Rasmussen2008 7750 Catalog

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

ZOOTAXA

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

CLAUS RASMUSSEN

Magnolia PressAuckland, New Zealand

1935


RASMUSSEN2 · Zootaxa 1935 © 2008 Magnolia Press

Claus RasmussenCatalog 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/

© 2008 Magnolia Press

All rights reserved.

No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any

means, without prior written permission from the publisher, to whom all requests to reproduce copyright

material should be directed in writing.

This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose

other than private research use.

ISSN 1175-5326 (Print edition)

ISSN 1175-5334 (Online edition)


Accepted by E. Almeida: 16 Oct 2008; published: 17 Nov. 2008 3

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2008 · Magnolia Press

Zootaxa 1935: 1–80 (2008) www.mapress.com/zootaxa/

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

CLAUS RASMUSSENDepartment of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, USA. E-mail: [email protected]

Table of contents

Abstract ............................................................................................................................................................................... 4Catalog format ..................................................................................................................................................................... 4Taxonomy and classification ............................................................................................................................................... 5Distribution and biology ..................................................................................................................................................... 6Acknowledgments ............................................................................................................................................................... 7Catalog of the Indo-Malayan/Australasian stingless bees .................................................................................................. 8

Austroplebeia ............................................................................................................................................................... 8Geniotrigona .............................................................................................................................................................. 11Heterotrigona ............................................................................................................................................................. 13Homotrigona .............................................................................................................................................................. 15Lepidotrigona ............................................................................................................................................................. 17Lisotrigona ................................................................................................................................................................. 21Lophotrigona .............................................................................................................................................................. 23Odontotrigona ............................................................................................................................................................ 24Papuatrigona ............................................................................................................................................................. 24Pariotrigona ............................................................................................................................................................... 25Platytrigona ............................................................................................................................................................... 26Sundatrigona .............................................................................................................................................................. 27Tetragonilla ................................................................................................................................................................ 28Tetragonula ................................................................................................................................................................ 31Tetrigona .................................................................................................................................................................... 50Meliponini indet ......................................................................................................................................................... 53Incertae sedis ............................................................................................................................................................ 53Nomina nuda .............................................................................................................................................................. 54

Appendix 1. Plants visited by stingless bees ..................................................................................................................... 55Appendix 2. Stingless bee taxon names ............................................................................................................................ 59Appendix 3. ...................................................................................................................................................................... 61References ......................................................................................................................................................................... 62



Abstract

This catalog of the published literature on stingless bees from the Indo-Malayan/Australasian region was compiled from520 references and deals with 129 proposed names for species-group taxa of which 89 are considered currently valid.Information is summarized on organisms associated with stingless bee colonies as well as the 225 plants from which thebees have been reported to forage. Lectotypes are designated for Trigona ornata described by Rayment and the followingspecies described by Friese: Trigona anamitica, T. australis, T. borneënsis, T. flaviventris, T. keyensis, T. luteiventris, T.pygmaea, T. reepeni, T. sericea, and T. versicolor.

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 andsubtropical parts of the world (Heard 1999a). They form large perennial colonies with complex social organi-zation (Michener 2007a), have a diverse nesting biology ranging from underground to exposed colonies(Schwarz 1948; Wille & Michener 1973; Camargo & Pedro 2003b; Roubik 2006; Rasmussen & Camargo2008), and are abundantly encountered within their native range (Roubik 2006). In addition to a well-corrobo-rated phylogeny, an up-to-date, comprehensive taxonomic and biological catalog is fundamental to any com-parative evolutionary, ecological, and behavioral research on any group of organisms. A catalog provides anindex to previous studies in taxonomy, behavioral research, and pollination ecology, thus consolidating theexisting knowledge in an accessible format. The extensive Neotropical fauna was recently cataloged byCamargo & Pedro (2007: 391 spp.) and a systematic revision of the Afrotropical stingless bees included anearly complete bibliography (Eardley 2004: 26 spp. incl. Madagascar). The present catalog treats the remain-ing Indo-Malayan/Australasian stingless bees (129 proposed species-group names, 89 currently acceptedvalid species) and completes efforts to catalog the global stingless bee fauna. Together with recent molecularphylogenies of the stingless bees (Rasmussen & Cameron 2007; Rasmussen 2008), it is expected that thesetools will spark renewed scientific interest in this diverse group of bees.

Catalog format

The catalog is sorted alphabetically by genus-group and currently valid species-group names; subspecies arenot recognized in the catalog. Genus-group headings are in boldface type. Following the accepted species-group names are listed synonyms, including validly proposed junior synonyms and nomina nuda. Referencesare sorted in chronological order and include citation, page numbers from publications, type-specimen infor-mation from both type-specimen labels, and the original publication where the species was first described, keywords, distribution, and floral records. Incertae sedis and nomina nuda are listed at the end of the catalog,except when the identity is known for a nomen nudum. Page numbers are not listed for brief works or if theparticular species is treated throughout the publication. Publication dates were determined by the publishersimprint date on the wrapper of the fascicle, when available, or information contained within the issue. Whenthe imprint year of the issue differed from the stated year of the volume, the latter are placed in parentheses.Asterisks (*) following page numbers indicate where the species discussion is located in major works. Keywords in parenthesis after a reference indicate the kind of information provided and associated organismsrecorded. Misidentifications are corrected in the key word section whenever necessary or known, and typo-graphical errors in spelling of taxa names in publications are noted followed by an exclamation mark. Distri-butions are listed alphabetically with country names in upper case type and type localities indicated as in theoriginal publication. Distributional records are not listed comprehensively, but rather are limited to records


Zootaxa 1935 © 2008 Magnolia Press · 5INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES

cited in either taxonomic works or faunal inventories. Spellings of floral records were corrected to conformwith the International Plant Name Index (www.ipni.org). Appendix 1 lists all plant species (including family)upon which stingless bees are known to forage in the region. Plant synonymies were not updated, but ratherleft as cited in the original publication.

Stingless bee type specimens are encountered in museums worldwide, as indicated in the catalog. Acro-nyms for institutions are abbreviated following Evenhuis (2007) and Tadauchi et al. (1998) for Sakagami-related collections (Appendix 3). Curators who assisted with loans and important information on types arealso listed following each institution. Of particular importance are the collections in the Natural HistoryMuseum (London), with 52 primary types, Museum für Naturkunde der Humboldt-Universität (Berlin), with15, and National Museum of Natural History (Washington, D.C.), with 10. As for the most prolific describersof new taxa, Theodore D.A. Cockerell described 38 new taxa, Heinrich Friese 17, Frederick Smith 14, HerbertF. Schwarz and Peter Cameron 11 each, and Shôichi F. Sakagami 10. The whereabouts of three Sakagami pri-mary types remain unknown (i.e., Trigona gressitti, T. pagdeniformis, and T. lieftincki) as these have not beenfound in the collections indicated by the late Sakagami nor in his private collection (Tadauchi et al. 1998). Itmay be that the types are in those collections by Tadauchi et al. (1998), but that they are not labeled as types.The type of Trigona kusutkana Dover is missing from the Raffles Museum of Biodiversity Research where itwas originally deposited. All of the types of Smith have been accounted for, although as stressed by Baker(1993), several of the specimens labeled as types in the Natural History Museum (London) may not be truetypes and putative type specimens from London needs to be examined and compared with specimens fromUniversity Museum of Natural History (Oxford) when in doubt. A number of species described before thestandardization of taxonomy with the International Code of Zoological Nomenclature (latest edition is 1999)are represented by syntypes (multiple primary types). In several cases a syntype series consists of multiplebiological and morphological species. Lectotypes are here designated for several species proposed by Frieseas the identity of his true types is often confusing (see Rasmussen & Ascher 2008).

Literature has been updated until mid-2007. References cited include those found in Zoological Record(Zoological Society of London 1864–present) and all references cited within these records, totaling 520papers. A number of popular travel accounts are listed in the nearly complete stingless bees bibliography bySchwarz (1948). These were all consulted, but only included if they provided either the scientific name orsome description of the natural history of the bees, which was infrequently the case.

Taxonomy and classification

I have followed the classification proposed by Moure (1961) and Rasmussen & Cameron (2007) in not treat-ing some of the Old World stingless bees as subgenera of Trigona Jurine (Table 1). The phylogenetic groupsof Rasmussen & Cameron (2007) are fully congruent with the systematic arrangement proposed by Moure(1961). Therefore, all supraspecific groups are treated at the generic level as in the New World stingless beecatalog (Camargo & Pedro 2007). Other studies propose a simpler classification to avoid a profusion of for-mal names, which is equally valid when based on monophyletic lineages. This dual approach to classifyinggenera (chiefly Moure 1961; Michener 1990, 2007a), however, complicates literature searches with specifictaxa appearing under several different generic names. The genus-group name or synonymy is listed for eachtaxon in Appendix 2.

Most synonymies here are based on the published examination of types by Schwarz (1937; 1939a) andMoure (1961). However, a number of species were only tentatively synonymized by them, and re-examinationof primary types was necessary for confirming and establishing their synonymies. In those cases, I indicatedthe types I examined. Just like the inconsistent usage of stingless bee generic combinations, some taxa havebeen treated as subspecies (e.g., Schwarz 1939a; Sakagami et al. 1990), species (e.g., Klakasikorn et al.



2005), or synonyms. I have therefore refrained from stating whether I provide a new combination (comb. nov.)or a new status (stat. nov.), as all taxa have been given full species status or presented in the current combina-tion in the references provided. I did, however, recognize all previously proposed subspecies and varieties(mainly from the works of Schwarz) at the specific level when considered a valid taxon. This also reflects thefact that about 80% of the New World taxa originally described by Schwarz as subspecies in light of addi-tional biological information are now regarded fully valid species (Camargo & Pedro 2007).

Identification of stingless bees from the Indo-Malay/Australasian region is best done with specializedkeys, although the fauna has not yet undergone the taxonomic scrutiny comparable to that of many NewWorld taxa (e.g., Camargo & Moure 1994; Camargo & Moure 1996; Pedro & Camargo 2003b). WhileMichener (2007a) included genus-group keys, the synonymy of a larger part of the regional fauna under thepolyphyletic subgenus Heterotrigona, sensu Michener, makes that key less useful. The only other genus-group key currently available is that of Moure (1961). However, as the fauna is not very extensive, species-group keys may be preferred (Schwarz 1939a; Sakagami et al. 1990), although additional literature may needto be consulted for poorly characterized genera like Tetragonula. For these genera, regional keys can be useful(Sakagami 1978; Dollin et al. 1997). No workable key exists for Austroplebeia (although see Rayment 1935:735), Platytrigona (although see Michener 1990: 127), and some of the varieties here considered as valid spe-cies.

Distribution and biology

The distribution of stingless bees in the Indo-Malay/Australasian region stretches from India to the SolomonIslands and from China (Yunnan, Hainan, Taiwan: Wu 2000) to Australia (New South Wales). Schwarz (1937)suggested that the greater abundance of stingless bees in Thailand and Malaysia, including all of Borneo, isdue to the abundance of resin-secreting trees (Dipterocarpaceae) and humid tropical climate, although mostlikely an array of factors are responsible for this concentration of species.

Natural history observations summarized from the present catalog are as follow:Colony pests: Achroea grisella (Lepidoptera: Tineidae), Ceria cf. ornatifrons (Diptera: Syrphidae) (larvae

feeding from provision), and Protaetia mandarinea (Coleoptera: Scarabaeidae). Insects and other organisms inhabiting nests or inquilines: Brachypeplus auritus, B. basalis, B. meyricki,

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 macrocer-cus, D. paradiseus (Dicruridae: Passeriformes), Dorniphora trigonae (Diptera: Phoridae) (of brood?), Helarc-tos malayanus (Carnivora: Ursidae), Hermetia illucens (Diptera: Stratiomyidae) (of brood), Meropsleschenaulti (Coraciiformes: Meropidae), Nephila maculata (Araneae: Nephilidae), Oecophylla smaragdina,O. sp. (Hymenoptera: Formicidae), Pahabengkakia piliceps (Hemiptera: Reduviidae), Phoridae (Diptera),small birds, Solenopsis geminata (Hymenoptera: Formicidae), Syrphidae (Diptera), and Thomisus spectabilis(Araneae: Thomisidae). A bee was also reported “clinging to the leg of a large green cicada”, Dundubia spic-ulata (Hemiptera: Cicadidae), although the details of that interaction was not provided.

Nest sites: aerial Crematogaster nests (Hymenoptera: Formicidae), bamboo stems and interspaces, beetleburrows, cavities among roots, coco nuts, cracks, crevices, holes in limestone, hollow pillars, hollow trunks(including Durio zibethinus, Bombacaceae; Eugenia aromatica, Myrtaceae; Ficus, Moraceae; Nephelium lap-paceum, Sapindaceae; Shorea laevis, Dipterocarpaceae), house posts, a human skull, and old tree stumps.



TABLE 1. Different classificatory schemes used in the literature.

Lastly, floral records are frequently reported and show that the stingless bees are truly polylectic with 69families and 179 genera of plants recorded as hosts in the Indo-Malayan/Australasian region. This includesplants visited for nectar without performing pollination. The diversity of plants includes both cycads (Cyca-daceae) and angiosperms, including Monocot (e.g., Arecaceae, Orchidaceae, and Poaceae) and Dicot families.

Acknowledgments

I am very grateful to the many curators who replied to my queries about their type holdings. In particular toFrank Koch for hosting me during a visit to the Museum für Naturkunde, David Furth and Brian Harris forhosting me multiple times at National Museum of Natural History, and Holger H. Dathe and Stephan Blankfor hosting a brief visit to the DEI. David Notton kindly provided much information and loan of types fromthe Natural History Museum. J.M.F. Camargo gave me unlimited access to his comprehensive stingless beelibrary and collection which was helpful in forming the foundation for the present catalog, Camargo and Sil-via Pedro also provided useful comments on this project. John S. Ascher, Sydney A. Cameron, Jim B. Whit-field, Chris Dietrich, Nils Cordes, and two anonymous reviewers made helpful comments on the manuscriptand Ying Wang helped with translation of Chinese. Visits to institutions were supported by a Francis M. andHarlie M. Clark Research Support Grant and a National Science Foundation grant (DEB 0446325) to S. A.Cameron who also encouraged this study.

Genus: This work and Moure (1961) Type species Genus: Michener (2007)

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)



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); Ray-ment 1935: 735 (key to species); Schwarz 1948: 117; Moure 1961: 196 (morphology, systematic posi-tion); 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); Wag-ner & 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); Michener1990: 134, 137 (illustration, morphology, systematic position); Cardale 1993: 318 (distribution, floralrecord, list); AUSTRALIA, NE coastal; Queensland; New South Wales; SE coastal; Murray-Darlingbasin; Floral record: Alphitonia; Angophora; Callistemon; Eucalyptus; Melaleuca; Rapistrum; Swain-sona; Wahlenbergia; Heard & Exley 1994: 97 (comparative note); Drumond et al. 1995: 43 (citation,comparative behavior); Heard 1996: (bee-keeping, nest); Dollin 1996a: (characteristics, nest); Dollin1996b: (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); Crane1999: 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); Serrão2001: (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 etal. 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 (dis-tribution); Cockerell 1929d: 299, 301 (common name (kootchar), distribution, nest (tree hollows), para-site? (Ceria cf. ornatifrons: Syrphidae: Diptera)); AUSTRALIA, Brisbane; Cockerell 1930b: 150(common name (kootchar), comparative note, floral record, nest); Floral record: Cassia; Eucalyptus;



Rayment 1930: 59 (comparative note); Rayment 1932a: 185 (common name (koochee)); Rayment1932d: (honey, pollen, wax); Rayment 1932b: (nest); Rayment 1932c: (behavioral comparison); Ray-ment 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, phys-iology); Wille & Michener 1973: 10 (synonymy); Camargo 1988: 372 (distribution); AUSTRALIA,Queensland; Michener 1990: 96, 101, 105, 133, 134 (illustration, morphology, systematic position); Sak-agami et al. 1990: 131 (see cacciae); Michener & Roubik 1993: 259, 262 (comparative note, morphol-ogy); Cardale 1993: 318-319 (distribution, floral record, list); AUSTRALIA, NE coastal; Queensland;Brisbane; Caloundra; Mackay; Floral record: Cassia; Eucalyptus; Cunha 1994: (systematics); Michener1997: 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 Wil-helmshafen)" (1 worker); Friese 1909("1908"): 354, 356 (distribution); INDONESIA, Irian Jaya/PAPUANEW GUINEA, Cyclopen Gebirge; Moaif; Moso; Sentani; Tawarin; Timena; Friese 1909: 272, 273 (cita-tion, distribution, key to species); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Cyclopen Gebirge;Moaif; Moso; Sentani: Tawarin; Timena; Cockerell 1910: 247 (misidentification, uncertain identity, pos-sibly 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 (probablymisidentification); INDONESIA (Irian Jaya)/PAPUA NEW GUINEA, Bewani Fluss; Cyclop Gebirge;Finschhafen; Friedrich Wilhelmshafen; Hussin; Jasa; Moaif; Moso; Njao; Sentani; Sepik-Bivak; Sim-bang; Stephansort; Tawarin; Timena; Zoutbron; Floral record: Eucalyptus "staunus"; Friese 1917: 9 (dis-tribution); AUSTRALIA, Kimberley (probable misidentification); Cockerell 1929e: 242 (distribution,taxonomy); PAPUA NEW GUINEA, Mt. Lamington; Schwarz 1932: 245; Rayment 1932f: 106 (taxon-omy); Rayment 1935: 735 (key to species); Schwarz 1948: 117; Moure 1961: 197 (morphology, system-atic 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; Sim-bag, Stephansort; Michener 1990: 97, 110, 133, 134 (systematic position); Michener & Roubik 1993:262 (comparative note, morphology); Cardale 1993: 319 (distribution (identity uncertain)); AUSTRA-LIA, 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. /



INS. COLL.", paratype (ANIC (1)) (illustration, taxonomy); Type locality: AUSTRALIA "Borroloola,North Australia (Gerald Hill, 25th September, 1911)" (worker); Rayment 1932c: (behavioral compari-son); 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; Dollin1997: (characteristics); Klumpp 2007: 5.

