: Coleoptera)); Cockerell 1914: 412-413; Floral record: Helianthus annuus; Friese 1917: 9 (distribution); Cockerell 1922a: 279 (comparative note); Salt 1929: 452, 453 (citation); Cockerell 1929e: 243 (distribution); AUSTRALIA, Ourimbah, New South Wales; Cockerell 1929d: 301 (common name (karbi), distribution, nest); AUSTRALIA, Moreton; Cockerell 1929a: 8 (comparative note (identification corrected in footnote)); Cockerell 1930b: 149 (common name (karbi), floral record, taxonomy); Floral record: Helianthus annuus; Cockerell 1930c: 149-150 (distribution, variation); Schwarz 1932: 235-236, 245; Rayment 1932a: 185, 186 (common name (koobee; karbi; baldies), morphology); Rayment 1932d: (honey, pollen, wax); Rayment 1932b: (nest); Rayment 1932c: (behavioral comparison); Rayment 1932e: (inquiline (Brachypeplus planus; Brachypeplus meyricki (or Epuraea): Nitidulidae: Coleoptera; Tribolium myrmecophilum: Tenebrionidae: Coleoptera)); Rayment 1932f: 106 (taxonomy); Cockerell 1934: 62, fig. 8 (illustration (mandible)); Rayment 1935: 735, 736, 737, 738, 739 (comparative note, key to species, taxonomy); Cannon 1937; Schwarz 1939a: 103 (synonymy); Rayment 1939: (distribution); Schwarz 1948: 22, 24, 29, 32, 37, 41, 60, 62, 67, 68, 77, 80, 84, 86, 91, 93, 94, 97, 107, 110, 117, 133, 166; Rayment 1953: (distribution); Moure et al. 1958: 484 (citation); Moure 1961: 209 (morphology, systematic position); Michener 1961: 3, 9, 10, 11, 15-23, 24, 25, 27, 39, 40, 42, 44 (aggressiveness, nest); AUSTRALIA, Beenleigh; Beerwah; Binna Burra (near Lamington National Park); Brisbane; Bundaberg; Caboolture; Caloundra; Capalaba; Mount Edwards; Gladstone; Ipswich; Jimboomba; Mackay; Maroochydore; Moggill (near Brisbane); Noosa; Petrie; Rockhampton; Sarina; Tamborine; Tibrogargen (near Beerwah); Yaamba; Yarriman; Wille 1961: 117; Rayment 1961: (nest); Wakefield 1962: (swarm (probably not stingless bees)); Michener 1964: 236 (nest); Michener 1965: 231 (citation); Pooley & Michener 1969: 429 (comparative note); Nogueira-Neto 1970: 218, 266, 314; Wille & Michener 1973: 13, 24, 47, 58, 70, 80, 81, 89, 92, 178 (nest); Michener 1974: 62, 341, 343 (colony, nest); McKenzie 1975: (common name (kabbai), honey); Kistner 1982: 122 (citation, inquiline (Brachypeplus auritus: Nitidulidae: Coleoptera)); Wagner & Dollin 1982a: (distribution); Evans & Matthews 1973: (predator (Bembix musca: Crabronidae: Hymenoptera)); Evans et al. 1982: 78 (predator (Bembix musca: Crabronidae: Hymenoptera)); Sakagami et al. 1983b: 20 (comparative note); Sakagami et al. 1983a: 106 (comparative note); Sakagami et al. 1983d: 454 (comparative note); Wagner & Briscoe 1983: (genetics); Hawkeswood 1983: (pollination); Floral record: Cupaniopsis anacardioides; Graur 1985: 194 (comparative note); Yong 1986: 628 (comparative note); Goebel 1986: (bee-keeping); Starr & Sakagami 1987: 97, 106 (citation); Yong et al. 1987: 467 (biochemistry); Fain & Heard 1987: 197, 199, 202 (inquiline (Cerophagus trigona: Acaridae: Acari)); Milborrow et al. 1987: 15, 22, 23, 25 (wax); Heard 1987: (nest); Camargo 1988: 372 (distribution, taxonomy); AUSTRALIA, Cape York, Gosford; Maitland, New South Wales; Heard 1988: (bee-keeping); Adams & Lawson 1988: (pollination); Michener 1990: 95, 101, 119, 120, 127, 128 (illustration, morphology, systematic position); Sakagami & Inoue 1990: 211, 212, 215 (comparative note); Wittmann et al. 1991: 802, 806 (citation); Harriss & Beatie 1991: (pollen transportation); Ornduff 1991: 207 (pollination); Floral record: Cycas media; Adams et al. 1992: (orchid pollination); Michener & Roubik 1993: 259, 260 (comparative note, morphology); Heard & Hendrikz 1993: (flight activity); Cardale 1993: 321-322 (distribution, floral record, list); AUSTRALIA, NE coastal; N coastal; New South Wales; Queensland; SE coastal; Western Australia; Murray-Darling basin; Floral record: Ageratum; Alocasia; Angophora; Banksia; Callistemon; Claoxylon; Cupaniopsis; Daviesia; Dendrobium; Eucalyptus; Hibbertia; Jacksonia; Leptospermum; Melaleuca;



33


Melastoma; Nymphaea; Oxylobium; Persoonia; Podolepis; Pultenaea; Rubus; Telopea; Xanthorrhoea; Williams & Adam 1993: 480 (floral record); Gross 1993: 471, 472, 473 (pollination); Floral record: Melastoma affine; Adams & Lawson 1993: (pollination); Floral record: Dendrobium teretifolium; Heard 1994: (pollination); Floral record: Macadamia integrifolia; Heard & Exley 1994: (pollination); Floral record: Macadamia integrifolia; Forster et al. 1994: 219 (pollination); Floral record: Cycas; Bartareau 1994: (pollination); Floral record: Dendrobium toressae; Yamane et al. 1995: (oviposition); Wallace & Trueman 1995: (seed dispersal (mellitochory)); Floral record: Eucalyptus torelliana; Disney & Bartareau 1995: (predator (Dorniphora trigonae: Phoridae: Diptera) (of brood?)); Bartareau 1995: (pollination); Floral record: Cymbidium madidum; Drumond et al. 1996a: 396, 399 (behavioral comparison); Kerr et al. 1996: 54 (citation); Bartareau 1996: (foraging); Yamane 1996: 14 (nest); Heard 1996: (beekeeping, nest); Dollin 1996a: (characteristics, nest); Dollin 1996b: (nest); Dollin 1996c; Nogueira-Neto 1997: 85, 204, 205, 334; Dollin et al. 1997: (nest, revision, taxonomy); AUSTRALIA, New South Wales; Queensland; Williams & Adam 1997: 75, 86, 88, 90-92 (competition, floral record); Floral record: Abrophyllum ornans; Acradenia euodiiformis; Alphitonia excelsa; Euroschinus falcata; Rhodomyrtus psidioides; Tristaniopsis laurina; Williams & Adam 1997: 75 (floral record); Floral record: Cuttsia viburnea; Dollin & Heard 1997: (domestication); Dollin & Heard 1997: (bee-keeping, predators (Phoriae; Syrphidae)); Dollin 1997: (characteristics); Heard & Dollin 1998: (citation); Heard & Dollin 1998: 6, 13 (foraging activity, pollination); Drumond et al. 1999: 235, 241 (comparative note); Heard 1999a: 186, 187, 195 (citation); Crane 1999: 89, 90; Dollin & Heard 1999: (bee-keeping); Michener 2000: 94, 783f, 800; Hilário et al. 2000: 304 (citation); Nieh et al. 2000: (communication, nest); Heard & Dollin 2000: (bee-keeping); Amano et al. 2000: (citation); Green et al. 2001: (microsatellite); Hilário et al. 2001: 195 (citation); Serrão 2001: (morphology (proventricular structure)); White et al. 2001: (flower constancy); Dollin et al. 2001: (bee-keeping); Green & Oldroyd 2002: (maternity of male); Wallace et al. 2002: (pollination); Floral record: Persoonia virgata; Pick & Blochtein 2002: 827, 831, 833, 834 (citation); Dollin 2002a: (bee-keeping); Dollin 2002b: (bee-keeping); Schmidt et al. 2003: 245 (citation); Gardener et al. 2003: 120 (citation); Williams & Walker 2003: 143, 144, 145 (floral record); Floral record: Pollia crispata; Heard 2003: (citation); Nieh 2004: 162, 164, 168 (citation, communication, comparative table); Franck et al. 2004: (genetics, microsatellite, nest); Cameron et al. 2004: 2358 (genetics); Tóth et al. 2004: 5, 6, 7 (citation); Sung et al. 2004: 263 (citation); Thomson et al. 2004: 622, 623 (pollination); Floral record: Cucumis sativus; Amano 2004: 3 (introduction, pollination); Floral record: Lycopersicon esculentum; Borges & Blochtein 2005: 683 (citation); Kajobe & Echazarreta 2005: 274 (citation); Bruce & Herberstein 2005: 3834, 3837, 3843 (predator (Argiope: Araneidae: Araneae)); Greco et al. 2005: 72 (citation (as stingless bees)); Roubik 2006: 133, 136 (nest); Hartfelder et al. 2006: 154 (citation); Arias et al. 2006: 267 (citation); Cortopassi-Laurino et al. 2006: (bee-keeping); Slaa et al. 2006: 297, 301, 306, 308 (citation); Bacelar-Lima et al. 2006: 344 (citation); Hoese et al. 2006: 769, 774 (predator (Argiope keyserlingi: Araneidae: Araneae)); Rasmussen & Cameron 2007: (systematics); Michener 2007a: 100, 807f, 824; Jongjitvimol & Wattanachaiyingcharoen 2007: 29 (citation); Stow et al. 2007: (antimicrobials, honey); Gloag et al. 2007: (reproduction); Fidalgo & Kleinert 2007: 138 (citation); Klumpp 2007; Wallace et al. 2008: (dispersal, pollination); Floral record: Corymbia torelliana; Irish et al. 2008: (honey properties). Tetragonula clypearis (Friese, 1909) Trigona laeviceps variety clypearis Friese 1909("1908"): 356, 358, fig. 15-3: Unknown; paralectotype (ZMHB (1)) (distribution, taxonomy); Type locality: INDONESIA (IRIAN JAYA) "Manikion, 14.-28. Februar 1903"; "Wendesi 29. Juni 1903, New Guinea" (worker); Friese 1909: 275; Moure 1961: 209 (systematic position); Michener 1961: 34 (synonymy); Michener 1965: 231 (citation); Wille &


RASMUSSEN


Michener 1973: 13, 24, 47, 58, 70, 94 (nest); Michener 1974: 331, 343 (nest); Starr & Sakagami 1987: 97 (citation); Sakagami et al. 1990: 129 (comparative note); Cardale 1993: 322 (distribution); AUSTRALIA, NE coastal; Queensland; Ikudome & Kusigemati 1996: 17, 19-20 (common name (bai), distribution, nest, taxonomy); PALAU ISLANDS; Dollin 1996b: (nest); Dollin et al. 1997: (distribution, nest, revision, taxonomy); AUSTRALIA, Queensland; INDONESIA, Ambon (probably distinct species); "NEW GUINEA"; PHILIPPINES (probably distinct species); Dollin 1997: (characteristics); Heard & Dollin 2000: (bee-keeping); Green et al. 2001: 91 (microsatellite); Palmer et al. 2002: (maternity); Dollin 2002a: (bee-keeping); Franck et al. 2004: (genetics, microsatellite); Tóth et al. 2004: 6 (citation); Sung et al. 2004: 268 (citation); Rasmussen & Cameron 2007: (systematics); Klumpp 2007: 4, 5, 82; Wallace et al. 2008: (dispersal, pollination); Floral record: Corymbia torelliana. Trigona wybenica Cockerell 1929d: 300: Holotype (USNM 54960, worker): examined; paratypes (BMNH (1), QM (3), ZMHB (1)) (common name (wyben), nest, taxonomy); Type locality: AUSTRALIA "Thursday island" (worker); Cockerell 1930b: 150 (taxonomy); Rayment 1932f: 107 (taxonomy); Rayment 1935: 735, 738 (distribution, key to species); Cockerell 1939b: 133 (pest (crawling into hair)); Moure 1961: 209 (morphology, synonymy); Michener 1961: 10, 34-36, 38, 39, 40, 42 (aggressiveness, nest); Michener 1964: 236 (nest); Michener 1965: 231 (citation); Wille & Michener 1973: 13 (synonymy); Wagner & Dollin 1982b: (swarming); Wagner & Dollin 1982c; Sakagami et al. 1983a: 106 (comparative note); Dollin & Dollin 1983: (nest); Cardale 1993: 322 (synonymy); Dollin et al. 1997: (revision, synonymy, taxonomy); Dollin 1997: (characteristics); Crane 1999: 89. Tetragonula dapitanensis (Cockerell, 1925) Trigona dapitanensis Cockerell 1925a: 492: Holotype (BMNH 17b.1135). Here considered a valid and larger form 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). Tetragonula davenporti (Franck, 2004) Trigona (Heterotrigona) davenporti Franck et al. 2004: 2325, 2326, 2327, 2328, 2330-2331, figs. 5a, 6a: Holotype (QM, worker); paratypes (ANIC (2), BMNH (2), MNHN (2), QM (2), "INRA collection, Montpellier" (8)) (genetics, illustration, microsatellite, nest, taxonomy); Type locality: AUSTRALIA "AUSTRALIA, QLD, Mudgeeraba, Austinville Road, V. 1990, P. Davenport leg." (worker); Klumpp 2007: 4, 5. Tetragonula devicta (Cockerell, 1921) 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 from Mr. R.C.J. Swinhoe" (worker); Cockerell 1922b: (age of fossil); Kerr & Maule 1964: (systematics, taxonomy); 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 & Ross 2000: 6, 13-14 (synonymy).