Austroplebeia essingtoni (Cockerell, 1905)

Trigona essingtoni Cockerell 1905: 220-221: Holotype (BMNH 17b.1138, worker): examined (head miss-ing), "Syntype" (blue border), "B.M. TYPE / HYM. / 14B.1138", "Pt. Essing / ton / 42-", "Trigona / ess-ingtoni 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); AUSTRA-LIA, Port Essington; Rayment 1930: 59 (comparative note); Rayment 1932f: 106 (taxonomy); Rayment1935: 735 (key to species); Rayment 1953: (distribution); Moure 1961: 196-197 (morphology, system-atic position, variation); Michener 1965: 230 (citation); Evans & Matthews 1973: (predator (Bembix fla-vipes: 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 posi-tion); Cardale 1993: 319-320 (distribution, list); AUSTRALIA, N coastal; NW coastal; Northern Terri-tory; Western Australia; Welch 1995: (rock art); Dollin 1997: (characteristics); Crane 1999: 89; Klumpp2007: 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 / cock-erelli / ornata", "MUS. VIC. / ENTO 2008-8 L"; Paralectotypes (MVMA (3)) (taxonomy (as manuscriptname)); Type locality: AUSTRALIA "Cape York, Queensland (Donald Thomson)" (workers); Rayment1935: 735 (key to species, taxonomy (described as new species again)); Moure 1961: 197 (possible syn-onymy); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution, list); AUSTRALIA, NEcoastal; Queensland; Dollin 1997: (characteristics); Klumpp 2007: 5.

Austroplebeia percinta (Cockerell, 1929)

Trigona cincta subsp. percinta nov. Cockerell 1929e: 242: Syntypes (ZMHB, 4 workers): examined, labeledby Friese as Trigona cincta? (comparative note (corrected identification of Cockerell 1910)); Type local-ity: AUSTRALIA "Hermannsberg, Finke River" (unknown); Cockerell 1930b: 150 (taxonomy); Ray-ment 1932f: 106 (taxonomy); Rayment 1935: 735 (key to species); Schwarz 1948: 92 (citation); Moure1961: 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



/ NT. QUEENS", "AUST. NAT. / INS. COLL." (taxonomy (as manuscript name)); Type locality: AUS-TRALIA "North Queensland" (worker); Rayment 1935: 735, 737-738 (key to species, taxonomy(described as new species again)); MacPherson & Rupp 1936: (pollination); Floral record: Dendrobiumspeciosum; 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, taxon-omy); 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: (sys-tematics); 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 / websteriRaym. / 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 (distribu-tion); 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 & Dollin2000: (bee-keeping); Maynard et al. 2004: 319 (citation); Roubik 2006: 126, 128 (citation, nest); Corto-passi-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, nativecollector, ded. F. Dupont" (3 workers, holotype, paratypes); "Todyamboe, 900m, C. Celebes" (3 workers);"Wuasa, Kab. Poso, Sulteng, Sulawesi" (9 workers); "Lorei Lindu Nat. Park, C. Sulawesi" (3 workers);Tadauchi et al. 1998: 242-243 (type depositories); Rasmussen & Cameron 2007: (systematics).

Geniotrigona lacteifasciata (Cameron, 1902)

Trigona borneënsis Friese 1933a: 46: Lectotype (ZMHB, worker): here designated, "Borneo / Sanggau / 24-7-32", "Trigona / borneensis / 1925 Friese det. Fr."; paralectotypes (DEI (4 (2 on 1 pin), ZMHB (2)); pos-sible 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 (synon-ymy); 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-



racica from the island of Borneo (taxonomy); Type locality: MALAYSIA "Borneo" (unknown); Cam-eron 1908: 193 (key to species); Cockerell 1919c: 78 (comparative note); Cockerell 1923: 242 (key tospecies); 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); Typelocality: MALAYSIA/SINGAPORE "Sandakan and Singapore" (unknown); Cockerell 1919c: 78 (tax-onomy (described in previous paper)); SINGAPORE; Cockerell 1920b: 228 (distribution); MALAYSIA,Sandakan; Cockerell 1929b: 139 (distribution); MALAYSIA, Kuala Lumpur; Cockerell 1930a: 157 (dis-tribution, nest (hollow tree), variation); CAMBODIA, Koh Po; Pattani; Cockerell 1934: 62, fig. 7 (com-parative 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 (synon-ymy).

Trigona thoracica Smith 1857: 50: Type (BMNH 17b.1181) (taxonomy); Type locality: SINGAPORE "Sin-gapore" (worker); Smith 1871: 395 (distribution); SINGAPORE; Dalla Torre 1896: 584 (citation); Bing-ham 1897: 561, 564 (distribution, key to species, taxonomy); MYANMAR, Tenasserim; SINGAPORE;Bingham 1905: 58 (citation, distribution); Friese 1909("1908"): 358 (comparative note); Cockerell1919c: 79 (distribution); Cockerell 1923: 240 (key to species); Dover 1929: 62 (citation, distribution);MALAYSIA, Kuala Tahan; Pahang; SINGAPORE; Friese 1933a: 46 (comparative note); Schwarz 1937:317, 318; Schwarz 1939a: 85, 90, 92, 104-106* (distribution, key to species, taxonomy); Floral record:Citrus limon; Tephrosia candida; Schwarz 1948: 83, 104, 114, 118; Moure 1961: 212, 213 (systematicposition); Yoshikawa et al. 1969: 172, 173 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang;Tapah; Touaran; Sakagami 1975: 56* (distribution); SINGAPORE; THAILAND; Wille 1979b: 247(morphology); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floralrecord: Xerospermum intermedium; Sakagami et al. 1983b: 2, 4, 13-15, 16, 20 (honey, nest (in Shorealaevis: Dipterocarpaceae)); Sakagami et al. 1985: (distribution, key to species); Baker & Delfinado-Baker 1985: 231 (inquiline (Neocypholaelaps phooni: Ameroseiidae: Acari)); Camargo 1988: 370 (dis-tribution, taxonomy); SINGAPORE; Sakagami & Inoue 1989: (morphology); Michener 1990: 105, 126(illustration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species);INDONESIA, Sumatra; MALAYSIA, Borneo; MYANMAR; THAILAND; Salmah et al. 1990: (distribu-tion, nest (hollow tree trunks; holes in limestone)); INDONESIA, Sumatera Barat; Inoue et al. 1990b:(population); Inoue 1990: 582 (nest (huge tree cavities), population); Johnson & Appanah 1990: (preda-tor (Oecophylla: Formicidae: Hymenoptera)); Michener & Roubik 1993: 260 (comparative note, mor-phology); Cunha 1994: (systematics); Yamane et al. 1995: 294 (synonymy); Roubik 1996: 61(distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433 (pollination); Floral record:Santiria laevigata; Michener 1997: 25 (type species); Nagamitsu & Inoue 1998: (foraging, morphol-ogy); Heard & Dollin 1998: (citation); Heard & Dollin 1998: 11 (pollination); Floral record: Averrhoacarambola; Liow 1999: 106, 127 (distribution); Nagamitsu et al. 1999: (floral records (not by species),foraging); Heard 1999a: 191, 192, 195 (citation); Michener 2000: 801-802; Chinajariyawong & Sai-boon 2000: (distribution, nest (as thorasica!)); THAILAND, Songkhla; Liow et al. 2001: 184; Nag-amitsu & Inoue 2002: 305 (foraging); Eltz et al. 2003: 309 (citation); Eltz 2004: 322 (abundance);Nagamitsu & Inoue 2005: 76, 81, 83, 86, plate 9 (distribution, foraging); Klakasikorn et al. 2005: 3, 6(floral record); Floral record: Dipterocarpus alatus; Hopea; Roubik 2006: 129, 130, 131 (nest); Slaa et



al. 2006: 297 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kancha-naburi, 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" (comparativenotes, key to species); Type locality: MALAYSIA "Penang Island (Baker 9068)" (worker); Cockerell1930a: 157 (distribution, floral record, variation); CAMBODIA, Tung Nui; Satul; Floral record: Melas-toma malabaricum; Schwarz 1937: 323, 324; Schwarz 1939a: 96 (synonymy); Moure 1961: 203 (mor-phology, 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?) (com-parative notes, taxonomy); Type locality: MALAYSIA "Sarawak (R. Shelford)" (worker); Cameron1908: 193, 194 (key to species); Cockerell 1920a: 115 (comparative note); Cockerell 1923: 242 (key tospecies); Schwarz 1937: 291, 319-320*, 323, 326, 327, plate 3, 6 (key to species, taxonomy); Floralrecord: Melastoma; Schwarz 1939a: 84, 85, 86, 90, 96, 100-101* (distribution, key to species, taxon-omy); Schwarz 1948: 118; Moure 1961: 203 (systematic position); Baltazar 1966: 408 (distribution);PHILIPPINES, Palawan; Jander & Jander 1970: 362, 363, 367 (orientation); Sakagami 1975: 55* (dis-tribution, variation); MALAYSIA; Wille 1979b: 265 (comparative note); Appanah 1982: (pollination);MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakag-ami et al. 1985: (distribution, key to species); Appanah et al. 1986: (foraging, nest); Floral record: Calo-phyllum; Elaeis; Elaeocarpus; Sindora; Xanthophyllum; Camargo 1988: 370 (distribution, taxonomy);MALAYSIA, Borneo, Sandakan; Sakagami & Inoue 1989: 617, 619 (comparative note); Michener1990: 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); BRU-NEI, Kuala Belalong; Bruijn & Sommeijer 1997: 44 (citation (as erythrogaster!)); Nagamitsu & Inoue1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Heard1999a: 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 spe-cies); 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);



Type locality: MALAYSIA "Sandakan, Borneo (Baker, 9593)" (worker); Cockerell 1920b: 228 (distri-bution); 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 bor-der), "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 spe-cies); Type locality: SINGAPORE "Singapore" (unknown); Cockerell 1919b: 241-242, 244 (taxonomy,variation); MALAYSIA, Penang; Cockerell 1919c: 78, 79 (comparative notes, key to species); Cockerell1919a: 121 (distribution, taxonomy, variation); INDONESIA, Pelaboean Ratoe, Java; SINGAPORE;Cockerell 1920a: 115 (comparative note); Cockerell 1925a: 492 (variation); MALAYSIA, Borneo, San-dakan; Cockerell 1927: 541 (distribution); MALAYSIA, Selangor, Pahang, near Genting, Sempat, Hillstream; 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, taxon-omy); Floral record: Mimosa pudica; Tephrosia candida; Sandhouse 1943: 558 (citation); Schwarz1948: 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); Appanah1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermumintermedium; Sakagami et al. 1983b: 2-4, 13, 19, 20 (nest); Sakagami et al. 1985: (distribution, key tospecies); 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 (Neocypholae-laps phooni: Ameroseiidae: Acari)); Atmowidjojo & Adisoemarto 1986: 227, 229 (pollination); Floralrecord: Indigofera; Sakagami & Inoue 1987: 625 (comparative note); Uji 1987: (pollination); INDONE-SIA, E Kalimantan; Floral record: Nephelium lappaceum; Camargo 1988: 370 (distribution, taxonomy);MALAYSIA, Kuala Lampur; SINGAPORE; Sakagami & Inoue 1989: 610, 613, 614, 616, 617 (mor-phology); Roubik 1989: 105, 133, 137, 156, 158; Michener 1990: 105, 120, 122, 125, 126, 127 (illustra-tion, 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, SumateraBarat; 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: (repro-duction); Roubik 1996: 61, 62 (distribution); BRUNEI, Kuala Belalong; Michener 1997: 29 (type spe-cies); Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 110, 127 (nest); Nagamitsu etal. 1999: (floral records (not by species), foraging); Heard 1999a: 196 (citation); Michener 2000: 801-802; 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);



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 etal. 2007: (foraging); Floral record: Averrhoa carambola; Cassia fistula; Michener 2007a: 826-827; Wat-tanachaiyingcharoen & 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 tospecies, taxonomy); Schwarz 1948: 118; Schwarz 1951: 63 (distribution); CAMBODIA, Angkor;Moure 1961: 201 (systematic position); Sakagami & Yoshikawa 1961: 434-435 (distribution, taxon-omy); THAILAND, Doi Inthanon; VIETNAM, Kompong Cham; Kamphangpet; Sakagami 1963: (mor-phology (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, paratypeworker); Schwarz 1939a: 104* (distribution, key to species, taxonomy); Moure 1961: 201 (systematicposition); 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."; para-lectotype (ZMHB (1)) (comparative note, taxonomy); Type locality: VIETNAM "Xom-Gom, Süd-Annam, 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); Sakagami1975: 52, 54 (distribution, variation); MALAYSIA, Borneo; VIETNAM; Camargo 1988: 371 (synon-ymy); Sakagami et al. 1990: 127 (synonymy).

Trigona melanotricha Cockerell 1918: 386, 387: Holotype (BMNH 17b.1095) (key to species, comparativenotes, 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 (synon-ymy); Moure 1961: 201 (systematic position); Sakagami 1975: 52 (synonymy); Camargo 1988: 370-371 (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-



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 "Sin-gapore" (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 spe-cies); Schwarz 1937: 283, 289, 304-307*, 320, 326, 327, plate 2, 4 (key to species, taxonomy); Schwarz1939a: 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 (compara-tive note); Sakagami 1963: (see aliceae); Sakagami 1975: 52-55* (distribution, morphology, variation);LAOS; MALAYSIA; SINGAPORE; THAILAND; VIETNAM; Sakagami et al. 1983b: 5 (nest); Sakag-ami et al. 1985: (distribution, key to species); Sakagami & Khoo 1987: 207 (comparative note); Cama-rgo 1988: 371 (synonymy); Roubik 1989: 133, 135, 417; Michener 1990: 105, 120, 125, 126 (systematicposition); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra;MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, SumateraBarat; Inoue et al. 1990b: (population); Roubik 1993: 541 (illustration, nest); Michener & Roubik 1993:260 (comparative note, morphology); Cunha 1994: (systematics); Roubik 1996: 61 (distribution); BRU-NEI, Kuala Belalong; Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: Santiria laevi-gata; 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; Rajit-parinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Green et al. 2001: 91 (micro-satellite); Eltz et al. 2002a: 30 (distribution, foraging, nest, population); MALAYSIA, Danum Valley;Sepilok; Deramakot; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh ForestReserve; Eltz et al. 2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baramarea; 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: Tec-tona grandis; Roubik 2006: 131 (nest); Jongjitvimol et al. 2006: (foraging); Jongjitvimol & Watta-nachaiyingcharoen 2006: (foraging); THAILAND, Phitsanulok; Floral record: Butea monosperma;Cassia bakeriana; Erythrina stricta; Fernandoa adenophylla; Ipomoea aquatica; Lagerstroemia calycu-lata; Lagerstroemia macrocarpa; Lagerstroemia tomentosa; Merremia vitifolia; Mimosa pigra; Mimosapudica; 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 (syn-onymy); 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)" (2workers); Schwarz 1937: 304, 306, 327, plate 4; Schwarz 1939a: 102, 103, 104 (synonymy); Schwarz1948: 118; Moure 1961: 201 (synonymy); Sakagami 1975: 52 (synonymy); Sakagami et al. 1990: 127(synonymy).



Homotrigona lutea (Bingham, 1897)

Trigona ferrea Cockerell 1929b: 139-140: Holotype (BMNH 17b.1117) (comparative note, taxonomy); Typelocality: 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 spe-cies, taxonomy; notice this is no longer a senior homonym of Trigona mocsaryi lutea Friese 1903, seeCamargo & Pedro 2007: 472); Type locality: MYANMAR "Northern Tenasserim; Karennee" (worker);Cockerell 1929b: 140 (comparative note); Moure 1961: 201 (systematic position); Wu 2000: 387, 390,plate (key to species, morphology).

Homotrigona indet. Roubik 2006: 130 (nest).

Lepidotrigona

Lepidotrigona arcifera (Cockerell, 1929)

Trigona arcifera Cockerell 1929c: 591-592: Holotype (BMNH 17b.1081). The ventralis species group (tax-onomy); Type locality: INDIA "Testa bridge, Himalayas, India, 10.1.97 (Sladen)" (worker); Schwarz1937: 295; Schwarz 1939a: 85, 88, 134, 135 (distribution, key to species, taxonomy); Nogueira-Neto1949: 121 (bee-keeping); Moure 1961: 199 (possible synonymy); Sakagami 1975: 67, 71 (synonymy);Sakagami et al. 1990: 126 (synonymy); Wu 2000: 387 (synonymy).

Lepidotrigona doipaensis (Schwarz, 1939)

Trigona (Lepidotrigona) ventralis variety doipaensis Schwarz 1939a: 85, 94, 136*: Holotype (USNM 53563,worker): examined, "Doi Pa / Mai Deng / Siam 750m / 12-29-32", "HughSmith / Coll", "Type No./ 53563/ U.S.N.M.", "Holotype", "Trigona ventralis / var. doipaensis / H.F. Schwarz". The ventralis species group(distribution, key to species, taxonomy); Type locality: THAILAND "SIAM.-Doi Pa, Mai Deng, 750meters, Dec. 29, 1932 (H. M. Smith)" (worker); Moure 1961: 199 (systematic position); Sakagami 1975:67, 69, 71 (distribution, variation); THAILAND, N; Sakagami & Yamane 1984: 35; Sakagami et al.1990: 126 (synonymy); Wu 2000: 387 (synonymy); Klakasikorn et al. 2005: 3, 6 (floral record); Floralrecord: Ficus religiosa; Homalium grandiflorum.

Lepidotrigona flavibasis (Cockerell, 1929)

Trigona flavibasis Cockerell 1929c: 592: Holotype (AMNH). The ventralis species group (taxonomy); Typelocality: THAILAND "Doi Sutep, Siam, Feb. 9 (Alice Mackie)" (worker); Schwarz 1939a: 85, 94, 134-135*, 136 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199 (systematicposition); Yoshikawa et al. 1969: 174 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang; Sakag-ami 1975: 67, 70, 71, 74 (distribution, variation); CAMBODIA; MALAYSIA; THAILAND; Sakagami& Yamane 1984: 38, 47; Sakagami et al. 1990: 126 (synonymy); Wu 2000: 387 (synonymy); Green etal. 2001: 91 (microsatellite); Chinh et al. 2005: 366, 368-371 (nest (as ventralis in part)); VIETNAM,Cucphuong National Forest, Red river delta; Chinh & Sommeijer 2005: (reproduction (as ventralis in



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 (varia-tion); Sakagami & Yamane 1984: 34-48; Sakagami et al. 1990: 126, 131 (distribution); Wu 2000: 387(synonymy); Chinh et al. 2005: 371.

Trigona ventralis hoozana Strand 1913b: 66: Syntypes (DEI, 9 workers). According to S. Blank (DEI)labeled "II. Hoozan Gormosa H. Sauter 10." (for February 1910); "Trigona ventralis Sm. Strand det.",some with "F. Sm." instead of "Sm.". Strand proposed the name conditionally (ICZN 11.5.1 and 15.1), butas it was proposed as a geograhical restricted (=subspecific) form before 1961, it is considered valid(45.6.1). The ventralis species group (taxonomy); Type locality: TAIWAN "Hoozan, Februar" ('a row' ofworkers); Sakagami et al. 1985: (key to species); Sakagami & Yamane 1987: (oviposition); Salmah etal. 1990: 155 (altitude); Bravo 1992: 876 (citation, altitude); Yamane et al. 1995: 294 (comparative note);Sung et al. 2004: (morphology); Nagamitsu & Inoue 2005: 76 (distribution, foraging); Chinh & Som-meijer 2005: 500 (citation); Sung et al. 2006c: (distribution, nest); Sung et al. 2006a; Sung et al. 2006b:167 (pollination); Floral record: Mangifera indica.

Lepidotrigona javanica (Gribodo, 1891)

Trigona javanica Gribodo 1891: 109: Holotype (MSNG, worker). The terminata species group (taxonomy);Type locality: INDONESIA "Giava" (=Java) (1 worker); Dalla Torre 1896: 579 (citation); Friese1914a: 13 (distribution); INDONESIA, Nusa Kambangan; Friese 1914b: 58 (nest); Cockerell 1929c: 591(comparative note); Schwarz 1937: 295, 296; Schwarz 1939a: 85, 90, 94, 140* (distribution, key to spe-cies, taxonomy); Moure 1961: 199 (systematic position); Wille & Michener 1973: 14, 24, 48 (nest);Sakagami 1975: 65 (comparative note).

Lepidotrigona latebalteata (Cameron, 1902)

Trigona latebalteata Cameron 1902: 130-131: Syntypes (BMNH 17b.1084, 2 workers). The terminata spe-cies group (taxonomy); Type locality: MALAYSIA "Kuching, Sarawak" (worker); Smith 1857: 50 (asventralis, in part); MALAYSIA, "Malacca (Mt. Ophir)"; Cameron 1908: 193 (key to species (as latibal-teata!)); Schwarz 1937: 282, 285, 288, 294, 295-297*, 326 (key to species, taxonomy); Schwarz 1939a:85, 94, 135, 137, 139-140* (distribution, key to species, taxonomy); Schwarz 1948: 118 (morphology);Moure 1961: 199 (systematic position); Sakagami 1975: 65 (comparative note).

Lepidotrigona latipes (Friese, 1900)

Trigona latipes Friese 1900: 384: Holotype (ZMHB, worker): examined, "India / Singapore / 1890", "Trig-ona / 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).



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); Schwarz1937: 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 nitidiventrisspecies 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, Pal-awan; Sakagami 1975: 64-65* (distribution, variation); MALAYSIA; THAILAND; Appanah 1982:(pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum interme-dium; Sakagami et al. 1985: (distribution, key to species); Sakagami & Yamane 1987: 25 (comparativenote); 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); Cunha1994: (systematics); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue1997: 434, 436 (pollination); Floral record: Santiria laevigata; Michener 1997: 32 (type species); Nag-amitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species),foraging); Michener 2000: 802; San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, PasohForest Reserve; Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Klakasikorn et al. 2005: 6(citation); Michener 2007a: 827; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparativenote).

Lepidotrigona palavanica (Cockerell, 1915)

Trigona palavanica Cockerell 1915: 2-3: Holotype (BMNH 17b.1120). The nitidiventris species group (com-parative notes, taxonomy); Type locality: PHILIPPINES "P. Princesa, Palawan (Baker coll 3839)"(worker); Cockerell 1919c: 77 (key to species); PHILIPPINES, Palawan, Puerto Princesa; Cockerell1920b: 229 (distribution); PHILIPPINES, Palawan; Schwarz 1939a: 85, 86, 133 (distribution, key to spe-cies, 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; nearNam 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); Cocker-ell 1929c: 591 (comparative note); Schwarz 1937: 282, 284, 288, 295, 296, 326; Schwarz 1939a: 85, 90,94, 95, 100, 127, 128, 133, 135, 136-139* (distribution, key to species, taxonomy); Schwarz 1948: 118;



Pagden 1957: 17 (comparative note); Sakagami 1960: 146 (distribution, variation); CAMBODIA, Snoul;Stung Treng; Moure 1961: 199 (systematic position); Wille 1961: 117, 121; Sakagami & Yoshikawa1961: 440 (distribution); LAOS, Thakhek; THAILAND, Mae Fack; Yoshikawa et al. 1969: 168, 169, 174(distribution, nest); MALAYSIA, George Town, Penang; Wille & Michener 1973: 14, 25, 49, 60 (nest);Sakagami 1975: 65-66*, 72, 73, 74 (distribution, variation); LAOS; THAILAND; VIETNAM; Sakag-ami 1978: 239 (comparative note); Sakagami et al. 1983b: 2, 15-19, 20 (nest); Sakagami & Yamane1984: 40, 47; Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts));Sakagami & Yamane 1987: 24, 25 (comparative note); Camargo 1988: 371 (distribution, taxonomy);INDONESIA, Java; MALAYSIA, Penang; Roubik 1989: 181; Michener 1990: 95, 105, 120, 122 (illus-tration, morphology, systematic position); Sakagami et al. 1990: (distribution, key to species); CAMBO-DIA; INDONESIA, Java; Sumatra; MALAYSIA; MYANMAR; THAILAND; Salmah et al. 1990:(distribution, nest (hollow tree trunks)); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population);Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Michener & Roubik1993: 260 (comparative note, morphology); Yong 1993: (biochemistry); Yamane et al. 1995: 294 (synon-ymy); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 434, 436(pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Liow1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener2000: 800; Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Wu 2000:386, 388-390 (key to species, morphology); Eltz et al. 2001b: 343, 345 (foraging); MALAYSIA, Borneo,Sabah; Green et al. 2001: 91 (microsatellite); Eltz et al. 2002a: 30, 31, 32 (distribution, foraging, nest,population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Floral record: Rhizophora apiculata;Klein et al. 2002: 1010 (floral record); Floral record: Coffea canephora; San & Osawa 2002: (popula-tion); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Kleinet al. 2003b: 957 (floral record); Floral record: Coffea canephora; Klein et al. 2003a: 841 (floral record);Floral record: Coffea canephora; Eltz 2004: 322 (abundance); Samejima et al. 2004: (population);MALAYSIA, Sarawak, Miri, Upper Baram area; Wattanachaiyingcharoen et al. 2004: (diversity, nest);Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Chinh et al. 2005: 368, 371 (citation);Klakasikorn et al. 2005: 3, 6 (floral record); Floral record: Ficus religiosa; Irvingia malayana; Slaa et al.2006: 297, 298 (citation); Tangmitcharoen et al. 2006: (pollination); Floral record: Tectona grandis;Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum;Rasmussen & Cameron 2007: (systematics); Michener 2007a: 824; Wattanachaiyingcharoen &Jongjitvimol 2007: 73 (comparative note).

Lepidotrigona trochanterica (Cockerell, 1920)

Trigona trochanterica Cockerell 1920a: 115: Holotype (BMNH 17b.1102) (taxonomy); Type locality:MALAYSIA "Sandakan, Borneo (Baker)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA,Sandakan; Schwarz 1937: 285, 287, 292-293*, 295, 296, 327 (key to species, taxonomy); Schwarz1939a: 85, 94, 133-134* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199(systematic position); Sakagami 1975: 65 (citation); Michener 1990: 122 (systematic position); Sakag-ami et al. 1990: (distribution, key to species, variation); INDONESIA, Sumatra; Salmah et al. 1990: (dis-tribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener 2000: 802;Nagamitsu & Inoue 2005: 76 (distribution, foraging); Michener 2007a: 827.

Lepidotrigona ventralis (Smith, 1857)

Trigona ventralis Smith 1857: 50: Type (BMNH 17b.1186); Mt. Ophir specimen is latebalteata. The ventra-



lis species group (taxonomy); Type locality: MALAYSIA "Borneo (Sarawak)"; "Malacca (Mt. Ophir)"(worker); Smith 1871: 395 (distribution); MALAYSIA, Borneo; Malacca; Gribodo 1891: 109 (compara-tive 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); Schwarz1939a: 85, 94, 134*, 135, 136, 139 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure1961: 199 (systematic position); Sakagami 1975: 66, 67-75* (distribution, morphology, variation); CAM-BODIA; LAOS; MALAYSIA; THAILAND; VIETNAM; Sakagami 1978: 239 (comparative note); Sak-agami 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: (distribu-tion, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener & Roubik 1993:260 (comparative note, morphology); Yong 1993: (biochemistry); Roubik 1996: 61 (distribution); BRU-NEI, Kuala Belalong; Nagamitsu & Inoue 1997: 434, 435, 436 (pollination); Floral record: Santiria lae-vigata; Nagamitsu & Inoue 1998: (foraging, morphology); Liow 1999: 106, 110, 127 (distribution (asventra! 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, mor-phology); 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!)); Santos-Filho et al. 2006: 208, 217 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAI-LAND, Kanchanaburi, Thong Pha Phum; Sung et al. 2006c: (see hoosana); Rasmussen & Cameron2007: (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 isunavailable for taxonomic purposes, as it must be deemed to be infrasubspecific (ICZN, art. 15.2 and45.6.3.) (variation); VIETNAM.

Melipona tunneli Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 13.1.1). The statement that it "is differentfrom the other two in appearance as it possesses somewhat yellow spots in the abdomen", however, couldmake it a nomen dubium, although most likely then a junior synonymon of arcifera (common name (ngapsiwor), honey, nest).

Lisotrigona

Lisotrigona cacciae (Nurse, 1907)

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



abad, Central Provinces" (workers); Moure 1961: 194, 195 (lectotype designation (see Engel 2000), mor-phology, systematic position); Sakagami et al. 1985: (key to species (as cassiae!)); Michener 1990: 131(systematic position); Sakagami et al. 1990: 131 (distribution (as cassiae!)); Cunha 1994: (systematics);Michener 1997: 33 (type species); Michener 2000: 793; Engel 2000: (key to species, taxonomy);MALAYSIA, Sandakan, Borneo; THAILAND, Pat Meeung mountains; Chantabari; "Yunzalin"; Jobiraj& Narendran 2004: 40, 43 (comparative note, key to species); Michener 2007b: (morphology);Michener 2007a: 817.

Trigona scintillans Cockerell 1920a: 116: Holotype (BMNH 17b.1115) (taxonomy); Type locality:MALAYSIA "Sandakan, Borneo (Baker)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA,Sandakan; Cockerell 1929c: 590 (distribution, ecology ("great nuisance in camp setting on one's handsand face")); CAMBODIA, Angkor; Pah Meeung Mt.; Pahtoop Mt.; Schwarz 1937: 289, 290, 307*, 327,plate 7 (key to species, taxonomy); Schwarz 1939a: 85, 86, 94, 130*, 132 (distribution, key to species,taxonomy); Schwarz 1948: 118; Moure 1961: 193, 195 (morphology, systematic position); Sakagami &Yoshikawa 1961: 440 (distribution); THAILAND, Bangkok; Sakagami 1975: 51-52* (distribution, mor-phology); LAOS; THAILAND; VIETNAM; Appanah 1982: (pollination); MALAYSIA, Negri Sembi-ran, Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution,key to species); Michener 1990: 96, 100, 105, 131 (illustration, morphology, systematic position); Saka-gami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra; LAOS, MALAY-SIA; THAILAND; VIETNAM; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat;Inoue et al. 1990b: (population); Oldroyd et al. 1992: 5 (pollination); THAILAND, Chanthaburi; Floralrecord: Archontophoenix alexandrae; Michener & Roubik 1993: 257, 258 (comparative note, morphol-ogy); Nagamitsu & Inoue 1997: 436 (pollination); Floral record: Santiria laevigata; Nagamitsu et al.1999: (floral records (not by species), foraging); Engel 2000: (synonymy); Rajitparinya 2000: (distribu-tion (as scintillan! in part)); Rajitparinya 2000: (distribution, nest); Samejima et al. 2004: (population);MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (distribution, foraging);Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAI-LAND, Kanchanaburi, Thong Pha Phum; Michener 2007b: 130 (synonymy).

Lisotrigona carpenteri Engel, 2000

Lisotrigona carpenteri Engel 2000: 232-235, 236, figs. 1-3: Holotype (IEBR); 3 paratypes (AMNH, IEBR)(illustration, key to species, taxonomy); Type locality: VIETNAM "VIETNAM: Nghê An Province: KheBo, 19 03N 104 43E, 25-28 April 1998, 123 m, James M. Carpenter" (worker); VIETNAM, Ha Thing, SEHuong Son; Jobiraj & Narendran 2004: 39 (key to species); Chinh et al. 2005: 363, 365-367, 370(nest); VIETNAM, Cucphuong National Forest, Red river delta; Michener 2007b: 130 (comparativenote).

Lisotrigona furva Engel, 2000

Lisotrigona furva Engel 2000: 235-236: Holotype (SEMC); 5 paratypes (AMNH, SEMC) (floral record, keyto species, taxonomy); Type locality: THAILAND "THAILAND: Nakhon Ratchasima Prov.: ThaChang, 10 February 1993, S. Boongird and C. Michener" (worker); THAILAND, Chiang Mai, W Fang;Floral record: Buddleja asiatica; Callistemon; Jobiraj & Narendran 2004: 40 (key to species);Michener 2007b: (morphology).

Lisotrigona mohandasi Jobiraj & Narendran, 2004



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, KeralaForest 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 spe-cies); 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 (syn-onymy); Moure 1961: 206 (synonymy); Camargo 1988: 370 (synonymy).

Trigona canifrons Smith 1857: 51: Syntype (OUMNH, 1 worker); additional putative type (BMNH17b.1183) (taxonomy); Type locality: MALAYSIA "Borneo (Sarawak)" (worker); Smith 1871: 395 (dis-tribution); MALAYSIA, Borneo; Dalla Torre 1896: 576 (citation); Bingham 1897: 560, 562 (distribu-tion, key to species, taxonomy); INDONESIA, Sumatra; MALAYSIA, Borneo; MYANMAR, Tenasserim;Cockerell 1905: 220 (see laeviceps); AUSTRALIA, Adelaide River (misidentification); SRI LANKA(misidentification, *iridipennis group); Bingham 1905: 58 (citation, distribution); Cameron 1908: 192,194 (comparative notes, key to species); Schulz 1909: (see laeviceps); Cockerell 1911: 176 (comparativenotes, taxonomy, misidentification, uncertain identity); Ihering 1912: 3 (citation); Friese 1914b: 53, 54,57 (identification, previous misidentification); Cockerell 1918: 385 (comparative notes); Cockerell1919b: 244 (comparative note); Prell 1919: (nest (in human skull)); Cockerell 1922a: 279 (misidentifica-tion); Cockerell 1923: 241, 242 (key to species); MALAYSIA, Sarawak; Cockerell 1929e: 242 (compar-ative note); Dover 1929: 62 (citation, distribution); SINGAPORE; Schwarz 1932: 246; Rayment 1932a:184 (citation); Rayment 1932f: 106 (taxonomy); Friese 1933a: 46 (distribution); INDONESIA, Meliau;Rayment 1935: 735, 740 (distribution, key to species, taxonomy); Schwarz 1937: 282, 289, 307-308*,320, 326, 327, plate 2, 5 (key to species, taxonomy); Floral record: Mallotus; Schwarz 1939a: 85, 88, 89,90, 92, 98, 101-102*, 111, 112 (distribution, key to species, taxonomy); Floral record: Mimosa pudica;Schwarz 1948: 18, 23, 117, 165; Moure 1961: 205, 206 (systematic position); Michener 1965: 231 (cita-tion); Yoshikawa et al. 1969: 173 (distribution); MALAYSIA, Kuala Lumpur; Michener 1974: 217; Sak-agami 1975: 55-56* (distribution, variation); MALAYSIA; THAILAND; Sakagami 1978: 239(comparative note); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve;Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Sakagami& Khoo 1987: 207 (comparative note); Uji 1987: (pollination); INDONESIA, E Kalimantan; Floralrecord: Nephelium lappaceum; Camargo 1988: 370 (synonymy); Roubik 1989: 105, 193; Michener1990: 88, 97, 105, 120, 126, 127 (systematic position); Sakagami et al. 1990: (distribution, key to spe-cies); INDONESIA, Sumatra; MALAYSIA, Borneo; MYANMAR; THAILAND; Salmah et al. 1990:(distribution, nest (hollow trunks of Ficus: Moraceae)); INDONESIA, Sumatera Barat; Inoue et al.1990b: (population); Roubik 1990b: 253 (defense); Inoue 1990: 582 (nest (huge tree cavities), popula-tion); Johnson & Appanah 1990: 586 (predator (Oecophylla: Formicidae: Hymenoptera) (as carni-frons!)); Roubik 1992: 345 (citation); Michener & Roubik 1993: 260, 264 (comparative note,



morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cunha 1994: (systematics); Yamane etal. 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 laevi-gata; Michener 1997: 34 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Momose etal. 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); MALAY-SIA, 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 Baramarea; 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, Kan-chanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics); Michener 2007a: 804, 824-826.