35


Tetragonula drescheri (Schwarz, 1939) 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; Sakagami 1959: 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 species, 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: Carnivora)); BRUNEI, Kuala Belalong; Michener & Boongird 2004: 144, 145 (comparative note); Nagamitsu & Inoue 2005: 77 (distribution, foraging). Tetragonula fuscobalteata (Cameron, 1908) Trigona atomella Cockerell 1919b: 243-244: Holotype (USNM 29467, worker): examined, "Island of / Penang / 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 clinging to the leg of a large green cicada (Platylomia spiculata (=Dundubia spiculata): Cicadidae: Hemiptera)")); CAMBODIA, Pahtoop Mt.; Cockerell 1930a: 157 (distribution, ecology (attracted to sweat)); CAMBODIA, Chun Het; Trang; Yasumatsu 1935: 94 (common name (mitsubachi), distribution); PALAU ISLANDS; Esaki 1936: 44, 46-37 (common name (galadngikl; chaladngikl; mitsubachi), nest); PALAU ISLANDS; 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 by man")); PALAU ISLANDS, Melekeiok on Babelthuap; Ngeremlengui, Ogiwal; Schwarz 1948: 69, 117; Krombein 1950: 133 (synonymy); Moure 1961: 209 (morphology, systematic position); Sakagami 1978: 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 no description 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, taxonomy); Type locality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker or queen); Cockerell 1918: 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). Trigona fusco-balteata Cameron 1908: 193, 194: Lectotype (BMNH 17b.1112): Designated Moure 1961 (key to species, taxonomy); Type locality: MALAYSIA "Medang, Sarawak (Hewitt)" (worker); Schwarz 1937: 290, 310-311*, 316, 326 (key to species, taxonomy); Schwarz 1939a: 85, 86, 88, 90, 93, 96, 107110*, 111, 113, 114, 115, 132 (distribution, key to species, taxonomy); Schwarz 1939b: 151-152 (distribution, taxonomy); PALAU ISLANDS, Arukoron; Babelthaob, Marukyoku-Kaisharu; Korror; Schwarz 1948: 40, 56, 69, 83, 90, 118; Krombein 1950: 102, 133, 135 (distribution, morphology); MICRONESIA, Caroline Islands; Nogueira-Neto 1951: (comparative note); Pagden 1957: 15, 17 (flight); Sakagami


RASMUSSEN


1959: 121 (comparative note); Moure 1961: 209 (morphology, systematic position); Michener 1961: 34 (comparative note); Sakagami & Yoshikawa 1961: 435 (comparative note); Baltazar 1966: 407 (distribution); PHILIPPINES, Mindanao, Lanao; Palawan; Yoshikawa et al. 1969: 168, 169, 172, 173, 174 (distribution, nest); MALAYSIA, Tapah; Tousaran; Jander & Jander 1970: 361, 362 (orientation); Wille & Michener 1973: 14, 24, 30, 59, 94 (nest); Michener 1974: 331, 343 (nest); Sakagami 1978: 166-194, 219, 220, 223-226*, 227, 228, 231, 232, 234, 236, 237, 238, 239, 240, 247, plate 5 (taxonomy); Sakagami 1982: 389 (citation); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Wille 1983: 50 (nest); Sakagami et al. 1983b: 2, 9-10, 20 (nest); Sakagami et al. 1983a: 106 (comparative note); Sakagami & Inoue 1985: 177 (comparative note); Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Yong 1986: (allozyme); Sakagami & Khoo 1987: 213 (comparative note); Sakagami & Inoue 1987: 626 (comparative note); Starr & Sakagami 1987: (common name (kiyot), nest); Yong et al. 1987: (biochemistry); Camargo 1988: 371 (distribution, synonymy); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1989: 266 (comparative note); Sakagami 1989: 164, 166 (nest); Michener 1990: 97, 105, 110, 119, 127, 131, 119 (illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species, variation); CAMBODIA; INDONESIA, Celebes; Sumatra; MALAYSIA; THAILAND; Sakagami & Inoue 1990: (oviposition); Salmah et al. 1990: (distribution, nest (bamboo stems; interspaces; cracks; crevices)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Roubik 1990c: 663 (citation); Wittmann et al. 1991: 802 (citation); Yong 1993: 54 (biochemistry); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Oliveira et al. 1995: 20 (citation); Yamane et al. 1995: 276, 284, 286, 292, 293 (comparative note); Roubik 1995: 41, 42; Roubik 1995: 151 (antropogenic nest); Roubik 1995: 41, 42; Drumond et al. 1996a: 395 (behavioral comparison); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Ikudome & Kusigemati 1996: 20 (taxonomy); Dollin et al. 1997: (nest, revision, taxonomy (Australian form is distinct)); AUSTRALIA, Queensland; INDONESIA, Sumatra; MALAYSIA; THAILAND; Nogueira-Neto 1954: 426 (citation); Nagamitsu & Inoue 1997: 433, 436 (pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 780; Chinajariyawong & Saiboon 2000: (distribution, nest); THAILAND, Songkhla; Amano et al. 2000: (citation); Michener 2001: 232 (morphology); Green et al. 2001: 91 (microsatellite); Eltz et al. 2003: 309 (comparative note); Franck et al. 2004: 2318 (citation); Cameron et al. 2004: 2358 (nest); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Amano 2004: 3 (introduction, pollination); Wattanachaiyingcharoen et al. 2004: (diversity, nest); Nagamitsu & Inoue 2005: 77, 78, 79, 80, 81, 83 (distribution, foraging); Klakasikorn et al. 2005: 3, 5, 6 (floral record); Roubik 2006: 130 (nest); Rasmussen & Cameron 2007: (systematics); Michener 2007b: 130 (comparative note); Michener 2007a: 806. Trigona pallidistigma Cameron 1908: 193, 195: Syntypes (BMNH 17b.1133, 2? workers) (key to species, taxonomy); Type locality: MALAYSIA "Sarawak, Borneo (R. Shelford)" (worker); Schwarz 1937: 291, 316* (key to species, taxonomy); Schwarz 1939a: 85, 108 (distribution, key to species, taxonomy); Moure 1961: 209 (synonymy); Sakagami 1978: 223, 224 (synonymy); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1990: 128 (synonymy). Trigona pfeifferi Friese 1925: 41: Lectotype (AMNH, worker): here designated, "Brasil / Campinas", "Trigona / pfeifferi / Friese det. 25 / Fr.", "Typus" (orange label), "Heterotrigona / cf. fuscobalteata / (erroneous locality) / det. J.S.Ascher"; paralectotypes (DEI (3)) (taxonomy); Type locality: UNCERTAIN "Campinas (São Paulo), S.-Brasil" (several workers; wrong locality label); Schwarz 1937: 285; Schwarz 1948: 195, 203; Wille 1962: 179 (comparative note); Wille 1979b: 264, 265 (morphology (pollen press)); Wille



37


1979a: (morphology (pollen press)); Michener & Roubik 1993: 260 (comparative note, morphology); Camargo & Pedro 2003a: (synonymy); Camargo & Pedro 2007: 524 (synonymy). Trigona pygmaea Friese 1933b: 147: Lectotype (ZMHB, worker): here designated "O-Sumatra / Mandau / 7.1933", "Trigona / pygmaea / Fr. / 1933 Friese det. / n."; paralectotype (DEI (1), ZMHB (1)) (common name (kloeloet itam ketjid)); Type locality: INDONESIA "Beringin, in der Wäldern oberhalb Mandau (Sumatra), v. Bengkalis. Im Juli 1933" (several workers); Schwarz 1937: 310; Schwarz 1939a: 107 (synonymy); Krombein 1950: 133 (synonymy); Moure 1961: 209 (possible synonymy); Sakagami 1978: 223 (synonymy); Sakagami et al. 1990: 128 (synonymy). Tetragonula geissleri (Cockerell, 1918) Trigona confusella Cockerell 1919b: 242: Holotype (USNM 40248) (taxonomy, previously identified as T. geissleri (in Cockerell 1918)); Type locality: SINGAPORE "Singapore (Baker)" (worker); Schwarz 1937: 282, 290, 311-313* (key to species, taxonomy); Schwarz 1939a: 101, 117 (synonymy); Moure 1961: 209 (synonymy); Sakagami 1978: 195, 196 (synonymy). Trigona geissleri Cockerell 1918: 385-386, 387: Unknown: Cockerell used a Friese manuscript name for his description (key to species, distribution, manuscript name of Friese, taxonomy, variation); Type locality: MALAYSIA "Sintang, North Borneo" (male); "Singapore by Baker" (couple of workers); Cockerell 1919b: 242 (taxonomy, variation); MALAYSIA, Sandakan; Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 311, 312, 313, 327; Schwarz 1939a: 85, 90, 95, 109, 114, 115, 117-119*, 120, 136 (distribution, key to species, taxonomy (probably several distinct species were included here)); Schwarz 1948: 83, 118, 233; Moure 1961: 209 (morphology, systematic position); Sakagami & Yoshikawa 1961: 435, 436 (comparative note, distribution, taxonomy); LAOS, Thakhek; Yoshikawa et al. 1969: 172, 173 (distribution); BRUNEI; MALAYSIA, Cameron Highland, Pahang; Fraser's Hill, Selagor-Pahang; Touaran; Kampong Astana, near Kuching; Sakagami 1978: 166-194, 195198*, 199, 200, 208, 209, 210, 226, 227, 228, 231, 234, 235, 236, 237, 238, 239 (taxonomy); MALAYSIA; SINGAPORE; THAILAND; Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1983b: 11, 20 (nest); Sakagami & Inoue 1985: 177 (comparative note); Sakagami et al. 1985: (distribution, key to species); Appanah et al. 1986: (foraging, nest); Floral record: Calamus; Calophyllum; Elaeis; Sakagami & Khoo 1987: 208, 209, 210, 211, 212 (morphology); Sakagami & Inoue 1987: 625, 626 (morphology); Starr & Sakagami 1987: 97 (citation); Camargo 1988: 371 (distribution); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1990: 128, 131 (distribution, misidentification); Oldroyd et al. 1992: 5 (pollination); THAILAND, Chanthaburi; Floral record: Archontophoenix alexandrae; Roubik 1995: 151 (antropogenic nest); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Bruijn & Sommeijer 1997: 44 (citation (as geisslerrii!)); Liow 1999: 106, 127 (distribution (as geisslerei! in part)); Nagamitsu et al. 1999: (floral records (not by species), foraging); Liow et al. 2001: 184; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Franck et al. 2004: 2318 (citation); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics). Tetragonula gressitti (Sakagami, 1978) Trigona (Tetragonula) gressitti Sakagami 1978: 166-194, 206, 214-216*, 226, 227, 228, 234, 235, 236, 237, 238, 239: Holotype (BPBM; not located, nor in SEHU); paratypes (SEHU) (taxonomy); Type locality: VIETNAM "Fyan-b, Viet Nam" (49 workers, holotype, paratypes); "Fyan-a" (1 worker, 1 male); "Dalat"