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: MALAY-SIA "Sarawak: Foot of Mt. Dulit, junction of rivers Tinjar and Lejok" (2 workers Aug. 2, 1932; 10 work-ers Aug 3, 1932); "Mt. Kalulong, 1800 ft." (1 worker); Schwarz 1939a: 85, 90, 102 (distribution, key tospecies, 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 (distri-bution); MALAYSIA, Sandakan; Schwarz 1937: 282, 383, 286, 291, 320*, 321, 327 (key to species, tax-onomy); 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); Camargo1988: 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 (notby species), foraging); Michener 2000: 801; Eltz et al. 2002a: 30 (distribution, foraging, nest, popula-tion); MALAYSIA, Danum Valley; Sepilok; Deramakot; Eltz et al. 2003: (nest, tree preference); Eltz2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baramarea; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Rasmussen & Cameron 2007: (systemat-ics); 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, desig-nated Michener 1990, "Manikion / 16.28.II.03", "Trigona / genalis / 1904 Friese det. / n. Fr.", "Type" (redlabel), "Coll. Friese", "Zool. Mus. / Berlin"; paralectotype (ZMHB (3)) (distribution, taxonomy); Typelocality: INDONESIA (IRIAN JAYA) "Manikion, 14.-28. Februar 1903 (Neu-Guinea)" (several work-



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 nearKerema; Michener 1965: 231 (citation); Wille & Michener 1973: 14, 24, 48, 59, 70 (nest); Michener1990: 95, 97, 105, 122, 127, 153-157 (distribution, illustration, lectotype designation, morphology, sys-tematic position); INDONESIA, Irian Jaya, Bodem; Bodem Sarmi area; Klamono oil fields; Waris S ofHollandia; PAPUA NEW GUINEA, Dreikikir; Marua river near Kerema; Northeast New Guinea; One;Sepik; Michener & Roubik 1993: 260 (comparative note, morphology); Snelling 1996: 131 (distribu-tion); 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 num-ber) (distribution, key to species, taxonomy); Type locality: MALAYSIA "MALAYA. State of Selangor:Bukit Kutu, 200 feet, Sept. 21, 1932, (H. M. Pendlebury)" (workers, holotype); "State of Pahang: KualaTeku, 500 feet, Dec. 8, 1921 (H. M. Pendlebury)" (workers); "N. BORNEO.-Bettotan, nr. Sandakan, (C.B. Kloss and H. M. Pendlebury)" (workers, July 28, 1927, and Aug. 15, 1927); Schwarz 1948: 118;Moure 1961: 193-194, 195 (morphology, systematic position); Yoshikawa et al. 1969: 174 (distribution);MALAYSIA, Fraser's Hill, Selagor-Pahang; Kiew 1985: (luring (unwashed shorts)); Michener 1990: 131(systematic position); Michener 2001: 235 (synonymy); San & Osawa 2002: (population); MALAYSIA,Negri Sembiran, Pasoh Forest Reserve; Inson & Malaipan 2006: (distribution, foraging, nest); THAI-LAND, Kanchanaburi, Thong Pha Phum.

Trigona (Hypotrigona) pendleburyi Schwarz 1939a: 85, 86, 94, 130-132*: Holotype (BMNH, no number);paratypes (BMNH (5)) (distribution, key to species, taxonomy); Type locality: MALAYSIA "CameronHighlands, Rhododendron Hill, 5000 feet (H. M. Pendlebury)" (workers, holotype, June 19,1923, andJune 22, 1923); MALAYA.-State of Pahang: Cameron Highlands, G. Tarbakar, 4500 feet, June 9, 1923 (H.M. Pendlebury); Cameron Highlands, Bukit Lendong, 5000 feet, May 21, 1931 (H. M. Pendlebury);Lubok Tamang, 3500 feet, March 5, 1924, and 4000 feet, June 10, 1923 (H. M. Pendlebury); Fraser's Hill,4600 feet, Pine Tree Hill, Oct. 25, 1932 (H. M. Pendlebury); Schwarz 1948: 118; Moure 1961: 192, 193-194, 195 (morphology, systematic position); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran,Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, keyto species); Michener 1990: 96, 100, 105, 129, 131 (illustration, morphology, systematic position); Saka-gami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; MALAYSIA; Salmah et al. 1990:(distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener & Rou-bik 1993: 257, 258, fig.15 (comparative note, morphology); Cunha 1994: (systematics); Roubik 1996:61, 62 (distribution, nest); BRUNEI, Kuala Belalong; Michener 1997: 46 (type species); Michener 2000:796; Engel 2000: (comparative note); Chinajariyawong & Saiboon 2000: (distribution, nest); THAI-LAND, Songkhla; Michener 2001: (distribution, morphology (male characters)); BRUNEI; INDONE-SIA, Sumatra; MALAYSIA, Sabah; THAILAND; Eltz et al. 2002a: 30 (distribution, foraging, nest,population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Eltz et al. 2003: (nest, tree preference);Jobiraj & Narendran 2004: 39; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Momose &Hamid 2005: 67, 71; Roubik 2006: 127 (nest); Inson & Malaipan 2006: (distribution, foraging, nest);THAILAND, Kanchanaburi, Thong Pha Phum; Michener 2007a: 821.



Platytrigona

Platytrigona atricornis (Smith, 1865)

Trigona atricornis Smith 1865: 94: Holotype (OUMNH, worker): According to Baker (1993) the specimen islabeled " 'N' [New Guinea (Allen); white disc] and 'Trigona atricornis' 'Smith' ". The atricornis speciesgroup (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 (compar-ative 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, sys-tematic position).

Platytrigona flaviventris (Friese, 1909)

Trigona flaviventris Friese 1909("1908"): 354, 356, 357-358, fig. 15-6: Lectotype (ZMHB, worker): heredesignated, "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); Friese1909: 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; Hohmann2003: (communication, nest); Michener 2007a: 824.

Platytrigona hobbyi (Schwarz, 1937)

Trigona hobbyi Schwarz 1937: 283, 288, 298-300*, 328: Holotype (BMNH 17b.1118). The hobbyi speciesgroup (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, 4000 feet, moss for-est, Oct. 18, 1932" (1 worker); Schwarz 1939a: 85, 129, 130* (distribution, key to species, taxonomy);Moure 1961: 205 (systematic position); Sakagami et al. 1985: (distribution); Michener 1990: 127 (sys-tematic position); Sakagami et al. 1990: 131 (distribution); Michener & Roubik 1993: 260 (comparativenote, morphology); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Samejima et al. 2004:(population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (distribu-tion, foraging); Rasmussen & Cameron 2007: (systematics).

Platytrigona keyensis (Friese, 1901)

Trigona keyensis Friese 1901: 271: Lectotype (ZMHB, worker): here designated, "AsiaArch. / Key Ins. /1900 / Kühn", "Trigona / keyensis / 1900 Friese det. / Fr.", "Type" (red label), "Coll. / Friese"; paralecto-type (AMNH (1)). The planifrons species group (manuscript name (latigena in litt.), taxonomy); Typelocality: INDONESIA "Key-Eilanden (Amboina, Nederland India) durch H. Kühn" (6 workers); Friese1909("1908"): 354, 356, fig. 15-7 (distribution); INDONESIA, Irian Jaya, Moaif; Timena; Friese 1909:272, 273, 277 (key to species, taxonomy); INDONESIA, Irian Jaya, Moaif; Timena; Strand 1911: 87 (dis-tribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Holl. N.G.; Pauwi; Samberi; Teba; Friese1912: 94 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, D. Neu Guinea; Pauwi; Sam-



beri; Teba; Strand 1912: (distribution); Friese 1915: 4 (distribution); INDONESIA, Irian Jaya/PAPUANEW 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; Hohmann2003: (communication, nest); Michener 2007a: 824.

Platytrigona lamingtonia (Cockerell, 1929)

Trigona lamingtonia Cockerell 1929e: 243: Holotype (AMS, worker): examined, "Mt. Lamington / NorthernDivision / Papua v.1927 / C.T. McNamara", "Trigona / lamingtonia / Ckll TYPE", "HOLOTYPE", "Aus-tralian Museum K 238299". The planifrons species group (taxonomy); Type locality: PAPUA NEWGUINEA "Mt. Lamington, May, 1927 (C.T.McNamara)" (workers); Michener 1965: 231 (citation).

Platytrigona planifrons (Smith, 1865)

Trigona planifrons Smith 1865: 93-94: Holotype (OUMNH, worker): According to Baker (1993) the speci-men is labeled " 'N' [New Guinea (Allen); white disc] and 'Trigona planifrons 'Smith' ". The BMNH"type" (17b.1180) indicated by Moure (1961), is not a true type (Baker, 1993). The planifrons speciesgroup (taxonomy); Type locality: INDONESIA (IRIAN JAYA) "New Guinea" (worker); Smith 1871:396 (distribution); "NEW GUINEA"; Dalla Torre 1896: 582 (citation); Friese 1898: 427, 428 (citation,key to species); MALAYSIA, Malacca (misidentification); "NEW GUINEA"; Friese 1909("1908"): 354,356, fig. 15-8 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai;Jamur Gebiet; Manikion; Manokwari; Merauke; Moaif; Tawarin; Friese 1909: 272, 273, 274 (distribu-tion, key to species); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai; JanourGebiet; Manokwari; Merauke, Moaif; Tawarin; Friese 1915: 4 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai; Manikion; Manokwari; Merauke; Moaif; Sattelberg;Sekofro; Simbang; Tawarin; Tjahe Fluss; Zoutbron; Cockerell 1923: 241 (key to species); Cockerell1929e: 242 (comparative note, distribution); PAPUA NEW GUINEA, Fly River; Mt. Lamington;Schwarz 1948: 83, 109, 118; Moure 1961: 203, 204 (systematic position); Michener 1965: 231 (cita-tion); Camargo 1988: 372 (distribution); PAPUA NEW GUINEA, N. Guinea, Simbang, Huon Golf;Michener 1990: 88, 105, 120, 127, 153 (illustration, morphology, systematic position); Michener &Roubik 1993: 260 (comparative note, morphology); Cunha 1994: (systematics); Michener 1997: 48(type species); Michener 2000: 800-801; Hohmann 2003: (communication, nest); Michener 2007a: 824,826.

Sundatrigona

Sundatrigona lieftincki (Sakagami & Inoue, 1987)

Trigona (Trigonella) lieftincki Sakagami & Inoue 1987: 610, 611, 613-615, 617-624: Holotype (RMNH, notlocated, nor in SEHU); Paratypes (RMNH, 3 workers, 2 males) (morphology, taxonomy); Type locality:INDONESIA "N.E. Sumatra, Tongkoh, probably Mt. Talamau (Pasaman), VI 1941, v.d. M. Mohr leg." (5workers, 4 males, holotype worker, paratypes); "Koto Atas, Solok, Sumatera Barat" (6 workers); Sakag-ami & Inoue 1989: 615 (comparative note); Michener 1990: 127 (systematic position); Sakagami et al.1990: (distribution, key to species); INDONESIA, Sumatra; Salmah et al. 1990: (distribution, nest);INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (population); Tadauchi



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); Sakagami1975: 56, 57 (synonymy); Sakagami et al. 1990: 127 (synonymy); Tadauchi et al. 1998: 243 (typedepositories).

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, nearsweat 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); Appanah1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermumintermedium; Sakagami et al. 1983c: (reproduction); Sakagami et al. 1985: (distribution, key to species);Inoue et al. 1985a: (aggressiveness, foraging, nest (in aerial Crematogaster: Formicidae: Hymenopteranests)); 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: (dis-tribution, key to species); INDONESIA, Sumatra; MALAYSIA; Salmah et al. 1990: (distribution, nest (inaerial Crematogaster: Formicidae: Hymenoptera nests)); INDONESIA, Sumatera Barat; Inoue et al.1990b: (population); Inoue 1990: 582 (nest (Crematogaster: Formicidae: Hymenoptera nests), popula-tion); Roubik 1990c: 663 (citation); Salmah et al. 1991: 199 (comparative note); Inoue & Sakagami1993: 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 hom-onym); Nagamitsu & Inoue 1998: (foraging, morphology); Cobara & Dejan 1998: 491 (citation); Nag-amitsu et al. 1999: (floral records (not by species), foraging); Heard 1999a: 187 (citation); Michener2000: 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 (distri-bution, foraging); Roubik 2006: 135 (nest); Rasmussen & Cameron 2007: (systematics); Michener2007a: 826.

Sundatrigona indet. Inoue & Sakagami 1993: 769 (citation); Inoue & Sakagami 1993: 769 (homonym (forTrigonella)); 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); Bing-ham 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:



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, tax-onomy); Floral record: Citrus limon; Schwarz 1948: 117, 290; Moure 1961: 210, 211 (morphology, sys-tematic position); Yoshikawa et al. 1969: 173 (distribution); MALAYSIA, Kuala Lumpur; George Town,Penang; Touaran, Sabah; Sakagami 1975: 59* (distribution); MALAYSIA; THAILAND; Sakagami1978: 230 (comparative note); Wille 1979b: 273 (citation, illustration); Wille 1979a: (morphology (pollenpress)); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floralrecord: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (lur-ing (unwashed shorts)); Sakagami & Khoo 1987: 207, 208, 209 (comparative note); Uji 1987: (pollina-tion); 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, morphol-ogy); Yong 1993: 54 (biochemistry); Cunha 1994: (systematics); Bruijn & Sommeijer 1997: 44 (cita-tion); Michener 1997: 57 (type species); Liow 1999: 127; Michener 2000: 801; Chinajariyawong &Saiboon 2000: (distribution, nest); THAILAND, Songkhla; San & Osawa 2002: (population); MALAY-SIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu & Inoue 2005: 76 (distribution, foraging);Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, foraging, nest); THAI-LAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007: (systematics); Michener2007a: 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 work-ers); Cockerell 1929b: 140 (distribution, nest); CAMBODIA, Angkor; Kum Puang Creek; Nan; Cocker-ell 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 synon-ymy); Sakagami 1975: 59 (synonymy); Sakagami & Khoo 1987: 207 (comparative note).

Trigona collina Smith 1857: 51-52: Type (OUMNH (=Wilson Saunders collection)) (taxonomy); Type local-ity: MALAYSIA "Malacca (Mount Ophir)" (worker); Smith 1871: 396 (distribution); MALAYSIA, Mal-acca; Dalla Torre 1896: 577 (citation); Bingham 1897: 560, 562 (distribution, key to species, taxonomy);MALAYSIA, Malacca; MYANMAR, S Tenasserim; Cameron 1901: 36 (distribution); THAILAND,Patalung; Waterhouse 1903: 133-134, plate 4 (see apicalis); Cameron 1908: 192 (key to species); Cock-erell 1918: 387 (comparative notes); Cockerell 1920a: 116 (comparative note); Cockerell 1923: 241, 242(key to species); Dover 1929: 62 (see apicalis); Schwarz 1937: 283, 288, 297-298*, 300, 301, 302, 326,327, plate 2, 5 (key to species, taxonomy); Schwarz 1939a: 85, 90, 94, 95, 123-125*, 126 (distribution,key to species, taxonomy); Floral record: Mimosa pudica; Schwarz 1948: 23, 117, 165; Schwarz 1951:63 (distribution); CAMBODIA, Angkor; Pagden 1957: 16-17 (flight); Sakagami 1960: 146 (distribu-tion); CAMBODIA, Stung Treng; Moure 1961: 211-212 (morphology, systematic position); Sakagami &Yoshikawa 1961: 437-439 (distribution, nest (only fig. 28)); CAMBODIA, Angkor Wat; Poipet; THAI-LAND, Chieng Mei; Doi Suthep; Li; Mae Hoi; Sara Buri; Yoshikawa et al. 1969: 168, 169, 173, 174 (dis-tribution, nest); MALAYSIA, Fraser's Hill, Selagor-Pahang; George Town, Penang; Nogueira-Neto 1970:291; Wille & Michener 1973: 13, 23, 47, 58, 70, 87 (nest); Sakagami 1975: 59-60*, 61 (distribution,variation); LAOS; MALAYSIA; THAILAND; VIETNAM; Sakagami 1978: 167-181, 231, 236, 237, 239(comparative note); Sakagami 1982: 389 (citation); Sakagami et al. 1983b: 5 (nest); Sakagami et al.



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 tospecies); 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), popula-tion); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Yong 1991: (bio-chemistry); 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 (com-parative 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 spe-cies), foraging); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Eltz etal. 2001a: (pollen foraging); Eltz et al. 2001b: (foraging); MALAYSIA, Borneo, Sabah; Floral record:Caesalpinia; Citrus; Clausena; Croton; Durio; Jasminium; Manihot esculenta; Mimosa pudica; Neonau-clea; Passiflora; Sterculia; Symplocos; Terminalia; Asteraceae; Leguminosae; Green et al. 2001: 91(microsatellite); Eltz et al. 2002b: (foraging (Rhizopus: Mucorales: Zygomycota)); Eltz et al. 2002a: (dis-tribution, 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, microsatel-lite); 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: Diospyrosrhodocalyx; 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 & Wattanachaiying-charoen 2006: (foraging); THAILAND, Phitsanulok; Floral record: Agave angustifolia; Alangium salvii-folium; Butea monosperma; Caryota bacsonensis; Cassia bakeriana; Coccinia grandis; Costusspeciosus; Croton roxburghii; Dalbergia lanceolaria; Delonix regia; Erythrina stricta; Fernandoa ade-nophylla; 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; Toreniafournieri; Tridax procumbens; Zingiber officinale; Rasmussen & Cameron 2007: (systematics); Leon-hardt et al. 2007: (foraging); Floral record: Archidendron jiringa; Jongjitvimol & Wattanachaiying-charoen 2007: (nest); Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (predator (Argiope:Araneidae: Araneae; Cypsiurus balasiensis: Apodiformes: Apodidae; Dicrurus macrocercus; Dicrurusparadiseus: Passeriformes: Dicruridae; Merops leschenaulti: Meropidae: Coraciiformes; Nephila macu-lata: 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



1948: 117; Moure 1961: 212 (systematic position); Yoshikawa et al. 1969: 173 (distribution); MALAY-SIA, 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); INDO-NESIA, 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, San-dakan; 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); Sakagami1975: 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); Yong1991: (biochemistry); Michener & Roubik 1993: 260 (comparative note, morphology (as rufibasis!));Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433, 436 (pollina-tion); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsuet al. 1999: (floral records (not by species), foraging); Nagamitsu & Inoue 2002: 305, 307, 308 (forag-ing); 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); Sakagamiet 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); Typelocality: PHILIPPINES "Philippinen (Schadenburg)" (few workers); "N-Guinea (Berlinhafen und



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, flo-ral record, key to species); PHILIPPINES, Bataan, Lamao; Luzon, Los Banos; Floral record: Nipa fruti-cans; Cockerell 1925c: 53, 55 (distribution); PHILIPPINES, Bangui, Ilocos norte; Limay; Cockerell &LeVeque 1925: 175 (distribution, floral record); PHILIPPINES, Loquilocon; Samar; Wright (formerParanas); Floral record: Citrus; Melicope triphylla; Mimosa pudica; Hedicke 1926: 423 (distribution);PHILIPPINES, Luzon; Mindanao; Siargao; Williams 1928: 91-92 (nest); Cockerell 1929d: 301 (compar-ative note (as birsi!)); Cockerell 1930a: 157 (distribution, floral record, uncertain identity); CAMBODIA,Tung Nui (misidentification); Satul (misidentification); Floral record: Melastoma malabaricum; Schwarz1939a: 85, 86, 90, 101, 119-120* (distribution, key to species, taxonomy); Floral record: Citrus; Melicopetriphylla; Mimosa pudica; Nipa fruticans; Moure 1961: 209 (systematic position); Michener 1961: 28(comparative note); Michener 1965: 231 (citation); Baltazar 1966: 406-407 (distribution); PHILIP-PINES, 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; Calliste-mon citrinus; Dillenia suffruticosa; Jatropha podagrica; Mussaenda; Nerine; Nicotiana tabacum; Plum-bago auriculata; Roubik 1995: 151 (antropogenic nest); Barile & Cervancia 1995: (ecology, flight,floral record, foraging, nest); Floral record: Antigonon leptopus; Asystasia gangetica; Callistemon citri-nus; Coccinia; Cocos nucifera; Dillenia suffruticosa; Euphorbia pulcherrima; Galphimia glauca; Hame-lia patens; Impatiens balsamina; Ixora; Jatropha hastata; Jatropha podagrica; Leucaena leucocephala;Mussaenda; Nerine; Nicotiana tabacum; Plumbago auriculata; Dollin et al. 1997: 877 (comparativenote, distribution); PHILIPPINES; New Guinea; Oshy & Arikumar 2000: 1651 (pollination); Floralrecord: 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 prop-erties).