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(5 workers); Balao (1 male); Sakagami & Inoue 1985: 177 (comparative note); Sakagami et al. 1985: (distribution); Sakagami & Inoue 1987: 626 (comparative note); Sakagami & Inoue 1989: 615 (comparative note); Sakagami et al. 1990: 131 (distribution); Sakagami & Inoue 1990: 202 (comparative note); Tadauchi et al. 1998: 243 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution, foraging). Tetragonula hirashimai (Sakagami, 1978) Trigona (Tetragonula) hirashimai Sakagami 1978: 166-194, 195, 196, 197, 198-200*, 202, 212, 215, 226, 227, 229, 234, 236, 237, 238, 239, 240: Holotype (SEHU) (taxonomy); Type locality: THAILAND "Chieng Dao-b, N. Thailand" (1 worker, holotype); "Doi-Suthep" (8 workers, 62 males)"; "Chieng Mai (1 worker); "Mae Klang" (2 workers); "Mae Fack" (1 worker); "Lamphun" (1 worker); "Uthaithani" (1 worker); Sakagami & Inoue 1985: 177, 182 (comparative note); Sakagami et al. 1985: (distribution); Sakagami & Inoue 1987: 625, 626 (morphology); Sakagami et al. 1990: 131 (distribution); Dollin et al. 1997: 873 (comparative note); Tadauchi et al. 1998: 243 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics). Tetragonula hockingsi (Cockerell, 1929) Trigona carbonaria hockingsi n. subsp. Cockerell 1929a: 8: Holotype (QM, worker); paratypes (BMNH (2), QM (1)) (comparative note, distribution, nest); Type locality: AUSTRALIA "Cape York Peninsula, Harold Hockings" (five workers); "Port Darwin"; Cockerell 1929d: 301 (comparative note, distribution, nest, redescription); AUSTRALIA, Cape York; Cockerell 1930b: 150 (distribution, nest, taxonomy); AUSTRALIA, Cape York, Port Darwin; Cockerell 1930c: 150 (comparative note); Rayment 1932f: 106 (taxonomy); Rayment 1935: 735, 737, 739-740 (key to species, taxonomy); MacPherson & Rupp 1936: (pollination); Floral record: Cymbidium iridifolium; Schwarz 1948: 91; Moure 1961: 209 (comparative note, morphology, systematic position); Michener 1961: 10, 18, 23-27, 29, 40, 42, 44 (aggressiveness, nest); Wille 1961: 117, 125, 129; Michener 1965: 231 (citation); Kerr et al. 1967: 283 (citation); Darchen 1969: 462 (illustration, physiology); Wille & Michener 1973: 14, 24, 38, 48, 59, 71, 94, 100 (nest); Michener 1974: 334, 343, 344 (nest); Wagner & Dollin 1982b: (swarming); Wagner & Dollin 1982c; Sakagami et al. 1983b: 20 (comparative note); Sakagami et al. 1983a: 106 (comparative note); Sakagami et al. 1983d: 454 (comparative note); Dollin & Dollin 1983: (nest); Starr & Sakagami 1987: 97 (citation); Michener 1990: 120 (systematic position); Sakagami & Inoue 1990: 211, 212, 215 (comparative note); Wittmann et al. 1991: 802 (citation); Cardale 1993: 323 (distribution, floral record, list); AUSTRALIA, NE coastal; N coastal; N Gulf; Northern Territory; Queensland; Floral record: Dendrobium; Bartareau 1994: (pollination); Floral record: Dendrobium toressae; Yamane et al. 1995: 294 (synonymy); Welch 1995: (rock art); Bartareau 1995: (pollination); Floral record: Cymbidium madidum; Dollin 1996a: (characteristics, nest); Dollin 1996b: (nest); Dollin 1996c; Dollin et al. 1997: (nest, revision, taxonomy); AUSTRALIA, Northern Territory; Queensland; Dollin & Heard 1997: (domestication); Dollin & Heard 1997: (bee-keeping, predators (Phoriae; Syrphidae)); Dollin 1997: (characteristics); Heard & Dollin 1998: 8, 13 (pollination); Floral record: Litchi chinensis; Crane 1999: 89; Dollin & Heard 1999: (bee-keeping); Michener 2000: 800; Heard & Dollin 2000: (bee-keeping); Green et al. 2001: 91 (microsatellite); Dollin et al. 2001: (bee-keeping); Palmer et al. 2002: (maternity); Dollin 2002a: (bee-keeping); Dollin 2002b: (bee-keeping); Gardener et al. 2003: (foraging, nectar choice); Franck et al. 2004: (genetics, microsatellite, nest); Tóth et al. 2004: 6 (citation); Sung et al. 2004: 268 (citation); Arias et al. 2006: 264 (citation); Cortopassi-Laurino et al. 2006: (bee-keeping); Rasmussen



39


& Cameron 2007: (systematics); Michener 2007a: 824-825; Klumpp 2007: 4, 5, 6, 8, 11, 15, 46, 77, 85; Wallace et al. 2008: (dispersal, pollination); Floral record: Corymbia torelliana. Tetragonula iridipennis (Smith, 1854) Trigona iridipennis Smith 1854: 413-414: Holotype (BMNH 17b.1114, worker): examined, "Type" (orange border), "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 species 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); Type locality: SRI LANKA "Ceylon" (worker); Tennent 1859: 282 (list); Tennent 1861: 454 (list); Castets 1893: 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 (synonymy (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); Cockerell 1929c: 590, 592 (distribution, floral record, taxonomy, variation); CAMBODIA, Angkor (misidentification); 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 (distribution, floral record); MYANMAR, Rangoon; Floral record: Helianthus annuus; Friese 1933a: 46 (see laeviceps); George 1933: (physiology (as irridipennis!)); George 1934: (brood, distribution, morphology, nest (crevice of wall), physiology (as irridipennis!)); INDIA, Coimbatore; Schwarz 1937: 283, 290, 308310*, 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 the specimens examined by Schwarz correspond to laeviceps, see laeviceps)); Schwarz 1939b: 151 (comparative 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!)); Krombein 1950: 133, 135 (comparative note); Krombein 1951: 278, 295 (morphology); Lindauer 1956: 546554, 555 (communication, nest); Pagden 1957: 16 (flight); Lindauer & Kerr 1958: 407, 410 (communication); Sakagami 1959: 120, 121 (comparative note); Portugal-Araújo & Kerr 1959: 226 (comparative note); Sakagami 1960: 146 (see laeviceps); Lindauer & Kerr 1960: 31, 32, 34 (communication); SRI LANKA; 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); Sakagami & Yoshikawa 1961: 434, 435, 436 (comparative note); Darchen & Louis 1961: 18, 21, 25-27 (biogeography, communication, nest); Kerr et al. 1963: 81 (citation); Kerr & Maule 1964: (comparative note); Schmid 1964: 588 (citation); Michener 1965: 231 (citation); Kerr & Esch 1965: (comparative note); 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); Sakagami 1971: 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: (systematics); 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,


RASMUSSEN


216-219*, 220, 221, 222, 225, 226, 227, 228, 230, 234, 235, 236, 237, 238, 239, 247, plate 5 (taxonomy); INDIA; SRI LANKA; Wille 1979b: 243 (comparative note); Koeniger & Vorvohl 1979: (foraging); SRI LANKA; Floral record: Boehmeria; Cassia; Cleome; Cocos; Erigeron; Homonoia; Lannea; Mimosa; Ocimum; Phyllanthus; Sapium; Smithia; Solanum; Syzygium; Taxillus; Toona; Tridax; Vitex; Divan 1981("1980"): (behavior); Sakagami & Ito 1981: 368, 373 (morphology); Wille 1983: 53 (population); Sakagami et al. 1983b: 6 (comparative note); Sakagami et al. 1983a: 101, 103, 106 (comparative note); Sakagami & Inoue 1985: 177, 182 (comparative note); Sakagami et al. 1985: (distribution); Utami 1986: (see laeviceps); Sakagami & Khoo 1987: 208, 209 (comparative note); Sakagami & Inoue 1987: 625, 626 (morphology); Uji 1987: (see laeviceps); Bichee & Sharma 1988: (pollination); Floral record: Helianthus annuus; Mohana Rao 1988("1987"): (foraging); Floral record: Trigonella corniculata; Camargo & Wittmann 1989: 223 (citation); Kumar & Kumar 1989: (foraging (as irridipensis!)); Floral record: Brassica campestris; Cajanus cajan; Mohana Rao & Suryanarayana 1989("1988"): 7 (pollination); Floral record: Citrullus lanatus; Utami & Kahono 1989: (pollination); INDIA, Bogor; Floral record: Areca vestiaria; Percy 1989: (bee-keeping); Michener 1990: 89, 119, 120, 126, 127 (systematic position); Sakagami et al. 1990: (misidentification, synonymy); Sakagami & Inoue 1990: 212 (taxonomy); Goel & Kumar 1990: 44 (pollination); INDIA, W Uttar Pradesh; Floral record: Helianthus annuus; Mohana Rao & Suryanarayana 1992("1990"): (pollination); Floral record: Guizotia abyssinica; Oldroyd et al. 1992: 6 (citation); Ramanujam et al. 1993("1992"): (pollination); Floral record: Ageratum conyzoides; Antigonon; Azadirachta indica; Bombax; Cocos nucifera; Cyanotis; Delonix regia; Eucalyptus globulus; Lawsonia alba; Leucaena leucocephala; Loranthus longiflorus; Peltophorum pterocarpum; Prosopis juliflora; Rotala densiflora; Tamarindus indica; Michener & Roubik 1993: 260 (comparative note, morphology); Cunha 1994: (systematics); Roubik 1995: 24, 41, 42 (pollination); Floral record: Allium; Roubik 1995: 24, 41, 42, 151 (antropogenic nest); Grimaldi et al. 1995: 262, 263; Ikudome & Kusigemati 1996: 20 (taxonomy); Smiley et al. 1996: 198, 200 (see laeviceps); NogueiraNeto 1997: 90; Dollin et al. 1997: (distribution, taxonomy); SRI LANKA; Michener 1997: 57 (type species); Rao et al. 1999("1997"): (pollination); Floral record: Crateva magna; Lakshmi & Suryanarayana 1999("1997"): (foraging (as inridipennis!)); Floral record: Ceiba pentandra; Joshi et al. 1998: (honey, pollen); INDIA, Pune, Maharashtra; Floral record: Ageratum conyzoides; Amaranthus; Bombax ceiba; Cocos nucifera; Moringa oleifera; Peltophorum pterocarpum; Sorghum vulgare; Zea mays; Heard 1999a: 193, 194, 197, 198 (citation); Crane 1999: 301; Michener 2000: 800-802; Jarau et al. 2000: 86 (citation); Wu 2000: 387, 392-393 (key to species, morphology); Atluri et al. 2000: (pollination, uncertain identity (unidentified)); INDIA, Visakhapatnam; Floral record: Helicteres isora; Chaudhary & Kumar 2000: 39, 40 (pollination); Floral record: Elettaria cardamomum; Mohan & Devanesan 2000: (bee-keeping, nest); Engel 2001: 176 (fossil (copal)); Koshy et al. 2001: (floral record, uncertain identity (as biroi)); INDIA, Cherthala, Alappuzha, Kerala; Thiruvananthapuram; Floral record: Bambusa vulgaris; Ochlandra travancorica; Wijesekara 2001: 153, 154 (distribution); Raju & Esradanam 2002: 1397 (floral record); Floral record: Jatropha curcas; Kuberappa et al. 2002: (pollination); Floral record: Psidium guajava; Hohmann 2003: (citation); Devanesan et al. 2003: (predator (Amblyseius: Phytoseiidae: Acarina (of inquilines?); Hermetia illucens: Stratiomyidae: Diptera (of brood); Solenopsis geminata: Formicidae: Hymenoptera)); INDIA, Vellayani, Kerala; Raju & Rao 2003: 3 (population); Biesmeijer & Slaa 2004: 148 (citation, foraging); Michener & Boongird 2004: 143 (comparative note); Sharma 2004: 7 (pollination); Floral record: Allium; Nagamitsu & Inoue 2005: 77 (distribution, foraging); Chinh et al. 2005: 367 (citation); Roubik 2005: 96 (distribution, food niche); Klakasikorn et al. 2005: 6 (citation); Eswarappa et al. 2005b: (pollination); INDIA, Bangalore; Floral record: Sechium edule; Eswarappa et al. 2005a: (pollination); INDIA, Bangalore; Floral record: Sechium edule; Eswarappa et al. 2005c: (pollination); INDIA, Bangalore; Floral record: Sechium edule; Raju et al. 2005: 1379 (pollination); Floral record: Wrightia tinctoria; Raju 2005: 106 (floral record); Floral record: Woodfordia floribunda; Raju et