Trigona bismarkiana Tadauchi et al. 1998: 245: Nomen nudum. The name is unavailable for taxonomic pur-poses, as no description or definition of the taxon was provided (ICZN, art. 15.1) (type depositories).

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 subspeciesof 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); Moure1961: 209 (morphology, systematic position); Michener 1965: 231 (citation); Camargo 1988: 372 (tax-onomy); 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 (cita-tion); Friese 1898: 428, 429, 430 (comparative note, key to species); Friese 1905: 118 (citation); Friese



1909: 272, 273-274 (key to species, taxonomy); AUSTRALIA, Mackay; Sydney; INDONESIA, Key-Inseln (misidentification); MICRONESIA, Karolinen (misidentification); PAPUA NEW GUINEA, Sat-telberg (misidentification); Simbang (misidentification); Cockerell 1910: 247 (comparative notes); Lea1910: 119, 207 (inquiline (Brachypeplus auritus; Brachypeplus basalis: Nitidulidae: Coleoptera)); Strand1911: 87 (distribution); Cockerell 1911: 176 (comparative notes, taxonomy); Friese 1912: 94 (distribu-tion); Strand 1912: (distribution); Cockerell 1912: 225 (comparative notes); Lea 1912: 38-39 (inquiline(Homalota trigonae: Staphylinidae: Coleoptera)); Cockerell 1914: 412-413; Floral record: Helianthusannuus; Friese 1917: 9 (distribution); Cockerell 1922a: 279 (comparative note); Salt 1929: 452, 453(citation); Cockerell 1929e: 243 (distribution); AUSTRALIA, Ourimbah, New South Wales; Cockerell1929d: 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, vari-ation); 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: (behav-ioral comparison); Rayment 1932e: (inquiline (Brachypeplus planus; Brachypeplus meyricki (orEpuraea): Nitidulidae: Coleoptera; Tribolium myrmecophilum: Tenebrionidae: Coleoptera)); Rayment1932f: 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 (synon-ymy); 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 LamingtonNational 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); Michener1965: 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 & Mat-thews 1973: (predator (Bembix musca: Crabronidae: Hymenoptera)); Evans et al. 1982: 78 (predator(Bembix musca: Crabronidae: Hymenoptera)); Sakagami et al. 1983b: 20 (comparative note); Sakagamiet al. 1983a: 106 (comparative note); Sakagami et al. 1983d: 454 (comparative note); Wagner & Briscoe1983: (genetics); Hawkeswood 1983: (pollination); Floral record: Cupaniopsis anacardioides; Graur1985: 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, mor-phology); 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;



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; Heard1994: (pollination); Floral record: Macadamia integrifolia; Heard & Exley 1994: (pollination); Floralrecord: Macadamia integrifolia; Forster et al. 1994: 219 (pollination); Floral record: Cycas; Bartareau1994: (pollination); Floral record: Dendrobium toressae; Yamane et al. 1995: (oviposition); Wallace &Trueman 1995: (seed dispersal (mellitochory)); Floral record: Eucalyptus torelliana; Disney & Bartar-eau 1995: (predator (Dorniphora trigonae: Phoridae: Diptera) (of brood?)); Bartareau 1995: (pollina-tion); 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: (bee-keeping, nest); Dollin 1996a: (characteristics, nest); Dollin 1996b: (nest); Dollin 1996c; Nogueira-Neto1997: 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: Abro-phyllum ornans; Acradenia euodiiformis; Alphitonia excelsa; Euroschinus falcata; Rhodomyrtus psidio-ides; Tristaniopsis laurina; Williams & Adam 1997: 75 (floral record); Floral record: Cuttsia viburnea;Dollin & Heard 1997: (domestication); Dollin & Heard 1997: (bee-keeping, predators (Phoriae; Syr-phidae)); 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); Gardeneret al. 2003: 120 (citation); Williams & Walker 2003: 143, 144, 145 (floral record); Floral record: Polliacrispata; 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); Flo-ral record: Cucumis sativus; Amano 2004: 3 (introduction, pollination); Floral record: Lycopersicon escu-lentum; 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); Ariaset 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 (Argi-ope keyserlingi: Araneidae: Araneae)); Rasmussen & Cameron 2007: (systematics); Michener 2007a:100, 807f, 824; Jongjitvimol & Wattanachaiyingcharoen 2007: 29 (citation); Stow et al. 2007: (antimi-crobials, honey); Gloag et al. 2007: (reproduction); Fidalgo & Kleinert 2007: 138 (citation); Klumpp2007; 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 &



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); AUS-TRALIA, NE coastal; Queensland; Ikudome & Kusigemati 1996: 17, 19-20 (common name (bai), distri-bution, 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); Ray-ment 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 (synon-ymy); 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 largerform of laeviceps from the Philippines (taxonomy); Type locality: PHILIPPINES "Dapitan, Mindanao(Baker, 23119)" (worker); Schwarz 1939a: 86, 112, 113 (synonymy); Moure 1961: 208 (morphology,systematic position).

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, Mont-pellier" (8)) (genetics, illustration, microsatellite, nest, taxonomy); Type locality: AUSTRALIA "AUS-TRALIA, 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 fromMr. R.C.J. Swinhoe" (worker); Cockerell 1922b: (age of fossil); Kerr & Maule 1964: (systematics, tax-onomy); Kerr & Esch 1965: (comparative note); Kerr 1969: 166 (systematics); Kerr & Cunha 1976:(systematics); Zeuner & Manning 1976: 222, 223, 224 (synonymy); Wille 1977: (systematics); Moure& Camargo 1978: 564-565 (morphology); Sakagami 1978: 216, 218-219 (synonymy); Wille 1979b: 243(comparative note); Grimaldi et al. 1995: 261, 262; Wu 2000: 392 (synonymy); Zherikhin & Ross2000: 6, 13-14 (synonymy).



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; Sakagami1959: 120 (citation); Moure 1961: 210 (morphology, systematic position); Yoshikawa et al. 1969: 174(distribution); MALAYSIA, Tapah; Sakagami 1978: 210 (synonymy); Sakagami et al. 1983b: 10-12, 20(nest); Sakagami & Inoue 1985: 174, 176, 177, 178, 179, 180, 181, 182-184*, 186, 187-188 (key to spe-cies, nest, taxonomy); Sakagami et al. 1985: (distribution, key to species); Sakagami & Inoue 1987: 626(comparative note); Starr & Sakagami 1987: 97 (citation); Roubik 1989: 105; Sakagami et al. 1989:266 (comparative note); Sakagami et al. 1990: (distribution, key to species); INDONESIA, Java;Sumatra; MALAYSIA, Borneo; Sakagami & Inoue 1990: 201 (mixed brood arrangement); Salmah et al.1990: (distribution, nest (beetle burrows; crevices)); INDONESIA, Sumatera Barat; Inoue et al. 1990b:(population); Roubik 1996: 61, 62 (distribution, nest, predator (Helarctos malayanus: Ursidae: Car-nivora)); BRUNEI, Kuala Belalong; Michener & Boongird 2004: 144, 145 (comparative note); Nag-amitsu & Inoue 2005: 77 (distribution, foraging).

Tetragonula fuscobalteata (Cameron, 1908)

Trigona atomella Cockerell 1919b: 243-244: Holotype (USNM 29467, worker): examined, "Island of / Pen-ang / Baker", "Type No. / 29467 / U.S.N.M.", "Trigona / atomella / Ckll TYPE"); 2 paratypes (USNM)(comparative notes, taxonomy); Type locality: MALAYSIA "Island of Penang (Baker, 9585)" (worker);Cockerell 1920a: 116 (comparative note); Cockerell 1929c: 590 (distribution, ecology ("found clingingto the leg of a large green cicada (Platylomia spiculata (=Dundubia spiculata): Cicadidae: Hemiptera)"));CAMBODIA, Pahtoop Mt.; Cockerell 1930a: 157 (distribution, ecology (attracted to sweat)); CAMBO-DIA, Chun Het; Trang; Yasumatsu 1935: 94 (common name (mitsubachi), distribution); PALAUISLANDS; Esaki 1936: 44, 46-37 (common name (galadngikl; chaladngikl; mitsubachi), nest); PALAUISLANDS; Schwarz 1937: 290, 310, 311*, 316, 328 (taxonomy); Schwarz 1939a: 107, 108 (synonymy);Schwarz 1939b: 152 (synonymy); Cockerell 1939a: 61, 62, 64 (distribution ("presumably introduced byman")); PALAU ISLANDS, Melekeiok on Babelthuap; Ngeremlengui, Ogiwal; Schwarz 1948: 69, 117;Krombein 1950: 133 (synonymy); Moure 1961: 209 (morphology, systematic position); Sakagami1978: 223, 224 (synonymy); Camargo 1988: 371 (synonymy, taxonomy); Sakagami et al. 1990: 128(synonymy).

Trigona brevis Friese 1925: 41: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 12.1) (comparative note).

Trigona erythrostoma Cameron 1908: 193-194: Syntypes (BMNH 17b.1113, 2 workers) (key to species, tax-onomy); Type locality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker or queen); Cockerell1918: 386 (comparative notes); Cockerell 1920a: 116 (comparative note); Schwarz 1937: 290, 310*, 326(key to species, taxonomy); Schwarz 1939a: 112 (synonymy); Moure 1961: 195, 209 (synonymy).

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); Schwarz1937: 290, 310-311*, 316, 326 (key to species, taxonomy); Schwarz 1939a: 85, 86, 88, 90, 93, 96, 107-110*, 111, 113, 114, 115, 132 (distribution, key to species, taxonomy); Schwarz 1939b: 151-152 (distri-bution, taxonomy); PALAU ISLANDS, Arukoron; Babelthaob, Marukyoku-Kaisharu; Korror; Schwarz1948: 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



1959: 121 (comparative note); Moure 1961: 209 (morphology, systematic position); Michener 1961: 34(comparative note); Sakagami & Yoshikawa 1961: 435 (comparative note); Baltazar 1966: 407 (distri-bution); PHILIPPINES, Mindanao, Lanao; Palawan; Yoshikawa et al. 1969: 168, 169, 172, 173, 174 (dis-tribution, 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); Sakagami1982: 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 (comparativenote); Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Yong1986: (allozyme); Sakagami & Khoo 1987: 213 (comparative note); Sakagami & Inoue 1987: 626(comparative note); Starr & Sakagami 1987: (common name (kiyot), nest); Yong et al. 1987: (biochem-istry); 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 spe-cies, 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, popu-lation); 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, taxon-omy (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 & Saiboon2000: (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 etal. 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, polli-nation); 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; Sakag-ami et al. 1990: 128 (synonymy).

Trigona pfeifferi Friese 1925: 41: Lectotype (AMNH, worker): here designated, "Brasil / Campinas", "Trig-ona / pfeifferi / Friese det. 25 / Fr.", "Typus" (orange label), "Heterotrigona / cf. fuscobalteata / (erroneouslocality) / det. J.S.Ascher"; paralectotypes (DEI (3)) (taxonomy); Type locality: UNCERTAIN "Campi-nas (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



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)) (commonname (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 (syn-onymy); 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); Schwarz1937: 282, 290, 311-313* (key to species, taxonomy); Schwarz 1939a: 101, 117 (synonymy); Moure1961: 209 (synonymy); Sakagami 1978: 195, 196 (synonymy).

Trigona geissleri Cockerell 1918: 385-386, 387: Unknown: Cockerell used a Friese manuscript name for hisdescription (key to species, distribution, manuscript name of Friese, taxonomy, variation); Type locality:MALAYSIA "Sintang, North Borneo" (male); "Singapore by Baker" (couple of workers); Cockerell1919b: 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 wereincluded here)); Schwarz 1948: 83, 118, 233; Moure 1961: 209 (morphology, systematic position); Sak-agami & 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, 195-198*, 199, 200, 208, 209, 210, 226, 227, 228, 231, 234, 235, 236, 237, 238, 239 (taxonomy); MALAY-SIA; SINGAPORE; THAILAND; Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, PasohForest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1983b: 11, 20 (nest); Sakag-ami & 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 & Khoo1987: 208, 209, 210, 211, 212 (morphology); Sakagami & Inoue 1987: 625, 626 (morphology); Starr &Sakagami 1987: 97 (citation); Camargo 1988: 371 (distribution); MALAYSIA, Borneo, Sandakan; Sak-agami et al. 1990: 128, 131 (distribution, misidentification); Oldroyd et al. 1992: 5 (pollination); THAI-LAND, Chanthaburi; Floral record: Archontophoenix alexandrae; Roubik 1995: 151 (antropogenic nest);Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Bruijn & Sommeijer 1997: 44 (citation (asgeisslerrii!)); Liow 1999: 106, 127 (distribution (as geisslerei! in part)); Nagamitsu et al. 1999: (floralrecords (not by species), foraging); Liow et al. 2001: 184; San & Osawa 2002: (population); MALAY-SIA, 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"



(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 (com-parative note); Sakagami et al. 1990: 131 (distribution); Sakagami & Inoue 1990: 202 (comparativenote); Tadauchi et al. 1998: 243 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution, forag-ing).

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 (1worker); "Mae Klang" (2 workers); "Mae Fack" (1 worker); "Lamphun" (1 worker); "Uthaithani" (1worker); 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: (system-atics).

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 (comparativenote, 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 & Dollin1982c; 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 (com-parative 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: Dendro-bium; Bartareau 1994: (pollination); Floral record: Dendrobium toressae; Yamane et al. 1995: 294 (syn-onymy); 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, revi-sion, 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); Dollin2002a: (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



& 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" (orangeborder), "iridipennis / Type Sm.", "B.M.TYPE / HYM. / 17B.1114", "Trigona / iridipennis / TYPE.Smith.", "Ceylon" (reverse side "53 / 23"). Citations for iridipennis are included in a broad sense, as spe-cies limits for this taxon is uncertain. To the same group belongs also T. bengalensis, T. praeterita, and T.ruficornis, as well as several undescribed species (see Rasmussen & Cameron (2007)) (taxonomy); Typelocality: SRI LANKA "Ceylon" (worker); Tennent 1859: 282 (list); Tennent 1861: 454 (list); Castets1893: 468, 481-488 (nest); Dalla Torre 1896: 579 (citation); Bingham 1897: 560, 563-564 (distribution,key to species, taxonomy); MYANMAR, Tenasserim; SRI LANKA; Friese 1904: 140 (see laeviceps);Friese 1909: 273 (comparative note); Strand 1913a: 149-150 (distribution (as laeviceps)); SRI LANKA;Friese 1914a: 12-13 (see laeviceps); Friese 1914a: 12 (synonymy (as canifrons)); Friese 1914a: 12 (syn-onymy (as laeviceps)); Friese 1914b: 49, 57 (citation (as laeviceps)); Friese 1914b: 43-58 (citation, nest);Friese 1914b: 53-54, 57 (citation, synonymy (as canifrons)); Cockerell 1919b: 242 (comparative notes);Cockerell 1919c: 79 (comparative notes, key to species); Cockerell 1919a: 121 (see laeviceps); Cocker-ell 1929c: 590, 592 (distribution, floral record, taxonomy, variation); CAMBODIA, Angkor (misidentifi-cation); Kum Puang Creek (misidentification); Nan (misidentification); Pahtoop Mt. (misidentification);INDIA, Bombay; Mangalore; PHILIPPINES, Palawan, Puerto Princesa (misidentification); SRI LANKA;THAILAND, Bangkok (misidentification); Floral record: Helianthus annuus; Cockerell 1930b: 149 (dis-tribution, floral record); MYANMAR, Rangoon; Floral record: Helianthus annuus; Friese 1933a: 46 (seelaeviceps); George 1933: (physiology (as irridipennis!)); George 1934: (brood, distribution, morphology,nest (crevice of wall), physiology (as irridipennis!)); INDIA, Coimbatore; Schwarz 1937: 283, 290, 308-310*, 314, 316, 326, 328, plate 3, 4 (key to species, taxonomy); Schwarz 1939a: 85, 86, 88, 89, 90, 93,96, 102, 108, 109, 110, 111-116*, 117, 118, 119, 120 (distribution, key to species, taxonomy (most of thespecimens examined by Schwarz correspond to laeviceps, see laeviceps)); Schwarz 1939b: 151 (compar-ative note); Schwarz 1945: 139, 141 (citation); Schwarz 1948: 4, 23, 31, 33, 39, 48, 50, 60, 62, 67, 68,71, 83, 84, 92, 109, 118, 141, 165, 233; Nogueira-Neto 1949: 121 (bee-keeping (as iridipenis!)); Krom-bein 1950: 133, 135 (comparative note); Krombein 1951: 278, 295 (morphology); Lindauer 1956: 546-554, 555 (communication, nest); Pagden 1957: 16 (flight); Lindauer & Kerr 1958: 407, 410 (communi-cation); Sakagami 1959: 120, 121 (comparative note); Portugal-Araújo & Kerr 1959: 226 (comparativenote); Sakagami 1960: 146 (see laeviceps); Lindauer & Kerr 1960: 31, 32, 34 (communication); SRILANKA; Lindauer & Kerr 1960: 31, 32, 34; Kerr 1960: 386 (citation); Moure 1961: 206, 207, 208(morphology, systematic position); Michener 1961: 3, 10, 28-33, 35, 38, 39, 42 (see laeviceps); Sakag-ami & Yoshikawa 1961: 434, 435, 436 (comparative note); Darchen & Louis 1961: 18, 21, 25-27 (bio-geography, communication, nest); Kerr et al. 1963: 81 (citation); Kerr & Maule 1964: (comparativenote); Schmid 1964: 588 (citation); Michener 1965: 231 (citation); Kerr & Esch 1965: (comparativenote); Baltazar 1966: 407 (see laeviceps); Phadke 1968: (honey, uncertain identity (unidentified));INDIA, Mysore, Madhya Pradesh; Kerr 1969: 166 (systematics); Darchen 1969: 482 (flight);Yoshikawa et al. 1969: 168, 169, 173 (see laeviceps); Jander & Jander 1970: 361 (citation); Sakagami1971: 345 (citation); Wille & Michener 1973: 14, 24, 30, 48, 59, 71, 94, 96, 98, 100, 102-103, 257 (nest);Michener 1974: 57, 153, 154, 343 (colony, nest); Sakagami 1975: 59 (comparative note); Kerr &Cunha 1976: (systematics); Zeuner & Manning 1976: 222-224, plate 2 (morphology); Wille 1977: (sys-tematics); Kshirsagar & Chauhan 1977: (oviposition); INDIA, Karnataka; Moure & Camargo 1978:565 (synonymy); Sakagami 1978: 165, 166-194, 196, 197, 200, 201, 202, 203, 205, 206, 208, 211, 212,