41


al. 2006: 940 (floral record); Floral record: Acacia caesia; Raju & Rao 2006: 963, 965 (pollination); Floral record: Pongamia pinnata; Inson & Malaipan 2006: (see laeviceps); Rasmussen & Cameron 2007: (systematics); Michener 2007a: 824, 826. Tetragonula laeviceps (Smith, 1857) Trigona læviceps Smith 1857: 51: Holotype (OUMNH). Moure (1961) indicated that the BMNH type specimen (17b.1184) came from Mt. Ophir, which was not the type locality and the specimen cannot be considered 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 be resolved in a separate paper with C.D. Michener. Citations below for laeviceps are in a broad sense, as species limits are uncertain and may include both valdezi and testaceitarsis. These taxa have previously been proposed as junior synonymons of laeviceps, but are slightly larger (taxonomy); Type locality: SINGAPORE "Singapore" (worker); Smith 1858: (propolis, wax); Smith 1859: 135 (distribution); INDONESIA, Aru; INDIA (misidentification); SINGAPORE; Smith 1865: 93 (distribution); "NEW GUINEA"; Parish 1866: (common name (pwai-ngyet), illustration, nest, propolis); Smith 1871: 395 (distribution); 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, Tenasserim; 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 (comparative notes, 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 species, nest (in structural wood)); INDONESIA, Ost-Java; Sumatra; Schulz 1909: (nest (as canifrons in part)); Friese 1912: 94 (distribution); PAPUA NEW GUINEA, Sepik-Bivak; Ihering 1912: 3 (citation (as laevicops!)); Strand 1913a: 149-150 (see iridipennis); Friese 1914a: 12-13 (distribution, floral record, uncertain identity (as iridipennis)); INDONESIA, Bandung; Batavia; Buitenzorg; Nongkodjadjar; Nusa Kambangan; Salatiga; Samarang; Floral record: Canna flavescens; Friese 1915: 4 (distribution, uncertain identity (as praeterita)); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Etna Bai; Jasa; Merauke; Njao; Zoutbron; Cockerell 1918: 385 (comparative notes); Cockerell 1919c: 77, 79 (distribution, previous misidentifications); Cockerell 1919a: 121 (comparative note, uncertain identity (as iridipennis)); INDONESIA, Java; Cockerell 1922a: 279 (distribution); Cockerell 1922b: (comparative note); Cockerell 1923: 241 (key to species); INDONESIA, Aru; Cockerell 1925a: 492 (comparative note); Cockerell 1925b: (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 (comparative 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 specimens examined 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-


RASMUSSEN


tion, uncertain identity (as iridipennis)); CAMBODIA, Stung Treng; Moure 1961: 208 (morphology, systematic 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; Palawan; 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 & Manning 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; THAILAND; 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; Sakagami et al. 1983d: (see minangkabau); Sakagami et al. 1983c: (reproduction); Inoue et al. 1984b: (see minangkabau); 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 (comparative note); 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 (comparative note); Starr & Sakagami 1987: 97 (citation); Salmah et al. 1987: 384 (synonymy); Uji 1987: (pollination, uncertain identity (as irridipenis!)); INDONESIA, E Kalimantan; Floral record: Nephelium lappaceum; Camargo 1988: 372 (distribution, synonymy); MALAYSIA, Sandakan, Penang; Sakagami & Inoue 1989: 615 (comparative note); Roubik 1989: 181, 205; Sakagami et al. 1989: 272 (misidentification); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDIA (misidentification); INDONESIA, Java; Sumatra; MALAYSIA; MYANMAR; THAILAND; Sakagami & Inoue 1990: (oviposition); Salmah et al. 1990: (distribution, nest (hollow pillars; cracks), also as laeviceps small); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Engels & Imperatriz-Fonseca 1990: (see minangkabau); Salmah et al. 1991: 196 (synonymy); Prasad & Hemanth 1992: (predator (Ardeola grayii: Ardeidae: Ciconiiformes) (as leviseps!)); Michener & Roubik 1993: 260 (comparative note, morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cardale 1993: 323-324 (see sapiens); 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); BRUNEI, 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-Neto 1997: 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: (floral records (not by species), foraging); Peters et al. 1999: 382 (citation); Crane 1999: 89; van Veen & Sommeijer 2000: 70, 74, 75 (citation); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Wu 2000: 387, 391-392, plate (key to species, morphology); Zherikhin & Ross 2000: 6, 1314 (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 (pollination); Floral record: Neuwiedia veratrifolia; Eltz et al. 2002a: 30 (distribution, foraging, nest, population);



43


MALAYSIA, Danum Valley; Sepilok; Deramakot; Nagamitsu & Inoue 2002: 309 (citation); San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Camargo & Pedro 2003a: (comparative notes, systematics, taxonomy); Hohmann 2003: (communication, nest); Biesmeijer & Slaa 2004: 148 (citation, foraging); Franck et al. 2004: (genetics, microsatellite); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Wattanachaiyingcharoen et al. 2004: (diversity, nest); Nagamitsu & Inoue 2005: 77, 78, 82, 83, 84, 85 (distribution, foraging); Chinh et al. 2005: 363, 364, 367-368, 370 (nest); VIETNAM, Cucphuong National Forest, Red river delta; Klakasikorn et al. 2005: 2, 3, 5, 6 (floral record); Floral record: Hevea brasiliensis; Roubik 2006: 134 (nest); Tangmitcharoen et al. 2006: (pollination); Floral record: Tectona grandis; Inson & Malaipan 2006: (distribution, foraging, nest, uncertain identity (as iridipennis)); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Ardisia pyramidalis; Diospyros durionoides; Ixora javanica; Pipturus; Thummajitsakul et al. 2008: 451 (genetic variation). Tetragonula melanocephala (Gribodo, 1893) Trigona melanocephala Gribodo 1893: 264: Holotype (MSNG, worker): The authentic holotype of this taxon is labeled "Bandjarmas" (?=Bandjarmasin, near Liangtelan) (F. Penati, pers. com.) (taxonomy); Type locality: MALAYSIA "Liangtelan (Borneo)" (1 worker); Dalla Torre 1896: 580 (citation); Cockerell 1919b: 242 (distribution); MALAYSIA, Sandakan; Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1937: 282, 283, 290, 315*, 316, 326 (key to species, taxonomy); Schwarz 1939a: 85, 93, 120* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 210 (systematic position); Yoshikawa et al. 1969: 172, 174 (distribution); MALAYSIA, Sabah, Touaran; Sakagami & Inoue 1985: 181 (comparative note); Sakagami et al. 1985: (distribution); Sakagami et al. 1990: 131 (distribution); Inoue et al. 1995: 97, 98 (pollination); Floral record: Dendrobium setifolium; Neuwiedia borneensis; Roubik 1996: 61, 62 (distribution, predator (Helarctos malayanus: Ursidae: Carnivora)); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433, 435, 436, 437 (pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Eltz et al. 2001a: (pollen foraging); Eltz et al. 2001b: 343, 345 (foraging); MALAYSIA, Borneo, Sabah; Eltz et al. 2002b: 28 (comparative note); Eltz et al. 2002a: 30, 31, 32 (distribution, foraging, nest, population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Floral record: Rhizophora apiculata; Nagamitsu & Inoue 2002: 305, 307, 308, 310, 312 (foraging); Eltz et al. 2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz 2004: 322 (abundance); Michener & Boongird 2004: 144, 145, 146 (comparative note); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 77, 78, 79, 80, 81, 82, 83, 84, 85, 86, 88 (distribution, foraging); Rasmussen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Diospyros durionoides. Trigona testaceinerva Cameron 1908: 193, 195: Type (BMNH 17b.1109) (key to species, taxonomy); Type locality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker); Schwarz 1937: 315, 326; Schwarz 1939a: 120 (synonymy); Moure 1961: 210 (synonymy); Michener & Boongird 2004: 146 (comparative note). Tetragonula melina (Gribodo, 1893) 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 carry no locality label (F. Penati, pers. com.) (taxonomy); Type locality: MALAYSIA "Liangtelan (Borneo)" (2


RASMUSSEN


workers); "Perak (Malacca)" (3 workers); Dalla Torre 1896: 580 (citation); Friese 1909("1908"): 354, 357 (comparative note, distribution); MALAYSIA, Borneo, Liangtelan; Malaka, Perak; Friese 1909: 272, 273, 276 (citation, key to species); MALAYSIA, Borneo, Liangtelan; Malacca; Perak; Cockerell 1918: 387 (distribution, key to species); MALAYSIA, Sandakan; Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Schwarz 1932: 245; Schwarz 1937: 283, 291, 316, 316*, 326 (key to species, taxonomy); Floral record: Melastoma; Schwarz 1939a: 85, 90, 93, 120* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 210 (systematic position); Sakagami 1978: 166-194, 198*, 226, 236, 237, 238 (taxonomy); MALAYSIA; THAILAND; Sakagami et al. 1983b: 2, 8, 20 (nest); Sakagami & Inoue 1985: 181 (comparative note); Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Starr & Sakagami 1987: 97 (citation); Camargo 1988: 372 (distribution); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener & Roubik 1993: 260 (comparative note, morphology); Salmah et al. 1996: 410 (comparative note); Roubik 1996: 61, 62, 63 (distribution, nest); BRUNEI, Kuala Belalong; Dollin et al. 1997: 886 (comparative note); Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Liow et al. 2001: 184; Eltz et al. 2001a: (pollen foraging); Eltz et al. 2001b: 343, 345 (foraging); MALAYSIA, Borneo, Sabah; Kocyan & Endress 2001: 848, 856, 864 (pollination); Floral record: Neuwiedia javanica; Eltz et al. 2002b: 28 (comparative note); Eltz et al. 2002a: 30, 31, 32 (distribution, foraging, nest, population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Floral record: Rhizophora apiculata; Nagamitsu & Inoue 2002: 305, 307, 308, 310, 312 (foraging); Eltz et al. 2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 77, 80, 82, 83, 84, 85, 86, 88 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics). Tetragonula mellipes (Friese, 1898) Trigona mellipes Friese 1898: 428, 429: Syntypes (ZMHB, 1 worker, 1 male): examined, "C. Austr. / v. Müller.93", "Trigona / mellipes / det. Friese 1897", "Zool. Mus. / Berlin", "Coll. / Friese" (only worker). Friese stated the type locality was "S. Australia", probably an error for C. Australia, as ZMHB specimens match the original description and the taxon is not known from southern Australia (key to species, taxonomy); Type locality: AUSTRALIA "S. Australia" (worker, male); Friese 1909: 272, 273, 276 (citation, key to species); Cockerell 1930b: 150 (taxonomy); Schwarz 1932: 245; Rayment 1932f: 107 (taxonomy (as melipes!)); Rayment 1932f: 107; Rayment 1935: 735 (key to species); Michener 1965: 231; Dollin & Dollin 1988: 223 (type locality); Gibbons 1992: (type depository); Cardale 1993: 324 (distribution, list); AUSTRALIA, N coastal; N Gulf; Northern Territory; Western Australia; Yamane et al. 1995: 294 (synonymy); Dollin et al. 1997: (nest, revision, taxonomy); AUSTRALIA, Northern Territory; Western Australia; Dollin 1997: (characteristics); Crane 1999: 89; Heard & Dollin 2000: (bee-keeping); Green et al. 2001: 91 (microsatellite); Serrão 2001: (morphology (proventricular structure)); Palmer et al. 2002: (maternity); Franck et al. 2004: (genetics, microsatellite, nest); Tóth et al. 2004: 6 (citation); Rasmussen & Cameron 2007: (systematics); Klumpp 2007: 4, 82. Tetragonula minangkabau (Sakagami & Inoue, 1985) Trigona (Tetragonula) minangkabau f. darek Sakagami & Inoue 1985: 176, 177, 178, 179, 180, 181, 186:



45


Nomen nudum. The name is unavailable for taxonomic purposes, as it must be deemed to be infrasubspecific (ICZN, art. 15.2 and 45.6.3.) (taxonomy); Sakagami et al. 1990: (distribution, key to species, variation); 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, ix 1981 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, reproduction (as laeviceps)); Salmah et al. 1984: (development (as laeviceps)); Sakagami et al. 1985: (distribution, 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); Salmah et 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 & Inoue 1990: 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: Neuwiedia borneensis; 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 depositories); Inoue et al. 1999: 208, 217 (citation, comparative table); Ranger & O'Donnell 1999: 189 (comparative 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 Forest 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: Formicidae: Hymenoptera)); Nagamitsu & Inoue 2005: 77, 78 (distribution, foraging); Hofstede & Sommeijer 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). Tetragonula minor (Sakagami, 1978) Trigona (Tetragonula) minor Sakagami 1978: 166-194, 213-214*, 224, 226, 227, 236, 237, 238: Holotype (SEHU); 32 paratypes (SEHU) (taxonomy); Type locality: MALAYSIA "Kuala Lumpur-2" (33 workers); Sakagami & Inoue 1985: 186 (comparative note); Sakagami et al. 1985: (distribution); Sakagami et al. 1990: 131 (distribution); Tadauchi et al. 1998: 244 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 2-3, 4, 5, 6 (floral record, morphology); Floral record: Homalium grandiflorum; Tectona grandis; Rasmussen & Cameron 2007: (systematics).


RASMUSSEN


Tetragonula pagdeni (Schwarz, 1939) Trigona (Tetragona) fusco-balteata variety pagdeni Schwarz 1939a: 85, 93, 96, 110-111*, 113: Holotype (USNM 53564, male): examined, "Nakon / SriTamarat / Siam 7/5/28", "HughSmith / coll", Type No. / 53564 / U.S.N.M.", "Holotype", "Trigona / fusco-balteata / var. pagdeni / H.F. Schwarz"; paratype (AMNH (1)) (distribution, key to species, taxonomy); Type locality: THAILAND "SIAM.-Nakon Sri Tamarat, July 5, 1928 (Hugh M. Smith)" (workers, males); "Singora, June 1929 (Hugh M. Smith)" (workers); Krombein 1951: 295 (comparative note); Schwarz 1951: 63 (distribution); THAILAND, Bangkok; Moure 1961: 208-9 (systematic position); Sakagami & Yoshikawa 1961: 435, 436-437 (comparative note, nest, taxonomy); CAMBODIA, Udong; THAILAND, Cheng Mai; Doi Dao; Kamphangpet; Korat; Yoshikawa et al. 1969: 168, 169 (nest); Wille & Michener 1973: 14, 24, 59 (nest); Sakagami 1978: 166194, 200, 211, 212, 213, 218, 219-223*, 224, 225, 226, 227, 228, 234, 236, 237, 238, 239, 240, 247, plate 5 (taxonomy); Sakagami et al. 1983b: 2, 6-7, 8, 20 (nest); Sakagami et al. 1983a: 103, 105, 106 (comparative note); Sakagami et al. 1983d: (oviposition behavior); Sakagami & Inoue 1985: 177 (comparative note); Sakagami et al. 1985: (distribution); Kiew 1985: (luring (unwashed shorts)); Yong 1986: 627 (comparative note); Sakagami & Khoo 1987: 213-214 (morphology); Sakagami & Inoue 1987: 626 (comparative note); Starr & Sakagami 1987: 97 (citation); Sakagami et al. 1990: 130, 131 (comparative note, distribution); Sakagami & Inoue 1990: 201, 202, 203, 204, 205, 206, 210, 211, 212, 215 (comparative note); Michener & Roubik 1993: 260 (comparative note, morphology); Yamane et al. 1995: 276 (comparative note); Drumond et al. 1996a: 399 (behavioral comparison); Dollin et al. 1997: 864 (comparative note, distribution); MALAYSIA; THAILAND; Wu 2000: 387, 390-391 (key to species, morphology); Franck et al. 2004: (genetics, microsatellite); Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics); Thummajitsakul et al. 2008: (genetic variation). Tetragonula pagdeniformis (Sakagami, 1978) Trigona (Tetragonula) pagdeniformis Sakagami 1978: 166-194, 209, 211-213*, 226, 227, 229, 234, 236, 237, 238, 240: Holotype (EKYU, not located, nor in SEHU); paratypes (BMNH (5)) (taxonomy); Type locality: THAILAND "Khao Chong-2, Peninsular Thailand" (1 worker, holotype); "Ulu Langat" (1 worker); "Gombak" (1 worker); "Kuala Lumpur" (1 worker); "Kuala Tranggan" (1 worker); "Kuala Tahan" (1 worker); "Taman Negara" (16 workers); "Kanching" (1 worker); "Penang" (8 worker); "Pasoh" (12 workers); "Songkhla" (3 workers); "Khoa Chong" (3 workers, 124 males); Sakagami & Inoue 1985: 177, 186 (comparative note); Sakagami et al. 1985: (distribution); Sakagami & Inoue 1987: 626 (comparative note); Sakagami et al. 1990: 130, 131 (comparative note, distribution); Michener & Roubik 1993: 260 (comparative note, morphology); Dollin et al. 1997: 873 (comparative note); Tadauchi et al. 1998: 244 (type depositories); San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Roubik 2006: 132 (nest). Tetragonula penangensis (Cockerell, 1919) Trigona penangensis Cockerell 1919c: 78, 79: Holotype (AMNH) (comparative notes, key to species); Type locality: MALAYSIA "Penang Island (Baker 9075)" (worker); Schwarz 1937: 309; Schwarz 1939a: 112 (synonymy); Moure 1961: 209 (systematic position); Sakagami 1978: 200 (synonymy); Sakagami et al. 1990: 129 (synonymy); Wu 2000: 391 (synonymy (as penengensis!)).



47


Tetragonula praeterita (Walker, 1860) Trigona præterita Walker 1860: 305-306: Holotype (BMNH 17b.1185, worker): examined, "63 / 52", "Type" (green border), "præterita / W.", "Trigona", "B.M. TYPE / HYM. / 17b.1185" (taxonomy); Type locality: SRI LANKA "Ceylon" (worker); Tennent 1859: 282 (list); Tennent 1861: 454 (list); Dalla Torre 1896: 582 (citation); Bingham 1897: 560, 564 (see laeviceps); Friese 1909("1908"): 354, 355 (see laeviceps); Friese 1915: 4 (see laeviceps); Schwarz 1937: 308, 309; Schwarz 1939a: 88, 111, 112 (synonymy); Moure 1961: 208 (synonymy); Sakagami 1978: 216 (synonymy); Wu 2000: 392 (synonymy); Wijesekara 2001: 153 (synonymy). Tetragonula reepeni (Friese, 1918) Trigona (Tetragonula) latigenalis Sakagami 1978: 166-194, 194-195*, 196, 227, 236, 237, 238, 247, plate 5: Holotype (SEHU) (taxonomy); Type locality: MALAYSIA "Frasers Hill-b, Malaya" (1 worker, holotype); "Malaya, Kepong" (1 worker); "Ulu Langat" (20 workers); "Gombak" (2 workers); "Cameron Highland" (4 workers); "Thailand, Khaophappha-Khaochong" (workers); Sakagami et al. 1985: (distribution, key to species); Sakagami & Khoo 1987: 207, 208, 210, 211, 212 (synonymy); Sakagami et al. 1990: 128 (synonymy); Tadauchi et al. 1998: 243 (type depositories); Wattanachaiyingcharoen et al. 2004: (diversity, nest); Klakasikorn et al. 2005: 6 (citation). Trigona reepeni Friese 1918: 519-520: Lectotype (ZMHB, worker): here designated, "Malacca / Taip. Hills / 2.1912 / Butt.-Reep.", "Trigona / reepeni / 1914 Friese det. / Fr.", "Type" (red label), "Coll. / Friese"; paralectotypes (DEI (1), USNM (1), ZMHB (7)) (taxonomy); Type locality: MALAYSIA "Taiping Hills (Upp. Perak) auf Malakka, im Februar 1912" (several workers); Cockerell 1926a: 224 (comparative note); Schwarz 1939a: 85, 93, 121* (distribution, key to species, taxonomy); Sakagami et al. 1985: (distribution); Sakagami & Khoo 1987: (morphology); Michener 1990: 120 (systematic position); Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; MALAYSIA; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Liow 1999: 110, 127 (distribution); Nagamitsu et al. 1999: (floral records (not by species), foraging); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 77 (distribution, foraging). Tetragonula ruficornis (Smith in Horne & Smith, 1870) Trigona ruficornis Horne & Smith 1870: 185, 194: Holotype (BMNH 17b.1187, worker): examined, "India" (reverse "69 / 86"), "Type" (red border), "Trigona / ruficornis / (Type) Smith", "B.M. TYPE / HYM. / 17b.1187") (common name (bhônga), honey, nest, taxonomy); Type locality: INDIA "Mainpurí, Northwest Provinces of India" (worker); Dalla Torre 1896: 583 (citation); Waterhouse 1903: 135-136 (nest (misidentification; probably fuscobalteata)); Schwarz 1939a: 88, 107 (synonymy); Schwarz 1948: 14, 23, 56, 165; Nogueira-Neto 1949: 121 (bee-keeping); Krombein 1950: 133 (synonymy); Nogueira-Neto 1951: (distribution); INDIA, Uttar Pradesh; Moure 1961: 208 (synonymy); Sakagami 1978: 216 (synonymy); Wu 2000: 392 (synonymy). Melipona smithii Bingham 1897: 560, 563: Unnecessary replacement name for Trigona ruficornis Smith, nec Lamarck. 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. ruficrus Latreille. BMNH has a labeled type for this replacement name (17b.1129) (distribution, key to species, taxonomy); INDIA, Mainpuri, NW provinces; MYANMAR, Tenasserim; Bingham 1905: 59 (citation, distribution); Schwarz 1939a: 88, 107, 108 (synonymy); Schwarz 1948: 14, 23, 56; Nogueira-