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); SRILANKA; Floral record: Boehmeria; Cassia; Cleome; Cocos; Erigeron; Homonoia; Lannea; Mimosa;Ocimum; Phyllanthus; Sapium; Smithia; Solanum; Syzygium; Taxillus; Toona; Tridax; Vitex; Divan1981("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); Utami1986: (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!)); Flo-ral record: Brassica campestris; Cajanus cajan; Mohana Rao & Suryanarayana 1989("1988"): 7 (polli-nation); Floral record: Citrullus lanatus; Utami & Kahono 1989: (pollination); INDIA, Bogor; Floralrecord: Areca vestiaria; Percy 1989: (bee-keeping); Michener 1990: 89, 119, 120, 126, 127 (systematicposition); Sakagami et al. 1990: (misidentification, synonymy); Sakagami & Inoue 1990: 212 (taxon-omy); Goel & Kumar 1990: 44 (pollination); INDIA, W Uttar Pradesh; Floral record: Helianthusannuus; Mohana Rao & Suryanarayana 1992("1990"): (pollination); Floral record: Guizotia abyssin-ica; Oldroyd et al. 1992: 6 (citation); Ramanujam et al. 1993("1992"): (pollination); Floral record:Ageratum conyzoides; Antigonon; Azadirachta indica; Bombax; Cocos nucifera; Cyanotis; Delonixregia; Eucalyptus globulus; Lawsonia alba; Leucaena leucocephala; Loranthus longiflorus; Peltophorumpterocarpum; Prosopis juliflora; Rotala densiflora; Tamarindus indica; Michener & Roubik 1993: 260(comparative note, morphology); Cunha 1994: (systematics); Roubik 1995: 24, 41, 42 (pollination); Flo-ral 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); Nogueira-Neto 1997: 90; Dollin et al. 1997: (distribution, taxonomy); SRI LANKA; Michener 1997: 57 (type spe-cies); Rao et al. 1999("1997"): (pollination); Floral record: Crateva magna; Lakshmi & Suryanarayana1999("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; Heard1999a: 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, uncer-tain 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: Psid-ium guajava; Hohmann 2003: (citation); Devanesan et al. 2003: (predator (Amblyseius: Phytoseiidae:Acarina (of inquilines?); Hermetia illucens: Stratiomyidae: Diptera (of brood); Solenopsis geminata: For-micidae: 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 etal. 2005a: (pollination); INDIA, Bangalore; Floral record: Sechium edule; Eswarappa et al. 2005c: (pol-lination); INDIA, Bangalore; Floral record: Sechium edule; Raju et al. 2005: 1379 (pollination); Floralrecord: Wrightia tinctoria; Raju 2005: 106 (floral record); Floral record: Woodfordia floribunda; Raju et



al. 2006: 940 (floral record); Floral record: Acacia caesia; Raju & Rao 2006: 963, 965 (pollination); Flo-ral 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 speci-men (17b.1184) came from Mt. Ophir, which was not the type locality and the specimen cannot be consid-ered a true type. Baker (1993) located three specimens in OUMNH and labeled one as the holotype.Unfortunately that specimen is identic to fuscobalteata according to Baker (1993), but this issue will beresolved in a separate paper with C.D. Michener. Citations below for laeviceps are in a broad sense, asspecies limits are uncertain and may include both valdezi and testaceitarsis. These taxa have previouslybeen proposed as junior synonymons of laeviceps, but are slightly larger (taxonomy); Type locality: SIN-GAPORE "Singapore" (worker); Smith 1858: (propolis, wax); Smith 1859: 135 (distribution); INDO-NESIA, Aru; INDIA (misidentification); SINGAPORE; Smith 1865: 93 (distribution); "NEWGUINEA"; Parish 1866: (common name (pwai-ngyet), illustration, nest, propolis); Smith 1871: 395 (dis-tribution); SINGAPORE; Anderson 1890: 24 (citation); Dalla Torre 1896: 580 (citation (as leviceps!));Bingham 1897: 560, 564 (distribution, key to species, taxonomy, uncertain identity (as praeterita));MYANMAR, Rangoon; SRI LANKA, Trincomali; Bingham 1897: 560, 563 (distribution, key to species,nest (often build in crevices in the brickwork of the walls of houses), taxonomy); MYANMAR, Tenasse-rim; SINGAPORE; Friese 1898: 427 (key to species); Friese 1904: 140 (floral record, uncertain identity(as iridipennis)); INDONESIA, Java; Floral record: Canna flavescens; Cockerell 1905: 220 (comparativenotes, synonymy, uncertain identity (as canifrons)); AUSTRALIA, Adelaide River (misidentification);SRI LANKA (misidentification, *iridipennis group); Bingham 1905: 59 (citation, distribution, nest(house-post in village)); Brown 1906: 686 (distribution); PHILIPPINES; Schulz 1907: (nest, predator(Oecophylla smaragdina: Formicidae: Hymenoptera)); Friese 1909("1908"): 354, 355 (distribution,uncertain identity (as praeterita)); INDONESIA, Irian Jaya, Etna Bai; Merauke; Friese 1909("1908"):354, 356 (distribution); INDONESIA, Irian Jaya, Manikion; Wendesi; Friese 1909: 272, 275 (key to spe-cies, nest (in structural wood)); INDONESIA, Ost-Java; Sumatra; Schulz 1909: (nest (as canifrons inpart)); Friese 1912: 94 (distribution); PAPUA NEW GUINEA, Sepik-Bivak; Ihering 1912: 3 (citation (aslaevicops!)); Strand 1913a: 149-150 (see iridipennis); Friese 1914a: 12-13 (distribution, floral record,uncertain identity (as iridipennis)); INDONESIA, Bandung; Batavia; Buitenzorg; Nongkodjadjar; NusaKambangan; Salatiga; Samarang; Floral record: Canna flavescens; Friese 1915: 4 (distribution, uncertainidentity (as praeterita)); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Etna Bai; Jasa; Merauke;Njao; Zoutbron; Cockerell 1918: 385 (comparative notes); Cockerell 1919c: 77, 79 (distribution, previ-ous misidentifications); Cockerell 1919a: 121 (comparative note, uncertain identity (as iridipennis));INDONESIA, Java; Cockerell 1922a: 279 (distribution); Cockerell 1922b: (comparative note); Cocker-ell 1923: 241 (key to species); INDONESIA, Aru; Cockerell 1925a: 492 (comparative note); Cockerell1925b: (misidentification (was a Diptera)); Cockerell 1926b; Salt 1929: 460 (citation); Cockerell 1929c:590, 592 (comparative note); Cockerell 1929e: 242, 243 (comparative note); Cockerell 1929d: 300 (com-parative note); Cockerell 1930b: 150 (comparative note (previously as canifrons)); Schwarz 1932: 245,246; Rayment 1932f: 106 (taxonomy); Friese 1933a: 46 (distribution, uncertain identity (as iridipennis));INDONESIA, Meliau; Rayment 1935: 735, 739 (key to species, taxonomy); Schwarz 1937: 307, 308,314, 315; Schwarz 1939a: 86, 88, 111, 112 (distribution, key to species, taxonomy (most of the specimensexamined by Schwarz and identified as iridipennis are actually laeviceps, see iridipennis)); Floral record:Typha angustifolia; Schwarz 1945: 141 (citation); Schwarz 1948: 23, 84, 141; Nogueira-Neto 1949: 121(synonymy); Sakagami 1959: 120 (comparative note); Sakagami 1960: 146 (comparative note, distribu-



tion, uncertain identity (as iridipennis)); CAMBODIA, Stung Treng; Moure 1961: 208 (morphology, sys-tematic position); Michener 1961: 3, 10, 28-33, 35, 38, 39, 42 (nest, uncertain identity (as iridipennis));PAPUA NEW GUINEA, Bisianumu, Sogeri Plateau; Bubia, vicinity of Lae; Kapagere, near Rigo;Koitakinumu Estate, Sogeri Plateau; Port Moresby; Michener 1965: 231 (citation); Baltazar 1966: 407(distribution, uncertain identity (as iridipennis)); PHILIPPINES, Mindanao, Lanao, Zamboanga; Pala-wan; Yoshikawa et al. 1969: 168, 169, 173 (distribution, nest, uncertain identity (as iridipennis));MALAYSIA, Fraser's Hill, Selagor-Pahang; Kuching; Bukit-Matajam, Prov. Wellesley; Zeuner & Man-ning 1976: 222, 223, 224 (synonymy); Wille 1977: 45 (synonymy); Sakagami 1978: 166-194, 196, 197,198, 200-208*, 209, 210, 211, 212, 213, 215, 216, 218, 219, 220, 221, 222, 226, 228, 229, 230, 234, 235,236, 237, 238, 239, 240, 247, plate 5 (nest, taxonomy); LAOS; MALAYSIA; SINGAPORE; THAI-LAND; VIETNAM; Sakagami & Ito 1981: 368, 373 (morphology); Sakagami et al. 1983b: 2, 6, 8, 10,13, 20 (nest); Sakagami et al. 1983a: (see minangkabau); INDONESIA, Sumatera Barat at Lubuk; Saka-gami et al. 1983d: (see minangkabau); Sakagami et al. 1983c: (reproduction); Inoue et al. 1984b: (seeminangkabau); Sakagami & Yamane 1984: 37, 48; Inoue et al. 1984a: (see minangkabau); Salmah et al.1984: (see minangkabau); Sakagami & Inoue 1985: 175, 176, 177, 180, 181, 182, 184, 186 (comparativenote); Sakagami et al. 1985: (distribution, key to species); Utami 1986: (pollination, uncertain identity(as irridipenis!)); INDONESIA, West Java, Pandeglang; Floral record: Metroxylon sagu; Sakagami &Khoo 1987: 208, 209, 210, 211, 212 (morphology); Sakagami & Inoue 1987: 611, 624 (comparativenote); Starr & Sakagami 1987: 97 (citation); Salmah et al. 1987: 384 (synonymy); Uji 1987: (pollina-tion, uncertain identity (as irridipenis!)); INDONESIA, E Kalimantan; Floral record: Nephelium lappa-ceum; Camargo 1988: 372 (distribution, synonymy); MALAYSIA, Sandakan, Penang; Sakagami &Inoue 1989: 615 (comparative note); Roubik 1989: 181, 205; Sakagami et al. 1989: 272 (misidentifica-tion); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDIA (misidentification);INDONESIA, Java; Sumatra; MALAYSIA; MYANMAR; THAILAND; Sakagami & Inoue 1990: (ovi-position); Salmah et al. 1990: (distribution, nest (hollow pillars; cracks), also as laeviceps small); INDO-NESIA, Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Johnson& Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera)); Engels & Imperatriz-Fonseca1990: (see minangkabau); Salmah et al. 1991: 196 (synonymy); Prasad & Hemanth 1992: (predator(Ardeola grayii: Ardeidae: Ciconiiformes) (as leviseps!)); Michener & Roubik 1993: 260 (comparativenote, morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cardale 1993: 323-324 (see sapi-ens); Dollin & Dollin 1994: (collecting males); Yamane et al. 1995: 276, 293 (comparative note);Drumond et al. 1995: 44 (citation, comparative behavior); Imperatriz-Fonseca & Zucchi 1995: 239(citation); Inoue et al. 1995: 98 (pollination); Floral record: Neuwiedia borneensis; Inoue et al. 1996b:643 (misidentification); Salmah et al. 1996: 402, 410 (synonymy); Roubik 1996: 61 (distribution); BRU-NEI, Kuala Belalong; Ikudome & Kusigemati 1996: 20 (taxonomy); Smiley et al. 1996: 198, 200(inquiline (Hypoaspis hoffmannae: Laelapidae: Acari), uncertain identity (as iridipennis)); MALAYSIA,Selangor; Okada et al. 1996: 178 (pollination); Floral record: Neuwiedia veratrifolia; Nogueira-Neto1997: 94 (citation); Dollin et al. 1997: (revision, synonymy, taxonomy); Nagamitsu & Inoue 1997: 435,436 (pollination); Floral record: Santiria laevigata; Dollin 1997: (compared to sapiens); Nagamitsu &Inoue 1998: (foraging, morphology); Liow 1999: 106, 116, 127 (nest); Nagamitsu et al. 1999: (floralrecords (not by species), foraging); Peters et al. 1999: 382 (citation); Crane 1999: 89; van Veen & Som-meijer 2000: 70, 74, 75 (citation); Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribu-tion, nest); Wu 2000: 387, 391-392, plate (key to species, morphology); Zherikhin & Ross 2000: 6, 13-14 (amber); Chinajariyawong & Saiboon 2000: (distribution, nest); THAILAND, Songkhla; Liow et al.2001: 184, 190; Green et al. 2001: 91 (microsatellite); Peeters & Ito 2001: 610-611 (citation);Wijesekara 2001: 154 (uncertain distribution); Kocyan & Endress 2001: 848, 856, 861, 864 (pollina-tion); Floral record: Neuwiedia veratrifolia; Eltz et al. 2002a: 30 (distribution, foraging, nest, population);



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); Hohmann2003: (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 (floralrecord); Floral record: Hevea brasiliensis; Roubik 2006: 134 (nest); Tangmitcharoen et al. 2006: (polli-nation); Floral record: Tectona grandis; Inson & Malaipan 2006: (distribution, foraging, nest, uncertainidentity (as iridipennis)); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007:(systematics); Leonhardt et al. 2007: (foraging); Floral record: Ardisia pyramidalis; Diospyros durion-oides; 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 taxonis labeled "Bandjarmas" (?=Bandjarmasin, near Liangtelan) (F. Penati, pers. com.) (taxonomy); Typelocality: MALAYSIA "Liangtelan (Borneo)" (1 worker); Dalla Torre 1896: 580 (citation); Cockerell1919b: 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 (systematicposition); 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; Neuwiediaborneensis; 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: (floralrecords (not by species), foraging); Eltz et al. 2001a: (pollen foraging); Eltz et al. 2001b: 343, 345 (forag-ing); 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; Floralrecord: 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 (abun-dance); Michener & Boongird 2004: 144, 145, 146 (comparative note); Samejima et al. 2004: (popula-tion); 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); Typelocality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker); Schwarz 1937: 315, 326; Schwarz1939a: 120 (synonymy); Moure 1961: 210 (synonymy); Michener & Boongird 2004: 146 (comparativenote).

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 carryno locality label (F. Penati, pers. com.) (taxonomy); Type locality: MALAYSIA "Liangtelan (Borneo)" (2



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, tax-onomy); 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); Sak-agami & Inoue 1985: 181 (comparative note); Sakagami et al. 1985: (distribution, key to species); Kiew1985: (luring (unwashed shorts)); Starr & Sakagami 1987: 97 (citation); Camargo 1988: 372 (distribu-tion); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1990: (distribution, key to species); INDONE-SIA, Sumatra; MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest);INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Michener & Roubik 1993: 260 (com-parative note, morphology); Salmah et al. 1996: 410 (comparative note); Roubik 1996: 61, 62, 63 (distri-bution, nest); BRUNEI, Kuala Belalong; Dollin et al. 1997: 886 (comparative note); Nagamitsu & Inoue1997: (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 (forag-ing); 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: Rhizophoraapiculata; Nagamitsu & Inoue 2002: 305, 307, 308, 310, 312 (foraging); Eltz et al. 2003: (nest, treepreference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz 2004: 322 (abundance); Samejima etal. 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 specimensmatch the original description and the taxon is not known from southern Australia (key to species, taxon-omy); 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; WesternAustralia; Dollin 1997: (characteristics); Crane 1999: 89; Heard & Dollin 2000: (bee-keeping); Green etal. 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:



Nomen nudum. The name is unavailable for taxonomic purposes, as it must be deemed to be infrasubspe-cific (ICZN, art. 15.2 and 45.6.3.) (taxonomy); Sakagami et al. 1990: (distribution, key to species, varia-tion); Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera Barat; Inoue et al. 1990b:(population); Tadauchi et al. 1998: 244 (type depositories); Nagamitsu & Inoue 2005: 77 (distribution,foraging).

Trigona (Tetragonula) minangkabau Sakagami & Inoue 1985: 175, 176, 177, 178, 179, 180, 181, 184-186*,187-188: Holotype (MZB, Hymn.0198, worker); paratypes (MZB (Hymn.0195-0197)) (key to species,nest, taxonomy); Type locality: INDONESIA "Lubuk Mintrun nr. Padang, Sumatera Barat, Indonesia, ix1981 in mass flight, S.F. Skagami" (1 worker, holotype); "Lubuk Mintrun" (many workers, 25 males,paratypes); Sakagami et al. 1983a: (nest, taxonomy (misidentification; as laeviceps)); INDONESIA,Sumatera Barat at Lubuk; Sakagami et al. 1983d: (oviposition behavior (misidentification; as laeviceps));Inoue et al. 1984b: (brood, nest, reproduction (as laeviceps)); Inoue et al. 1984a: (physiology, reproduc-tion (as laeviceps)); Salmah et al. 1984: (development (as laeviceps)); Sakagami et al. 1985: (distribu-tion, key to species); Inoue et al. 1985a: (aggressiveness, foraging, nest); Inoue et al. 1985b: (foraging);Sakagami & Inoue 1987: 626 (comparative note); Starr & Sakagami 1987: 97, 106 (citation); Salmahet al. 1987: 383, 388, 389, 390 (development); Roubik 1989: 86, 133, 137; Sakagami et al. 1989: 271,272 (comparative note); Bego et al. 1989: 48, 53 (foraging, greenhouse, introduction); Sakagami 1989:164; Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; Sakagami & Inoue1990: 201, 202, 203, 205, 210, 211, 212, 215 (comparative note); Salmah et al. 1990: (distribution, nest(hollow rambutan branches (Nephelium lappaceum: Sapindaceae); bamboo; interspaces)); INDONESIA,Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Suka et al. 1990:(nestmate recognition); Roubik 1990a: 115 (citation); Engels & Imperatriz-Fonseca 1990: 186, 207(citation (as laeviceps)); Inoue et al. 1990a: 229; Salmah et al. 1991: (reproduction); Kakutani et al.1993: (pollination); Floral record: Fragaria; Suka & Inoue 1993: (behavior); Inoue et al. 1993: (nest(hollow rambutan branches (Nephelium lappaceum: Sapindaceae))); INDONESIA, Sumatra; Hoshiba &Imai 1993: 469, 477, 481, 487 (chromosome); Yamane et al. 1995: 276, 282, 293 (comparative note);Roubik 1995: 151 (antropogenic nest); Inoue et al. 1995: 98 (pollination); Floral record: Neuwiediaborneensis; Inoue et al. 1996b: (development, nest, variation); Salmah et al. 1996: 402, 407, 408, 409,410 (comparative note); Inoue et al. 1996a: (population); Tadauchi et al. 1998: 243-244 (type deposito-ries); Inoue et al. 1999: 208, 217 (citation, comparative table); Ranger & O'Donnell 1999: 189 (compar-ative note); Heard 1999a: 187, 193 (citation); Breed et al. 1999: 772-773 (citation, foraging); van Veen& Sommeijer 2000: 70, 74, 75 (citation (as laeviceps)); Amano et al. 2000: (citation); Nagamitsu &Inoue 2002: 309 (citation); San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh For-est Reserve; Rocha et al. 2003: 317 (citation); Tóth et al. 2003: 366 (citation); Costa et al. 2004: 187(citation); Tóth et al. 2004: 5, 6 (citation); Tsuji et al. 2004: (predator (Oecophylla smaragdina: Formi-cidae: Hymenoptera)); Nagamitsu & Inoue 2005: 77, 78 (distribution, foraging); Hofstede & Som-meijer 2006: 388 (citation); Cepeda 2006: 176, 184 (citation); Cortopassi-Laurino et al. 2006: 278(citation); Slaa et al. 2006: 297, 305, 306 (citation); Leonhardt et al. 2007: 126 (citation).

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 work-ers); Sakagami & Inoue 1985: 186 (comparative note); Sakagami et al. 1985: (distribution); Sakagamiet 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); Floralrecord: Homalium grandiflorum; Tectona grandis; Rasmussen & Cameron 2007: (systematics).



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 SriTamarat, July 5, 1928 (Hugh M. Smith)" (workers, males); "Singora, June 1929 (Hugh M. Smith)" (work-ers); Krombein 1951: 295 (comparative note); Schwarz 1951: 63 (distribution); THAILAND, Bangkok;Moure 1961: 208-9 (systematic position); Sakagami & Yoshikawa 1961: 435, 436-437 (comparativenote, 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: 166-194, 200, 211, 212, 213, 218, 219-223*, 224, 225, 226, 227, 228, 234, 236, 237, 238, 239, 240, 247, plate5 (taxonomy); Sakagami et al. 1983b: 2, 6-7, 8, 20 (nest); Sakagami et al. 1983a: 103, 105, 106 (compar-ative note); Sakagami et al. 1983d: (oviposition behavior); Sakagami & Inoue 1985: 177 (comparativenote); 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 (comparativenote, distribution); Sakagami & Inoue 1990: 201, 202, 203, 204, 205, 206, 210, 211, 212, 215 (compara-tive 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 (com-parative note, distribution); MALAYSIA; THAILAND; Wu 2000: 387, 390-391 (key to species, mor-phology); Franck et al. 2004: (genetics, microsatellite); Nagamitsu & Inoue 2005: 77 (distribution,foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics); Thumma-jitsakul 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 local-ity: 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" (1worker); "Taman Negara" (16 workers); "Kanching" (1 worker); "Penang" (8 worker); "Pasoh" (12 work-ers); "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 (comparativenote); 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 ForestReserve; 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); Typelocality: 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!)).



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, holo-type); "Malaya, Kepong" (1 worker); "Ulu Langat" (20 workers); "Gombak" (2 workers); "CameronHighland" (4 workers); "Thailand, Khaophappha-Khaochong" (workers); Sakagami et al. 1985: (distri-bution, 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"; para-lectotypes (DEI (1), USNM (1), ZMHB (7)) (taxonomy); Type locality: MALAYSIA "Taiping Hills(Upp. Perak) auf Malakka, im Februar 1912" (several workers); Cockerell 1926a: 224 (comparativenote); Schwarz 1939a: 85, 93, 121* (distribution, key to species, taxonomy); Sakagami et al. 1985: (dis-tribution); Sakagami & Khoo 1987: (morphology); Michener 1990: 120 (systematic position); Sakag-ami 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); Nag-amitsu 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, North-west 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-Neto1951: (distribution); INDIA, Uttar Pradesh; Moure 1961: 208 (synonymy); Sakagami 1978: 216 (synon-ymy); Wu 2000: 392 (synonymy).

Melipona smithii Bingham 1897: 560, 563: Unnecessary replacement name for Trigona ruficornis Smith, necLamarck. Lamarck 1817 (and repeated in 1835) only listed Melipona favosa, M. amalthea, M. ruficrus, M.postica, and M. pallida. The replacement name therefore remains enigmatic, unless mistaken for M. rufi-crus Latreille. BMNH has a labeled type for this replacement name (17b.1129) (distribution, key to spe-cies, taxonomy); INDIA, Mainpuri, NW provinces; MYANMAR, Tenasserim; Bingham 1905: 59(citation, distribution); Schwarz 1939a: 88, 107, 108 (synonymy); Schwarz 1948: 14, 23, 56; Nogueira-



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 local-ity: SOLOMON ISLANDS "Solomon Islands, July-August, 1909 (Froggatt)" (worker); Cockerell1929e: 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); Krombein1951: 295 (synonymy); Moure 1961: 209 (systematic position); Michener 1965: 231 (citation); Sakag-ami 1978: 221, 222; Dollin & Dollin 1983: (nest (as laeviceps)); Sakagami & Inoue 1987: 626 (compar-ative 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; AdelaideRiver; Cape York; Gordonvale; Thursday Is.; Oliveira et al. 1995: 20 (citation); Snelling 1996: 131 (dis-tribution); 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 & Cam-eron 2007: (systematics); Klumpp 2007: 4, 82; Wallace et al. 2008: (dispersal, pollination); Floralrecord: 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); Schwarz1939a: 85, 92, 106* (distribution, key to species, taxonomy); Schwarz 1948: 118; Sakagami 1959: 119-120 (distribution, morphology, variation); SINGAPORE; Moure 1961: 210 (morphology, systematicposition); Yoshikawa et al. 1969: 168, 169 (nest); Wille & Michener 1973: 14, 25, 59 (nest); Sakagami1978: 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 & Sakagami1987: 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, Thai-land (3), SEMK (11) (taxonomy); Type locality: THAILAND "Thailand, on the Isthmus of Kra: RanongProvince: Muang district, 9–13 November 2002" (11 workers); "Ngow Waterfall National Park" (7 work-



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 (morphol-ogy, systematic position); Sakagami 1978: 200 (as testaceicornis!); Sakagami et al. 1990: 129 (synon-ymy (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 (com-parative note, floral record, taxonomy); CAMBODIA, Angkor; Nan; Pah Meeung; Floral record: Poinset-tia; 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 (distribu-tion, variation); MICRONESIA, Caroline Islands, Toloas; Truk; Cockerell 1939a: 61, 62, 64 (distribution("presumably introduced by man"), variation); MICRONESIA, Dublon Island; Tarik Island; Schwarz1948: 118, 233; Krombein 1950: 102, 133, 135 (distribution, morphology); MICRONESIA, CarolineIslands; Schwarz 1951: 63 (distribution); CAMBODIA, Angkor; Mt. Suept; Sakagami 1960: 146 (com-parative note); Moure 1961: 208 (morphology, systematic position); Michener 1961: 28 (comparativenote); 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 etal. 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" (1worker); "Templer Park" (17 workers); Yoshikawa et al. 1969: 173 (citation (as iridipennis)); Sakagamiet al. 1985: (distribution); Sakagami et al. 1990: 131 (distribution); Tadauchi et al. 1998: 244 (typedepositories); 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: (tax-onomy); Roubik 2006: 126, 127, 130, 132 (nest).

Tetrigona

Tetrigona apicalis (Smith, 1857)



Trigona apicalis Smith 1857: 51: Holotype (OUMNH); additional putative type (BMNH 17b.1188) (taxon-omy); Type locality: MALAYSIA "Borneo (Sarawak)" (worker); Smith 1871: 395 (distribution);MALAYSIA, Borneo; Dalla Torre 1896: 575 (citation); Bingham 1897: 560, 562 (common name (dam-mar bee), distribution, key to species, taxonomy (see also binghami)); MALAYSIA, Borneo; MYAN-MAR, Tenasserim; Waterhouse 1903: 133-134, plate 4 (morphology (male), nest (as collina)); Beccari1904: 162 (nest); Bingham 1905: 59 (citation, distribution); Cockerell 1918: 387 (see binghami); Cock-erell 1920a: 116 (comparative note); Cockerell 1920b: 228 (distribution); MALAYSIA, Sandakan; Cock-erell 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; Schwarz1937: 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); CAM-BODIA, 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 (distribu-tion, nest); MALAYSIA, George Town, Penang; Wille & Michener 1973: 13, 23, 47, 58, 87 (nest); Sak-agami 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 interme-dium; Sakagami et al. 1983b: 2, 5-6, 13, 19, 20 (nest); Inoue et al. 1984a: 136 (comparative note); Saka-gami 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 posi-tion); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Java; Sumatra;MALAYSIA, Borneo; MYANMAR; THAILAND; Salmah et al. 1990: (distribution, nest (hollow trunksof Ficus: Moraceae; Durio zibethinus: Bombacaceae; crevices)); INDONESIA, Sumatera Barat; Inoue etal. 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: San-tiria 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); Michener2000: 801; Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Chinajariya-wong & 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: (distribu-tion, 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; Merre-mia 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



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 MaaHia; 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); Saka-gami 1975: 60, 61 (synonymy); Sakagami et al. 1990: 127 (synonymy).

Trigona sericea Friese 1933a: 45-46: Lectotype (ZMHB, worker): here designated, "Borneo / Sanggau / 24-7-32", "Trigona / sericea / 1925 Friese det. / Fr."; paralectotypes (ZMHB (5), DEI (4)) (taxonomy); Typelocality: 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); Moure1961: 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 api-calis)); MALAYSIA, Sandakan; Schwarz 1939a: 85, 94, 126, 128-129* (distribution, key to species, tax-onomy); 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, treepreference); 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; Rasmus-sen & Cameron 2007: (systematics); Leonhardt et al. 2007: (foraging); Floral record: Archidendron jir-inga; 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 synonymonof 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, sys-tematic position); Sakagami 1975: 61, 62-64* (distribution, morphology, variation); LAOS; THAI-LAND; 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: (for-aging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Rajitparinya2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Nagamitsu & Inoue 2005: 76, 83 (dis-tribution, foraging); Klakasikorn et al. 2005: 6 (citation); Inson & Malaipan 2006: (distribution, forag-ing, 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-



tion); Type locality: MALAYSIA "Perak, F.M.S., Batang Padang, Jor Camp, 1800 ft., June 4, 1923(Pendlebury)"; CAMBODIA, Patalung; MALAYSIA, Kualu Lumpur, Gombak valley; Schwarz 1937:302; Schwarz 1939a: 85, 94, 102, 127, 129-130* (distribution, key to species, taxonomy); Schwarz 1948:117; Moure 1961: 216 (morphology, systematic position); Sakagami 1975: 61, 62, 63, 64* (distribution,morphology, variation); MALAYSIA; THAILAND; Sakagami et al. 1985: (distribution, key to species);Michener 1990: 127 (systematic position); Sakagami et al. 1990: 131, 136 (distribution, key to species);San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu &Inoue 2005: 76 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation).

Tetrigona vidua (Lepeletier de Saint Fargeau, 1836)

Melipona vidua Lepeletier de Saint Fargeau 1836: 429: Syntype (MNHN, 1 worker): putative syntypeexamined, "Museum Paris / Bengale / Diard & Duvaucel 1815", "vidua", "Diard et Duvaucel", "type","TYPE", "M. Vidua / Lep. S. Farg / Bengale". I consider this a true type: Alfred Duvaucel (1793-1825)and Pierre-Médard Diard (1794-1863) arrived on their first expedition to Kolkata, India, in January 1818and spent years collecting in the Indo-Malayan region. The label date is wrong as they were still in Francein 1815, but did reside for long periods in India later (Claeys: 1954) and I consider the label provenancemerely the port of embarkation of the specimen, rather than the collecting locality. The type is similar tomelanoleuca and could be considered a senior synonymon of that. However, melanoleuca is a variabletaxon, and is here retained until further studies may document if in fact it is only a single taxon (taxon-omy); Type locality: INDONESIA/E TIMOR "Ile de Timor" (unknown); Spinola 1853: 94 (citation);Smith 1854: 413 (citation); Smith 1871: 395 (distribution); INDONESIA/EAST TIMOR, Timor; Castets1893: 481 (citation); Dalla Torre 1896: 585 (citation); Bingham 1897: 560, 561 (distribution, key to spe-cies, taxonomy, uncertain identity); INDIA (misidentification); INDONESIA; MALAYSIA; MYAN-MAR, Tenasserim; Haddon 1901: 87 (common name (manyi), bad omen for Ibans ("if a swarm of beessettled underneath a house that had recently been built it would be considered a bad sign"), uncertain iden-tity); Bingham 1905: 59 (citation, distribution); Friese 1914a: 13 (distribution (maybe itama?)); INDO-NESIA, Gunung Ungaran; Friese 1918: 519 (comparative note (maybe itama?)); Cockerell 1918: 387(comparative notes); Cockerell 1926a: 224 (comparative note); Salt 1929: 454 (citation); Schwarz 1937:299-300; Schwarz 1939a: 86, 88, 98, 129 (synonymy, taxonomic position); Schwarz 1948: 422; Moure1961: 217 (morphology, systematic position); Sakagami & Khoo 1987: 207 (comparative note); Cama-rgo 1988: 372 (distribution); INDONESIA, Sumatra; Wu 2000: 386, 390, plate (key to species, morphol-ogy).

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 (keyto species, taxonomy (as Melipona)); Waterhouse 1900: (nest); Cockerell 1920a: 116 (distribution);Cockerell 1930b: 149; Rayment 1932a: (morphology, nest); Bibby 1947: 78 (distribution); Michener1961: (common name (native bee; sweat bee), nest); Michener 1965: 231 (citation); Michener 1965:230; Smythe 1970: (pollination (as Trigona)); Floral record: Cymbidium madidum; Dendrobium; Frith1978: (nest (in Rhizophora apiculata: Rhizophoraceae); predator (Oecophylla smaragdina: Formicidae:Hymenoptera)); Schneider 1982: (pollination); Appanah 1982: (pollination); MALAYSIA, Negri Sem-biran, 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);



Raju & Reddi 1989: (pollination); Floral record: Anisomeles indica; Anisomeles malabarica; Heard etal. 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 & Law-son 1993: (pollination (as Trigona)); Floral record: Caladenia carnea; Cymbidium madidum; Cymbidiumsuave; 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); Viraktamathet al. 1999: (pollen); Heard 1999b: (pollination (as spp.)); Floral record: Mangifera indica; Yumoto2000: (pollination); Heard 2000: (pollination (as spp.)); Jagadish et al. 2002: (common name (dammerbee), 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 seri-cea; 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 basimaculataBingham 1905: 60, plate A11 (citation).

Trigona pallidicincta Cockerell 1918: 386: Holotype (BMNH 17b.1134, male) (key to species, comparativenotes, taxonomy); Type locality: SINGAPORE "Singapore (Baker)" (male).

Nomina nuda

Melipona pulla Illiger 1806: 158: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 12.1) (citation); INDONESIA, Sumatra;Melipona pulla Dalla Torre 1896: 582 (citation).

Melipona Khasiana Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 13.1.1) (common name (nagp hamang),honey, nest).

Melipona terrestris Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as nodescription or definition of the taxon was provided (ICZN, art. 13.1.1) (common name (nagp khyndew),honey, nest).

Trigona (Pacificotrigona) okazawai Tadauchi et al. 1998: 245: Nomen nudum (as Trigona (Pacificotrigona)okazawai irianensis). The name is unavailable for taxonomic purposes, as no description or definition ofthe taxon was provided (ICZN, art. 15.1).