RASMUSSEN


Neto 1949: 121 (synonymy); Krombein 1950: 133 (synonymy); Moure 1961: 208 (synonymy); Jain & Kushwaha 1972: (morphometrics); Wille & Michener 1973: 14, 25, 49, 60 (nest); Sakagami 1978: 216 (synonymy); Starr & Sakagami 1987: 97 (citation); Nogueira-Neto 1954: 426 (citation); Wu 2000: 392 (synonymy). Tetragonula sapiens (Cockerell, 1911) Trigona sapiens Cockerell 1911: 176: Holotype (ANIC, worker) (comparative notes, taxonomy); Type locality: SOLOMON ISLANDS "Solomon Islands, July-August, 1909 (Froggatt)" (worker); Cockerell 1929e: 242-243 (comparative note, distribution, taxonomy); SOLOMON ISLANDS, Lavoro Plantations, Gaudalcanal Is.; New Georgia, W. Solomon; Cockerell 1936: 225 (distribution); SOLOMON ISLANDS, Halaita, Nggela; Schwarz 1939a: 111, 113 (synonymy); Krombein 1949: 14 (distribution); Krombein 1951: 295 (synonymy); Moure 1961: 209 (systematic position); Michener 1965: 231 (citation); Sakagami 1978: 221, 222; Dollin & Dollin 1983: (nest (as laeviceps)); Sakagami & Inoue 1987: 626 (comparative note); Starr & Sakagami 1987: (common name (kiyot), nest); Roubik 1990c: 663 (citation); Michener & Roubik 1993: 260 (comparative note, morphology); Cardale 1993: 323-324 (distribution, list (as laeviceps)); AUSTRALIA, NE coastal; N coastal; Northern Territory; Queensland; Adelaide River; Cape York; Gordonvale; Thursday Is.; Oliveira et al. 1995: 20 (citation); Snelling 1996: 131 (distribution); Dollin et al. 1997: (nest, revision, taxonomy); AUSTRALIA, Queensland; INDONESIA, Moluccas; "NEW GUINEA"; PHILIPPINES; SOLOMON ISLANDS; Dollin 1997: (characteristics); Green et al. 2001: 91 (microsatellite); Palmer et al. 2002: (maternity); Franck et al. 2004: (genetics, microsatellite); Cameron et al. 2004: 2358 (nest); Sung et al. 2004: 268 (citation); Rasmussen & Cameron 2007: (systematics); Klumpp 2007: 4, 82; Wallace et al. 2008: (dispersal, pollination); Floral record: Corymbia torelliana. 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). Tetragonula sarawakensis (Schwarz, 1937) Trigona sarawakensis Schwarz 1937: 283, 290, 313-315*, 316, 328, plate 2, 4: Holotype (BMNH 17b.1110); paratypes (AMNH, BMNH?) (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, 4000 ft." (2 workers Oct. 18, 1932; 2 workers Oct. 19, 1932; 1 worker Oct. 22, 1932); Schwarz 1939a: 85, 92, 106* (distribution, key to species, taxonomy); Schwarz 1948: 118; Sakagami 1959: 119120 (distribution, morphology, variation); SINGAPORE; Moure 1961: 210 (morphology, systematic position); Yoshikawa et al. 1969: 168, 169 (nest); Wille & Michener 1973: 14, 25, 59 (nest); Sakagami 1978: 166-194, 210-211*, 228, 229, 230, 236, 237, 238 (taxonomy); Sakagami et al. 1983b: 2, 10 (nest); Sakagami et al. 1983a: 106 (comparative note); Sakagami & Inoue 1985: 174, 176, 177, 178, 179, 180, 181, 182*, 184, 186 (key to species, taxonomy); Sakagami et al. 1985: (distribution); Starr & Sakagami 1987: 97 (citation); Sakagami et al. 1990: 131; Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics). Tetragonula sirindhornae (Michener & Boongird, 2004) Trigona (Tetragonula) sirindhornae Michener & Boongird 2004: 143-146: Holotype (SEMC, worker); paratypes (AMNH (3), Insect Museum, Chakthong Building, Department of Agriculture, Bangkok, Thailand (3), SEMK (11) (taxonomy); Type locality: THAILAND "Thailand, on the Isthmus of Kra: Ranong Province: Muang district, 9–13 November 2002" (11 workers); "Ngow Waterfall National Park" (7 work-



49


ers); Klakasikorn et al. 2005: 5, 6 (citation); Rasmussen & Cameron 2007: (systematics). Tetragonula testaceitarsis (Cameron, 1901) Trigona testaceitarsis Cameron 1901: 36: Type (BMNH 17b.1121) (taxonomy); Type locality: MALAYSIA "Patani, Malay Peninsula" (worker); Cockerell 1923: 242 (key to species); Moure 1961: 208 (morphology, systematic position); Sakagami 1978: 200 (as testaceicornis!); Sakagami et al. 1990: 129 (synonymy (as testaceicornis!)); Wu 2000: 391 (synonymy (as testaceicornis!)). Tetragonula valdezi (Cockerell, 1918) Trigona valdezi Cockerell 1918: 387: Holotype (USNM 40249, worker): examined; 3 paratypes (USNM) (distribution, key to species, uncertain identity); Type locality: SINGAPORE "Singapore" (unknown); Cockerell 1919c: 78, 79 (comparative notes, key to species); SINGAPORE; Cockerell 1929c: 592 (comparative note, floral record, taxonomy); CAMBODIA, Angkor; Nan; Pah Meeung; Floral record: Poinsettia; Cockerell 1934: 62, fig. 6 (illustration (mandible)); Schwarz 1937: 309; Schwarz 1939a: 85, 88, 90, 93, 110, 113, 116, 117, 118 (distribution, key to species, taxonomy); Schwarz 1939b: 151, 152 (distribution, variation); MICRONESIA, Caroline Islands, Toloas; Truk; Cockerell 1939a: 61, 62, 64 (distribution ("presumably introduced by man"), variation); MICRONESIA, Dublon Island; Tarik Island; Schwarz 1948: 118, 233; Krombein 1950: 102, 133, 135 (distribution, morphology); MICRONESIA, Caroline Islands; Schwarz 1951: 63 (distribution); CAMBODIA, Angkor; Mt. Suept; Sakagami 1960: 146 (comparative note); Moure 1961: 208 (morphology, systematic position); Michener 1961: 28 (comparative note); Sakagami & Yoshikawa 1961: 435 (comparative note, distribution); CAMBODIA, Phnom Penh; LAOS, Thakhek; THAILAND, Mae Fack; Baltazar 1966: 407 (distribution); PHILIPPINES, Luzon, Laguna; Mindanao, Davao; Sakagami 1978: 166, 197, 200, 201, 205, 206, 208 (synonymy); Sakagami et al. 1983a: 101 (comparative note); Camargo 1988: 372 (synonymy); Sakagami et al. 1990: 129, 130 (synonymy, variation); Ikudome & Kusigemati 1996: 20 (taxonomy); Wu 2000: 391 (synonymy); Klakasikorn et al. 2005: 6 (citation). Tetragonula zucchii (Sakagami, 1978) Trigona (Tetragonula) zucchii Sakagami 1978: 166-194, 196, 197, 198, 208-210*, 228, 229, 236, 237, 238, 247, plate 5: Holotype (SEHU) (nest, taxonomy); Type locality: MALAYSIA "Fraser's Hill-b, Malaya" (9 workers, holotype, 8 paratypes); "Cameron Highlands" (8 worker); "Gombak" (1 worker); "Penang" (1 worker); "Templer Park" (17 workers); Yoshikawa et al. 1969: 173 (citation (as iridipennis)); Sakagami et al. 1985: (distribution); Sakagami et al. 1990: 131 (distribution); Tadauchi et al. 1998: 244 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution, foraging); Rasmussen & Cameron 2007: (systematics). 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: (taxonomy); Roubik 2006: 126, 127, 130, 132 (nest).

Tetrigona Tetrigona apicalis (Smith, 1857)


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Trigona apicalis Smith 1857: 51: Holotype (OUMNH); additional putative type (BMNH 17b.1188) (taxonomy); Type locality: MALAYSIA "Borneo (Sarawak)" (worker); Smith 1871: 395 (distribution); MALAYSIA, Borneo; Dalla Torre 1896: 575 (citation); Bingham 1897: 560, 562 (common name (dammar bee), distribution, key to species, taxonomy (see also binghami)); MALAYSIA, Borneo; MYANMAR, Tenasserim; Waterhouse 1903: 133-134, plate 4 (morphology (male), nest (as collina)); Beccari 1904: 162 (nest); Bingham 1905: 59 (citation, distribution); Cockerell 1918: 387 (see binghami); Cockerell 1920a: 116 (comparative note); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Cockerell 1923: 241 (key to species); Cockerell 1927: 541 (comparative note); Salt 1929: 454 (citation); Dover 1929: 62 (citation, distribution (as collina)); MALAYSIA, Semangko Pass, Selangor Pahang; Kedah Peak; Friese 1933a: 45, 46 (comparative note, distribution); INDONESIA, Adjongan; Schwarz 1937: 282, 283, 288, 297, 298, 300-302*, 303, 304, 326 (key to species, taxonomy); Schwarz 1939a: 85, 94, 95, 100, 121, 122, 126-128*, 129, 130, 133 (distribution, key to species, taxonomy); Schwarz 1948: 19, 23, 91, 117, 141, 165, 290; Pagden 1957: 17-18 (flight); Sakagami 1960: 146 (distribution); CAMBODIA, Stung Treng; Moure 1961: 215, 216 (systematic position); Wille 1961: 117, 121; Sakagami & Yoshikawa 1961: 439 (distribution, nest (in Ficus: Moraceae; only fig. 29) (as collina)); THAILAND, Doi Suthep; Kamphangpet; Mae Fack; Mae Hoi; Thoen; Yoshikawa et al. 1969: 168, 173, 174 (distribution, nest); MALAYSIA, George Town, Penang; Wille & Michener 1973: 13, 23, 47, 58, 87 (nest); Sakagami 1975: 60-62*, 63 (distribution, morphology, variation); LAOS; MALAYSIA; THAILAND; VIETNAM; Sakagami 1978: 239 (comparative note); Sakagami 1982: 389 (citation); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1983b: 2, 5-6, 13, 19, 20 (nest); Inoue et al. 1984a: 136 (comparative note); Sakagami et al. 1985: (distribution, key to species); Sakagami & Khoo 1987: 207, 208, 209, 210 (morphology); Camargo 1988: 372 (distribution); MALAYSIA, Sandakan; SINGAPORE; Roubik 1989: 417; Michener 1990: 95, 105, 116, 120, 124, 125, 126, 127 (illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Java; Sumatra; MALAYSIA, Borneo; MYANMAR; THAILAND; Salmah et al. 1990: (distribution, nest (hollow trunks of Ficus: Moraceae; Durio zibethinus: Bombacaceae; crevices)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Roubik 1993: 541 (illustration, nest); Michener & Roubik 1993: 260 (comparative note, morphology); Cunha 1994: (systematics); Yamane et al. 1995: 294 (synonymy); Roubik 1996: 61, 62 (distribution, nest); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: Santiria laevigata; Michener 1997: 57 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener 2000: 801; Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Chinajariyawong & Saiboon 2000: (distribution, nest); THAILAND, Songkhla; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); 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, 82, 83, 84, 85 (distribution, foraging); Klakasikorn et al. 2005: 2, 3, 6 (floral record); Floral record: Dipterocarpus alatus; Tectona grandis; Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Jongjitvimol et al. 2006: (foraging); Jongjitvimol & Wattanachaiyingcharoen 2006: (foraging); THAILAND, Phitsanulok; Floral record: Agave angustifolia; Alangium salviifolium; Coccinia grandis; Costus speciosus; Croton roxburghii; Delonix regia; Erythrina stricta; Fernandoa adenophylla; Ipomoea aquatica; Ixora grandifolia; Merremia vitifolia; Millettia brandisiana; Mimosa pigra; Mimosa pudica; Paederia linearis; Ruellia tuberose; Senna siamea; Tectona grandis; Torenia fournieri; Tridax procumbens; Rasmussen & Cameron 2007: (systematics); Michener 2007a: 826; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative



51


note). Trigona hemileuca Cockerell 1929b: 140: Holotype (BMNH 17b.1098) (comparative note, taxonomy); Type locality: THAILAND "Siam: Nan, Dec. 31 and Jan. 31 (Cockerell)" (worker); CAMBODIA, Ban Maa Hia; Mekami river; Cockerell 1929c: 591 (distribution); CAMBODIA, W Weing Sa; Schwarz 1937: 300, 302; Schwarz 1939a: 126 (synonymy); Moure 1961: 216, 217 (morphology, systematic position); Sakagami 1975: 60, 61 (synonymy); Sakagami et al. 1990: 127 (synonymy). Trigona sericea Friese 1933a: 45-46: Lectotype (ZMHB, worker): here designated, "Borneo / Sanggau / 247-32", "Trigona / sericea / 1925 Friese det. / Fr."; paralectotypes (ZMHB (5), DEI (4)) (taxonomy); Type locality: MALAYSIA/INDONESIA "Meliau (Borneo) 4. Juni 1932"; "Sanggau (Borneo) 24. Juli 1932" (several workers); Schwarz 1937: 282, 300, 301, 302, 327; Schwarz 1939a: 126 (synonymy); Moure 1961: 216 (synonymy). Tetrigona binghami (Schwarz, 1937) Trigona apicalis variety binghami Schwarz 1937: 288, 300, 301, 303-304*, 328, plate 2, 5, 7: Holotype (BMNH 17b.1142); paratypes (AMNH, BMNH, USNM) (key to species, taxonomy); Type locality: MYANMAR "Tenasserim: Dawnat Range, Jan., 1891 (Col. C.T. Bingham)" (1 worker); Bingham 1897: 560 (as apicalis, but most belong to this taxon); Cockerell 1918: 387 (distribution, key to species (as apicalis)); MALAYSIA, Sandakan; Schwarz 1939a: 85, 94, 126, 128-129* (distribution, key to species, taxonomy); Schwarz 1948: 117; Moure 1961: 216 (morphology, systematic position); Sakagami 1975: 61 (variation); Sakagami et al. 1990: 131 (distribution); Eltz et al. 2001b: 343, 345 (foraging); MALAYSIA, Borneo, Sabah; Eltz et al. 2002a: 30, 31, 32 (distribution, foraging, nest, population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Floral record: Rhizophora apiculata; Eltz et al. 2003: (nest, tree preference); Eltz 2004: 322 (abundance); Nagamitsu & Inoue 2005: 76 (distribution, foraging); Klakasikorn et al. 2005: 2, 3, 5, 6 (floral record, morphology); Floral record: Tectona grandis; Rasmussen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Archidendron jiringa; Ardisia elliptica; Cassia fistula; Colocasia gigantea; Diospyros durionoides; Allophylus rubifolius; Peltophorum pterocarpum. Tetrigona melanoleuca (Cockerell, 1929) Trigona melanoleuca Cockerell 1929b: 140-141: Holotype (BMNH 17b.1119). Possibly a junior synonymon of vidua. See below (comparative note, taxonomy); Type locality: THAILAND "Siam: Nan, Jan. 13, 1928 (Alice Mackie)" (worker); Schwarz 1937: 300, 302, 304; Schwarz 1939a: 85, 94, 95, 98, 129*, 130 (distribution, key to species, taxonomy); Schwarz 1948: 117; Moure 1961: 216-217 (morphology, systematic position); Sakagami 1975: 61, 62-64* (distribution, morphology, variation); LAOS; THAILAND; Sakagami et al. 1985: (distribution, key to species); Michener 1990: 124, 126, 127 (illustration, morphology, systematic position); Sakagami et al. 1990: 131, 136 (distribution, key to species); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Nagamitsu & Inoue 2005: 76, 83 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics). Tetrigona peninsularis (Cockerell, 1927) Trigona apicalis peninsularis subsp.n. Cockerell 1927: 541: Holotype (BMNH 17b.1099) (taxonomy, varia-


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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 syntype examined, "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 1818 and spent years collecting in the Indo-Malayan region. The label date is wrong as they were still in France in 1815, but did reside for long periods in India later (Claeys: 1954) and I consider the label provenance merely the port of embarkation of the specimen, rather than the collecting locality. The type is similar to melanoleuca and could be considered a senior synonymon of that. However, melanoleuca is a variable taxon, and is here retained until further studies may document if in fact it is only a single taxon (taxonomy); 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; Castets 1893: 481 (citation); Dalla Torre 1896: 585 (citation); Bingham 1897: 560, 561 (distribution, key to species, taxonomy, uncertain identity); INDIA (misidentification); INDONESIA; MALAYSIA; MYANMAR, Tenasserim; Haddon 1901: 87 (common name (manyi), bad omen for Ibans ("if a swarm of bees settled underneath a house that had recently been built it would be considered a bad sign"), uncertain identity); Bingham 1905: 59 (citation, distribution); Friese 1914a: 13 (distribution (maybe itama?)); INDONESIA, 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; Moure 1961: 217 (morphology, systematic position); Sakagami & Khoo 1987: 207 (comparative note); Camargo 1988: 372 (distribution); INDONESIA, Sumatra; Wu 2000: 386, 390, plate (key to species, morphology).

Meliponini Meliponini indet. Westwood 1840: 282-284 (honey); AUSTRALIA; Anderson 1890: 23-24 (common name (dammer; pwai-ngyet), wax); Veth & Ritsema 1892: 6, 16 (distribution); Bingham 1897: 559-561 (key to species, taxonomy (as Melipona)); Waterhouse 1900: (nest); Cockerell 1920a: 116 (distribution); Cockerell 1930b: 149; Rayment 1932a: (morphology, nest); Bibby 1947: 78 (distribution); Michener 1961: (common name (native bee; sweat bee), nest); Michener 1965: 231 (citation); Michener 1965: 230; Smythe 1970: (pollination (as Trigona)); Floral record: Cymbidium madidum; Dendrobium; Frith 1978: (nest (in Rhizophora apiculata: Rhizophoraceae); predator (Oecophylla smaragdina: Formicidae: Hymenoptera)); Schneider 1982: (pollination); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Pande & Sumita 1987("1985")a: 14 (pollination); Floral record: Cajanus cajan; Pande & Sumita 1987("1985")b: 42, 43 (insecticide);



53


Raju & Reddi 1989: (pollination); Floral record: Anisomeles indica; Anisomeles malabarica; Heard et al. 1990: 1006, 1007, 1012 (pollination); Floral record: Anacardium occidentale; Boonithee et al. 1991: (pollination); Adams et al. 1992: (orchid pollination); Appanah 1993: 466 (pollination); Adams & Lawson 1993: (pollination (as Trigona)); Floral record: Caladenia carnea; Cymbidium madidum; Cymbidium suave; Dendrobium adae; Dendrobium kingianum; Dendrobium monophyllum; Dendrobium speciosum; Dendrobium lichenastrum; Sarcochilus moorei; Pomatocalpa macphersonii; Heard 1993: (pollination (as spp.)); Floral record: Macadamia integrifolia; Forster et al. 1994: (pollination); Rinderer et al. 1996: (foraging); Aluri & Subba Reddi 1996: (pollination); Momose et al. 1998: (pollination); Heard 1998: (pollination (as spp.)); Floral record: Macadamia integrifolia; Crane 1999: 86-90 (honey); Viraktamath et al. 1999: (pollen); Heard 1999b: (pollination (as spp.)); Floral record: Mangifera indica; Yumoto 2000: (pollination); Heard 2000: (pollination (as spp.)); Jagadish et al. 2002: (common name (dammer bee), pollination); Floral record: Hygrophila auriculata; Leucas aspera; Tangmitcharoen et al. 2002; Chourykaew et al. 2002: 35, 36, 38, 41 (pollination (contrary to conclusion)); Floral record: Afgekia sericea; Cunningham et al. 2002: 898 (citation); Corlett 2004: (pollination); Maynard et al. 2004: 319 (citation); Momose & Hamid 2005; Blamires et al. 2008: (predator (Argiope keyserlingi: Araneidae: Araneae)).

Incertae sedis Melipona basimaculata Bingham 1903: vii: Holotype (BMNH 17b.1127, male) (taxonomy); Type locality: MALAYSIA "Biserat, Jalor; Telôm, Perak-Pahang boundary, 4,000 feet" (male); Melipona basimaculata Bingham 1905: 60, plate A11 (citation). Trigona pallidicincta Cockerell 1918: 386: Holotype (BMNH 17b.1134, male) (key to species, comparative notes, taxonomy); Type locality: SINGAPORE "Singapore (Baker)" (male).

Nomina nuda Melipona pulla Illiger 1806: 158: Nomen nudum. The name is unavailable for taxonomic purposes, as no description 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 no description 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 no description 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 of the taxon was provided (ICZN, art. 15.1). Trigona (Pacificotrigona) okazawai irianensis Tadauchi et al. 1998: 245: Nomen nudum. The name is


RASMUSSEN


unavailable for taxonomic purposes, as no description or definition of the taxon was provided (ICZN, art. 15.1) (type depositories).

Appendix 1. Plants visited by stingless bees Acacia caesia (Mimosaceae) Afgekia sericea (Leguminosae) Agave angustifolia (Agavaceae) Ageratum (Asteraceae) Ageratum conyzoides (Asteraceae) Alangium salviifolium (Alangiaceae) Allium (Alliaceae) Allophylus rubifolius (Sapindaceae) Alocasia (Araceae) Alphitonia (Rhamnaceae) Amaranthus (Amaranthaceae) Anacardium occidentale (Anacardiaceae) Angophora (Myrtaceae) Anisomeles indica (Lamiaceae) Anisomeles malabarica (Lamiaceae) Antigonon (Polygonaceae) Antigonon leptopus (Polygonaceae) Archidendron jiringa (Leguminosae) Archontophoenix alexandrae (Arecaceae) Ardisia elliptica (Myrsinaceae) Ardisia pyramidalis (Myrsinaceae) Areca vestiaria (Arecaceae) Asystasia gangetica (Acanthaceae) Averrhoa carambola (Oxalidaceae) Azadirachta indica (Meliaceae) Bambusa vulgaris (Poaceae) Banksia (Proteaceae) Boehmeria (Urticaceae) Bombax (Bombacaceae) Bombax ceiba (Bombacaceae) Brassica campestris (Brassicaceae) Buddleja asiatica (Buddlejaceae) Butea monosperma (Leguminosae) Caesalpinia (Leguminosae) Cajanus cajan (Leguminosae) Caladenia carnea (Orchidaceae) Calamus (Araceae) Callistemon (Myrtaceae) Callistemon citrinus (Myrtaceae) Calophyllum (Clusiaceae) Canna flavescens (Cannaceae) Caryota bacsonensis (Arecaceae) Cassia (Leguminosae) Cassia bakeriana (Leguminosae) Cassia fistula (Leguminosae) Ceiba pentandra (Bombacaceae) Citrullus lanatus (Cucurbitaceae) Citrus (Rutaceae) Citrus limon (Rutaceae) Claoxylon (Euphorbiaceae) Clausena (Rutaceae)



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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Cleome (Capparaceae) Coccinia (Cucurbitaceae) Coccinia grandis (Cucurbitaceae) Cocos (Arecaceae) Cocos nucifera (Arecaceae) Coffea canephora (Rubiaceae) Colocasia gigantea (Araceae) Corymbia torelliana (Myrtaceae) Costus speciosus (Costaceae) Crateva magna (Capparaceae) Croton (Euphorbiaceae) Croton roxburghii (Euphorbiaceae) Cucumis sativus (Cucurbitaceae) Cupaniopsis (Sapindaceae) Cupaniopsis anacardioides (Sapindaceae) Cyanotis (Commelinaceae) Cycas (Cycadaceae) Cycas media (Cycadaceae) Cymbidium iridifolium (Orchidaceae) Cymbidium madidum (Orchidaceae) Cymbidium suave (Orchidaceae) Dalbergia lanceolaria (Leguminosae) Daviesia (Leguminosae) Delonix regia (Leguminosae) Dendrobium (Orchidaceae) Dendrobium adae (Orchidaceae) Dendrobium kingianum (Orchidaceae) Dendrobium lichenastrum (Orchidaceae) Dendrobium monophyllum (Orchidaceae) Dendrobium setifolium (Orchidaceae) Dendrobium speciosum (Orchidaceae) Dendrobium teretifolium (Orchidaceae) Dendrobium toressae (Orchidaceae) Dillenia excelsa (Dilleniaceae) Dillenia suffruticosa (Dilleniaceae) Diospyros durionoides (Ebenaceae) Diospyros rhodocalyx (Ebenaceae) Dipterocarpus alatus (Dipterocarpacea) Dryobalanops aromatica (Dipterocarpacea) Durio (Bombacaceae) Elaeis (Arecaceae) Elaeocarpus (Elaeocarpaceae) Elettaria cardamomum (Zingiberaceae) Erigeron (Asteraceae) Erythrina stricta (Leguminosae) Eucalyptus (Myrtaceae) Eucalyptus globulus (Myrtaceae) Eucalyptus torelliana (Myrtaceae) Euphorbia pulcherrima (Euphorbiaceae) Fernandoa adenophylla (Bignoniaceae) Ficus religiosa (Moraceae) Fragaria (Rosaceae) Galphimia glauca (Malpighiaceae) Guizotia abyssinica (Asteraceae) Hamelia patens (Rubiaceae) Helianthus annuus (Asteraceae) Helicteres isora (Sterculiaceae) Hevea brasiliensis (Euphorbiaceae) Hibbertia (Dilleniaceae)


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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Homalium grandiflorum (Flacourtiaceae) Homonoia (Euphorbiaceae) Hopea (Dipterocarpacea) Hygrophila auriculata (Acanthaceae) Impatiens balsamina (Balsaminaceae) Indigofera (Leguminosae) Ipomoea aquatica (Convolvulaceae) Irvingia malayana (Irvingiaceae) Ixora (Rubiaceae) Ixora grandifolia (Rubiaceae) Ixora javanica (Rubiaceae) Jacksonia (Leguminosae) Jasminium (Oleaceae) Jatropha curcas (Euphorbiaceae) Jatropha hastata (Euphorbiaceae) Jatropha podagrica (Euphorbiaceae) Lagerstroemia calyculata (Lythraceae) Lagerstroemia macrocarpa (Lythraceae) Lagerstroemia tomentosa (Lythraceae) Lannea (Anacardiaceae) Lawsonia alba (Lythraceae) Leptospermum (Myrtaceae) Leucaena leucocephala (Leguminosae) Leucas aspera (Lamiaceae) Litchi chinensis (Sapindaceae) Loranthus longiflorus (Loranthaceae) Lycopersicon esculentum (Solanaceae) Macadamia integrifolia (Proteaceae) Mallotus (Euphorbiaceae) Mangifera indica (Anacardiaceae) Manihot esculenta (Euphorbiaceae) Melaleuca (Myrtaceae) Melastoma (Melastomataceae) Melastoma affine (Melastomataceae) Melicope triphylla (Rutaceae) Memecylon malabaricum (Melastomataceae) Merremia vitifolia (Convolvulaceae) Metroxylon sagu (Arecaceae) Millettia brandisiana (Leguminosae) Mimosa (Leguminosae) Mimosa pigra (Leguminosae) Mimosa pudica (Leguminosae) Moringa oleifera (Moringaceae) Mussaenda (Rubiaceae) Neonauclea (Rubiaceae) Nephelium lappaceum (Sapindaceae) Nerine (Amaryllidaceae) Neuwiedia borneensis (Orchidaceae) Neuwiedia javanica (Orchidaceae) Neuwiedia veratrifolia (Orchidaceae) Nicotiana tabacum (Solanaceae) Nipa fruticans (Arecaceae) Nymphaea (Nymphaeaceae) Ochlandra travancorica (Poaceae) Ocimum (Lamiaceae) Oxylobium (Leguminosae) Paederia linearis (Rubiaceae) Passiflora (Passifloraceae) Peltophorum pterocarpum (Leguminosae)



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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. Persoonia (Proteaceae) Persoonia virgata (Proteaceae) Phyllanthus (Euphorbiaceae) Pipturus (Urticaceae) Plumbago auriculata (Plumbaginaceae) Podolepis (Asteraceae) Poinsettia (Euphorbiaceae) Pollia crispata (Commelinaceae) Pomatocalpa macphersonii (Orchidaceae) Pongamia pinnata (Leguminosae) Prosopis juliflora (Leguminosae) Psidium guajava (Myrtaceae) Pultenaea (Leguminosae) Rapistrum (Brassicaceae) Rhizophora apiculata (Rhizophoraceae) Rotala densiflora (Lythraceae) Rubus (Rosaceae) Ruellia tuberose (Acanthaceae) Santiria laevigata (Burseraceae) Sapium (Euphorbiaceae) Sarcochilus moorei (Orchidaceae) Sechium edule (Cucurbitaceae) Senna siamea (Leguminosae) Sindora (Leguminosae) Smithia (Leguminosae) Solanum (Solanaceae) Sorghum vulgare (Poaceae) spp (Asteraceae) spp (Leguminosae) Sterculia (Sterculiaceae) Swainsona (Leguminosae) Symplocos (Symplocaceae) Syzygium (Myrtaceae) Tamarindus indica (Leguminosae) Taxillus (Loranthaceae) Tectona grandis (Lamiaceae) Telopea (Proteaceae) Tephrosia candida (Leguminosae) Terminalia (Combretaceae) Thunbergia laurifolia (Acanthaceae) Toona (Meliaceae) Torenia fournieri (Scrophulariacea) Tridax (Asteraceae) Tridax procumbens (Asteraceae) Trigonella corniculata (Leguminosae) Typha angustifolia (Typhaceae) Vitex (Lamiaceae) Wahlenbergia (Campanulaceae) Woodfordia floribunda (Lythraceae) Wrightia tinctoria (Apocynaceae) Xanthophyllum (Polygalaceae) Xanthorrhoea (Xanthorrhoeacea) Xerospermum intermedium (Sapindaceae) Zea mays (Poaceae) Zingiber officinale (Zingiberaceae)


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Appendix 2. Stingless bee taxon name followed by current combination or synonymy aliceae, Homotrigona aliceae (Cockerell, 1929) ambusta, see thoracica anamitica, Homotrigona anamitica (Friese, 1909) angophorae, see carbonaria apicalis, Tetrigona apicalis (Smith, 1857) arcifera, Lepidotrigona arcifera (Cockerell, 1929) atomella, see fuscobalteata atricornis, Platytrigona atricornis (Smith, 1865) atripes, Tetragonilla atripes (Smith, 1857) australis, Austroplebeia australis (Friese, 1898) bakeri, Heterotrigona bakeri (Cockerell, 1919) basimaculata, incertae setae bengalensis, Tetragonula bengalensis (Cameron, 1897) binghami, Tetrigona binghami (Schwarz, 1937) biroi, Tetragonula biroi (Friese, 1898) bismarkiana, nomen nudum, see biroi borneensis, see lacteifasciata breviceps, see itama brevis, nomen nudum, see fuscobalteata busara, see canifrons cacciae, Lisotrigona cacciae (Nurse, 1907) cambodiensis, see collina canifrons, Lophotrigona canifrons (Smith, 1857) carbonaria, Tetragonula carbonaria (Smith, 1854) carpenteri, Lisotrigona carpenteri Engel, 2000 cassiae, Austroplebeia cassiae (Cockerell, 1910) castanea, see fimbriata cincta, Austroplebeia cincta (Mocsáry in Friese, 1898) clypearis, Tetragonula clypearis (Friese, 1909) cockerelli, Austroplebeia cockerelli (Rayment, 1930) collina, Tetragonilla collina (Smith, 1857) confusella, see geissleri dapitanensis, Tetragonula dapitanensis (Cockerell, 1925) darek, nomen nudum, see minangkabau davenporti, Tetragonula davenporti (Franck, 2004) devicta, Tetragonula devicta (Cockerell, 1921) doipaensis, Lepidotrigona doipaensis (Schwarz, 1939) drescheri, Tetragonula drescheri (Schwarz, 1939) dulitae, see haematoptera erythrogastra, Heterotrigona erythrogastra (Cameron, 1902) erythrostoma, see fuscobalteata essingtoni, Austroplebeia essingtoni (Cockerell, 1905) ferrea, see lutea fimbriata, Homotrigona fimbriata (Smith, 1857) flavibasis, Lepidotrigona flavibasis (Cockerell, 1929) flavistigma, see fimbriata flaviventris, Platytrigona flaviventris (Friese, 1909) fulvomarginata, see terminata fulvopilosella, see nitidiventris furva, Lisotrigona furva Engel, 2000 fuscibasis, Tetragonilla fuscibasis (Cockerell, 1920) fuscobalteata, Tetragonula fuscobalteata (Cameron, 1908) geissleri, Tetragonula geissleri (Cockerell, 1918) genalis, Papuatrigona genalis (Friese, 1909) gressitti, Tetragonula gressitti (Sakagami, 1978) haematoptera, Odontotrigona haematoptera (Cockerell, 1919)



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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. hemileuca, see apicalis hirashimai, Tetragonula hirashimai (Sakagami, 1978) hobbyi, Platytrigona hobbyi (Schwarz, 1937) hockingsi, Tetragonula hockingsi (Cockerell, 1929) hoosana, see hoozana hoozana, Lepidotrigona hoozana (Strand, 1913) incisa, Geniotrigona incisa (Sakagami & Inoue, 1989) irianensis, nomen nudum iridipennis, Tetragonula iridipennis (Smith, 1854) itama, Heterotrigona itama (Cockerell, 1918) javanica, Lepidotrigona javanica (Gribodo, 1891) keyensis, Platytrigona keyensis (Friese, 1901) khasiana, nomen nudum klossi, see pendleburyi kusutkana, see aliceae lacteifasciata, Geniotrigona lacteifasciata (Cameron, 1902) laeviceps, Tetragonula laeviceps (Smith, 1857) lamingtonia, Platytrigona lamingtonia (Cockerell, 1929) latebalteata, Lepidotrigona latebalteata (Cameron, 1902) latigenalis, see reepeni latipes, Lepidotrigona latipes (Friese, 1900) lieftincki, Sundatrigona lieftincki (Sakagami & Inoue, 1987) lutea, Homotrigona lutea (Bingham, 1897) luteiventris, see erythrogastra matsumurai, see moorei melanocephala, Tetragonula melanocephala (Gribodo, 1893) melanoleuca, Tetrigona melanoleuca (Cockerell, 1929) melanotricha, see anamitica melina, Tetragonula melina (Gribodo, 1893) mellipes, Tetragonula mellipes (Friese, 1898) minangkabau, Tetragonula minangkabau (Sakagami & Inoue, 1985) minor, Tetragonula minor (Sakagami, 1978) mohandasi, Lisotrigona mohandasi Jobiraj & Narendran, 2004 moorei, Sundatrigona moorei (Schwarz, 1937) nigribasis, nomen nudum, see Lepidotrigona nitidiventris, Lepidotrigona nitidiventris (Smith, 1857) okazawai, nomen nudum ornata, Austroplebeia ornata (Rayment, 1932) pagdeni, Tetragonula pagdeni (Schwarz, 1939) pagdeniformis, Tetragonula pagdeniformis (Sakagami, 1978) palavanica, Lepidotrigona palavanica (Cockerell, 1915) pallidicincta, incertae setae pallidistigma, see fuscobalteata penangensis, Tetragonula penangensis (Cockerell, 1919) pendleburyi, Pariotrigona pendleburyi (Schwarz, 1939) peninsularis, Tetrigona peninsularis (Cockerell, 1927) percincta, Austroplebeia percinta (Cockerell, 1929) pfeifferi, see fuscobalteata planifrons, Platytrigona planifrons (Smith, 1865) praeterita, Tetragonula praeterita (Walker, 1860) pulla, nomen nudum pygmaea, see fuscobalteata reepeni, Tetragonula reepeni (Friese, 1918) rufibasalis, Tetragonilla rufibasalis (Cockerell, 1918) ruficornis, Tetragonula ruficornis (Smith in Horne, 1870) sandacana, see erythrogastra sapiens, Tetragonula sapiens (Cockerell, 1911) sarawakensis, Tetragonula sarawakensis (Schwarz, 1937) scintillans, see cacciae


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TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. sericea, see apicalis sirindhornae, Tetragonula sirindhornae (Michener & Boongird, 2004) smithii, see ruficornis symei, Austroplebeia symei (Rayment, 1932) terminata, Lepidotrigona terminata (Smith, 1878) terrestris, nomen nudum testaceinerva, see melanocephala testaceitarsis, 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 fimbriata vidua, Tetrigona vidua (Lepeletier de Saint Fargeau, 1836) wallacei, nomen nudum, see sapiens websteri, Austroplebeia websteri (Rayment, 1932) wybenica, see clypearis zucchii, 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, Max Beatson) ANIC Australia, Australian Capital Territory, Canberra City, CSIRO, Australian National 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 Department DEI 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 Resources MNHN 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 Museum RMNH Netherlands, Leiden, Nationaal Natuurhistorische Museum (“Naturalis”) (Cornelius van Achterberg) SEHU Japan, Sapporo, Hokkaido University Museum, Systematic entomology (Masahiro Ohara) SEMC USA, Kansas, Lawrence, University of Kansas, Snow Entomological Museum



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SFS SMNS USNM ZMHB ZRC

Shôichi F. Sakagami’s collection (see SEHU) Germany, Stuttgart, Staatliches Museum für Naturkunde USA, Washington D.C., National Museum of Natural History, (formerly, United States National Museum) (David Furth, Brian Harris) Germany, Berlin, Museum für Naturkunde der Humboldt-Universität (Frank Koch) Singapore, National University of Singapore, Raffles Museum of Biodiversity Research, Zoological Reference Collection (Lua Hui Kheng)

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