Trigona (Pacificotrigona) okazawai irianensis Tadauchi et al. 1998: 245: Nomen nudum. The name is



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)



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)



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)



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)



Appendix 2. Stingless bee taxon name followed by current combination or synonymy

aliceae, Homotrigona aliceae (Cockerell, 1929) ambusta, see thoracicaanamitica, Homotrigona anamitica (Friese, 1909) angophorae, see carbonariaapicalis, Tetrigona apicalis (Smith, 1857) arcifera, Lepidotrigona arcifera (Cockerell, 1929) atomella, see fuscobalteataatricornis, 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 biroiborneensis, see lacteifasciatabreviceps, see itamabrevis, nomen nudum, see fuscobalteatabusara, see canifronscacciae, Lisotrigona cacciae (Nurse, 1907) cambodiensis, see collinacanifrons, Lophotrigona canifrons (Smith, 1857) carbonaria, Tetragonula carbonaria (Smith, 1854) carpenteri, Lisotrigona carpenteri Engel, 2000 cassiae, Austroplebeia cassiae (Cockerell, 1910) castanea, see fimbriatacincta, Austroplebeia cincta (Mocsáry in Friese, 1898) clypearis, Tetragonula clypearis (Friese, 1909) cockerelli, Austroplebeia cockerelli (Rayment, 1930) collina, Tetragonilla collina (Smith, 1857) confusella, see geissleridapitanensis, Tetragonula dapitanensis (Cockerell, 1925) darek, nomen nudum, see minangkabaudavenporti, Tetragonula davenporti (Franck, 2004) devicta, Tetragonula devicta (Cockerell, 1921) doipaensis, Lepidotrigona doipaensis (Schwarz, 1939) drescheri, Tetragonula drescheri (Schwarz, 1939) dulitae, see haematopteraerythrogastra, Heterotrigona erythrogastra (Cameron, 1902) erythrostoma, see fuscobalteataessingtoni, Austroplebeia essingtoni (Cockerell, 1905) ferrea, see luteafimbriata, Homotrigona fimbriata (Smith, 1857) flavibasis, Lepidotrigona flavibasis (Cockerell, 1929) flavistigma, see fimbriataflaviventris, Platytrigona flaviventris (Friese, 1909) fulvomarginata, see terminatafulvopilosella, see nitidiventrisfurva, 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)



hemileuca, see apicalishirashimai, Tetragonula hirashimai (Sakagami, 1978) hobbyi, Platytrigona hobbyi (Schwarz, 1937) hockingsi, Tetragonula hockingsi (Cockerell, 1929) hoosana, see hoozanahoozana, 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 pendleburyikusutkana, see aliceaelacteifasciata, Geniotrigona lacteifasciata (Cameron, 1902) laeviceps, Tetragonula laeviceps (Smith, 1857) lamingtonia, Platytrigona lamingtonia (Cockerell, 1929) latebalteata, Lepidotrigona latebalteata (Cameron, 1902) latigenalis, see reepenilatipes, Lepidotrigona latipes (Friese, 1900) lieftincki, Sundatrigona lieftincki (Sakagami & Inoue, 1987) lutea, Homotrigona lutea (Bingham, 1897) luteiventris, see erythrogastramatsumurai, see mooreimelanocephala, Tetragonula melanocephala (Gribodo, 1893) melanoleuca, Tetrigona melanoleuca (Cockerell, 1929) melanotricha, see anamiticamelina, 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 fuscobalteatapenangensis, Tetragonula penangensis (Cockerell, 1919) pendleburyi, Pariotrigona pendleburyi (Schwarz, 1939) peninsularis, Tetrigona peninsularis (Cockerell, 1927) percincta, Austroplebeia percinta (Cockerell, 1929) pfeifferi, see fuscobalteataplanifrons, Platytrigona planifrons (Smith, 1865) praeterita, Tetragonula praeterita (Walker, 1860) pulla, nomen nudum pygmaea, see fuscobalteatareepeni, Tetragonula reepeni (Friese, 1918) rufibasalis, Tetragonilla rufibasalis (Cockerell, 1918) ruficornis, Tetragonula ruficornis (Smith in Horne, 1870) sandacana, see erythrogastrasapiens, Tetragonula sapiens (Cockerell, 1911) sarawakensis, Tetragonula sarawakensis (Schwarz, 1937) scintillans, see cacciae



sericea, see apicalissirindhornae, Tetragonula sirindhornae (Michener & Boongird, 2004) smithii, see ruficornissymei, Austroplebeia symei (Rayment, 1932) terminata, Lepidotrigona terminata (Smith, 1878) terrestris, nomen nudum testaceinerva, see melanocephalatestaceitarsis, Tetragonula testaceitarsis (Cameron, 1901) thoracica, Geniotrigona thoracica (Smith, 1857) trochanterica, Lepidotrigona trochanterica (Cockerell, 1920) tunneli, nomen nudum, see Lepidotrigona valdezi, Tetragonula valdezi (Cockerell, 1918) ventralis, Lepidotrigona ventralis (Smith, 1857) versicolor, see fimbriatavidua, Tetrigona vidua (Lepeletier de Saint Fargeau, 1836) wallacei, nomen nudum, see sapienswebsteri, Austroplebeia websteri (Rayment, 1932) wybenica, see clypeariszucchii, Tetragonula zucchii (Sakagami, 1978)

Appendix 3. Museum acronyms. Curators and sources of information concerning the collection are listed in parenthesis

AMNH USA, New York, New York, American Museum of Natural History (Jerome G.Rozen, John S. Ascher)

AMS Australia, New South Wales, Sydney, Australian Museum (David Britton, MaxBeatson)

ANIC Australia, Australian Capital Territory, Canberra City, CSIRO, AustralianNational Insect Collection (John La Salle, Nicole Fisher)

BMNH United Kingdom, London, The Natural History Museum (David Notton)BPBM USA, Hawaii, Honolulu, Bernice P. Bishop Museum (Shepherd P. Myers)Calicut India, Kerala, Calicut University, Zoology DepartmentDEI Germany, Müncheberg, Deutsches Entomologisches Institut im ZALF (Holger

H. Dathe, Stephan Blank)EKYU Japan, Fukuoka, Kyushu University, Entomological laboratory (Tadauchi et al.

1998, website)HNHM Hungary, Budapest, Hungarian Natural History Museum (Sándor Csõsz)IEBR Vietnam, Hanoi, Institute of Ecology and Biological ResourcesMNHN France, Paris, Muséum National d'Histoire Naturelle (Claire Villemant)MSNG (Collezione Gribodo)Italy, Genova, Museo Civico di Storia Naturale "Giacomo Doria" (Fabio Penati)MVMA Australia, Victoria, Abbotsford, Museum of Victoria (Ken Walker)MZB Indonesia, Bogor, Museum Zoologicum Bogoriense (Yayuk R. Suhardjono)OUMNH (=Wilson Saunders collection)United Kingdom, Oxford, University Museum of Natural History

(James Hogan)QM Australia, Queensland, South Brisbane, Queensland MuseumRMNH Netherlands, Leiden, Nationaal Natuurhistorische Museum (“Naturalis”) (Cor-

nelius van Achterberg)SEHU Japan, Sapporo, Hokkaido University Museum, Systematic entomology (Masa-

hiro Ohara)SEMC USA, Kansas, Lawrence, University of Kansas, Snow Entomological Museum



SFS Shôichi F. Sakagami’s collection (see SEHU)SMNS Germany, Stuttgart, Staatliches Museum für NaturkundeUSNM USA, Washington D.C., National Museum of Natural History, (formerly, United

States National Museum) (David Furth, Brian Harris)ZMHB Germany, Berlin, Museum für Naturkunde der Humboldt-Universität (Frank

Koch)ZRC Singapore, National University of Singapore, Raffles Museum of Biodiversity

Research, Zoological Reference Collection (Lua Hui Kheng)

References

Adams, P.B., Bartareau, T., & Walker, K.L. (1992) Pollination of Australian orchids by Trigona (Tetragona) Jurine bees(Hymenoptera: Apidae). Australian Entomological Magazine, 19, 97–101.

Adams, P.B. & Lawson, S.B. (1993) Pollination in Australian orchids: a critical assessment of the literature 1882–1992.Australian Journal of Botany, 41, 553–575.

Adams, P.B. & Lawson, S.D. (1988) Multiple bee pollinators of Dendrobium kingianum Bidw. in the natural habitat. TheOrchadian (Australasian Native Orchid Society), 9, 103, 105–107.

Aluri, R.J.S. & Subba Reddi, C. (1996) Floral biology and pollination in Gliricidia sepium (Fabaceae). Journal of NatureConservation, 8, 65–67.

Amano, K. (2004). Attempts to introduce stingless bees for the pollination of crops under greenhouse conditions inJapan. Food and Fertilizer Technology Center, Taipei.

Amano, K., Nemoto, T., & Heard, T.A. (2000) What are stingless bees, and why and how to use them as crop pollinators?– a review –. Japan Agricultural Research Quarterly, 34, 183–190.

Anderson, D.L., Sedgley, M., Short, J.R.T., & Allwood, A.J. (1982) Insect pollination of mango in Northern Australia.Australian Journal of Agricultural Research, 33, 541–548.

Anderson, J. (1890) The Selungs of the Mergui archipelago. Trübner & Co., London, 47.Appanah, S. (1982) Pollination of androdioecious Xerospermum intermedium Radlk. (Sapindaceae) in a rain forest. Bio-

logical Journal of the Linnean Society, 18, 11–34.Appanah, S. (1993) Mass flowering of dipterocarp forests in the aseasonal tropics. Journal of Bioscience, 18, 457–474.Appanah, S., Willemstein, S.C., & Marshall, A.G. (1986) Pollen foraging by two Trigona colonies in a Malaysian rain

forest. Malayan Nature Journal, 39, 177–191.Arias, M.C., Brito, R.M., Francisco, F.d.O., Moretto, G., Oliveira, F.F.d., Silvestre, D., & Sheppard, W.S. (2006) Molecu-

lar markers as a tool for population and evolutionary studies of stingless bees. Apidologie, 37, 259–274.Ashmead, W.H. (1905) Additions to the recorded hymenopterous fauna of the Philippine islands, with descriptions of

new species. Proceedings of the United States National Museum, 28, 957–971.Atluri, J.B., Rao, S.P., & Reddi, C.S. (2000) Pollination ecology of Helicteres isora Linn. (Sterculiaceae). Current Sci-

ence (India), 78, 713–718.Atmowidjojo, A.H. & Adisoemarto, S. (1986) Potential pollen–transferring insects of Indigofera spp. Treubia, 29, 225–

235.Bacelar–Lima, C.G., Freire, D.d.C.B., Coletto–Silva, A., Costa, K.B., Laray, J.P.B., Vilas–Boas, H.C., & Carvalho–Zilse,

G.A. (2006) Melitocoria de Zygia racemosa (Ducke) Barneby & Grimes por Melipona seminigra merrillae Cocker-ell, 1919 y Melipona compressipes manaosensis Schwarz, 1932 (Hymenoptera, Meliponina) en la Amazonía Cen-tral, Brasil. Acta Amazonica, 36, 343–348.

Baker, D.B. (1993) The type material of the nominal species of exotic bees described by Frederick Smith (Hymenoptera,Apoidea). Oxford University, Oxford, UK, (i)–vi, 1–311, 14 pl.

Baker, E.W. & Delfinado–Baker, M. (1985) An unusual new species of Neocypholaelaps (Acari: Ameroseiidae) from thenests of stingless bees (Apidae: Meliponinae). International Journal of Acarology, 11, 227–232.

Baltazar, C.R. (1966) A catalogue of Philippine Hymenoptera (with a bibliography, 1758–1963). Pacific Insects Mono-graph, 8, 1–488.

Barile, G.E. & Cervancia, C.R. (1995) Foraging and nesting behavior of Trigona biroi Friese (Hymenoptera: Apidae).Philippine Entomologist, 9, 555–566.

Bartareau, T. (1994) The reproductive ecology of Dendrobium toressae (Bailey) Dockr., a geographically restricted spe-cies in north–east Queensland. The Orchadian (Australasian Native Orchid Society), 11, 106–111.

Bartareau, T. (1995) The pollination of Cymbidium madidum Lindley and C. suave R. Br. by native bees – a case of floralreward deception? The Orchadian (Australasian Native Orchid Society), 11, 487–495.



Bartareau, T. (1996) Foraging behaviour of Trigona carbonaria (Hymenoptera: Apidae) at multiple choice feeding sta-tions. Australian Journal of Zoology, 44, 143–153.

Beccari, O. (1904) Wanderings in the great forests of Borneo: Travels and researches of a naturalist in Sarawak [fromitalian "Nelle foreste di Borneo, viaggi e ricerche di un naturalista"]. A. Constable & co., London, xxiv, 423.

Bego, L.R., Maeta, Y., Tezuka, T., & Ishida, K. (1989) Floral preference and flower constancy of a Brazilian stinglessbee, Nannotrigona testaceicornis, kept in a greenhouse (Hymenoptera, Apidae). Bulletin of the Faculty of Agricul-ture (Shimane University), 23, 46–54.

Bibby, F.F. (1947) Notes on the insect fauna of the Samar group, Philippines. Philippine Journal of Science, 77, 61–81.Bichee, S.L. & Sharma, M. (1988) Effect of different modes of pollination in sunflower Helianthus annus L. (Composi-

tae) sown on different dates by Trigona iridipennis Smith. Apiacta (Rome), 23, 65–68.Biesmeijer, J.C. & Slaa, E.J. (2004) Information flow and organization of stingless bee foraging. Apidologie, 35, 143–

157.Bingham, C.T. (1897). Hymenoptera. Wasps and bees. In: W.T. Blandford (Ed.), The fauna of British India, including

Ceylon and Burma. Vol. 1, Taylor & Francis, London, pp. 579.Bingham, C.T. (1903). Diagnoses of aceuleate Hymenoptera. In: N. Annandale & H.C. Robinson (Eds.), Fasciculi

Malayenses, Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States,1901–1902. Zoology. Vol. I (appendix), University Press of Liverpool, London, pp. iii–vii.

Bingham, C.T. (1905). Report on the aceuleate Hymenoptera. In: N. Annandale & H.C. Robinson (Eds.), FasciculiMalayenses, Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States,1901–1902. Zoology. Vol. III, University Press of Liverpool, London, pp. 16–60, plate A.

Blamires, S.J., Hochuli, D.F., & Thompson, M.B. (2008) Why cross the web: decoration spectral properties and preycapture in an orb spider (Argiope keyserlingi) web. Biological Journal of the Linnean Society, 94, 221–229.

Boonithee, A., Juntawong, N., Peckhacker, H., & Hüttinger, E. (1991) Floral visits to select crops by four Apis speciesand Trigona sp. in Thailand. Acta Horticulturae (Wageningen), 288, 74–80.

Borges, F.v.B. & Blochtein, B. (2005) Atividades externas de Melipona marginata obscurior Moure (Hymenoptera, Api-dae), em distintas épocas do ano, em São Francisco de Paula, Rio Grande do Sul, Brasil. Revista Brasileira de Zoo-logia, 22, 680–686.

Bravo, F. (1992) Sistemática e distribução de Parapartamona Schwarz (Hymenoptera, Apidae, Meliponinae). RevistaBrasileira de Entomologia, 36, 863–878.

Breed, M.D., McGlynn, T.P., Sanctuary, M.D., Stocker, E.M., & Cruz, R. (1999) Distribution and abundance of coloniesof selected meliponine species in a Costa Rican tropical wet forest. Journal of Tropical Ecology, 15, 765–777.

Brown, R.E. (1906) A catalogue of Philippine Hymenoptera, with descriptions of new species. Philippine Journal of Sci-ence, 1, 683–695.

Bruce, M.J. & Herberstein, M.E. (2005) Web decoration polymorphism in Argiope Audouin, 1826 (Araneidae) spiders:ontogenetic and interspecific variation. Journal of Natural History, 39, 3833–3845.

Bruijn, L.L.M. & Sommeijer, M.J. (1997) Colony foraging in different species of stingless bees (Apidae, Meliponinae)and the regulation of individual nectar foraging. Insectes Sociaux, 44, 35–47.

Camargo, J.M.F. (1988) Meliponinae (Hymenoptera, Apidae) da coleção do "Instituto di entomologia agraria", Portici,Itália. Revista Brasileira de Entomologia, 32, 351–374.

Camargo, J.M.F., Grimaldi, D.A., & Pedro, S.R.M. (2000) The extinct fauna of stingless bees (Hymenoptera: Apidae:Meliponini) in Dominican amber: two new species and redescription of the male of Proplebeia dominicana (Willeand Chandler). American Museum Novitates, 3293, 1–24.

Camargo, J.M.F. & Moure, J.S. (1994) Meliponinae neotropicais: os gêneros Paratrigona Schwarz, 1938 e AparatrigonaMoure, 1951 (Hymenoptera, Apidae). Arquivos de Zoologia, 32, 33–109.

Camargo, J.M.F. & Moure, J.S. (1996) Meliponini neotropicais: o gênero Geotrigona Moure, 1943 (Apinae, Apidae,Hymenoptera), com especial referência à filogenia e biogeografia. Arquivos de Zoologia, 33, 95–161.

Camargo, J.M.F. & Pedro, S.R.M. (2003a). Sobre as relações filogenéticas de Trichotrigona Camargo & Moure(Hymenoptera, Apidae, Meliponini). In: G.A.R. Melo & I. Alves-dos-Santos (Eds.), Apoidea neotropica: Homena-gem aos 90 anos de Jesus Santiago Moure, Editora UNESC, Criciúma, pp. 109–122.

Camargo, J.M.F. & Pedro, S.R.M. (2003b) Meliponini neotropicais: o gênero Partamona Schwarz, 1939 (Hymenoptera,Apidae, Apinae) — bionomia e biogeografia. Revista Brasileira de Entomologia, 47, 311–372.ß

Camargo, J.M.F. & Pedro, S.R.M. (2007). Meliponini. In: J.S. Moure, D. Urban & G.A.R. Melo (Eds.), Catalogue ofbees (Hymenoptera, Apoidea) in the Neotropical region, Sociedade Brasileira de Entomologia, Curitiba (Paraná),pp. 272–578.

Camargo, J.M.F. & Wittmann, D. (1989) Nest architecture and distribution of the primitive stingless bee, Mourella caer-ulea (Hymenoptera, Apidae, Meliponinae): Evidence for the origin of Plebeia (s.lat.) on the Gondwana continent.Studies on Neotropical Fauna and Environment, 24, 213–229.

Cameron, E.C., Franck, P., & Oldroyd, B.P. (2004) Genetic structure of nest aggregations and drone congregations of thesoutheast Asian stingless bee Trigona collina. Molecular Ecology, 13, 2357–2364.



Cameron, P. (1897) Hymenoptera Orientalia, or contributions to a knowledge of the Hymenoptera of the Oriental zoolog-ical region. Part V. Memoirs and Proceedings of the Manchester Literary & Philosophical Society, 41, 15–144.

Cameron, P. (1901) On the Hymenoptera collected during the "Skeat Expedition" to the Malay Peninsula, 1899–1900.Proceedings of the Zoological Society of London, 1901, 16–44.

Cameron, P. (1902) On the Hymenoptera collected by Mr. Robert Shelford at Sarawak, and on the Hymenoptera of theSarawak museum. Journal of the Straits Branch of the Royal Asiatic Society, 37, 29–131.

Cameron, P. (1908) On some bornean species of Trigona (Apidae). The Entomologist, 41, 192–195.Cannon, R.C. (1937) Insect fauna of the Upper Walsh River, north Queensland, continued. The North Queensland Natu-

ralist (The Journal and Magazine of the North Queensland Naturalist Club), 5, 1–3.Cardale, J.C. (1993). Hymenoptera: Apoidea. In: W.W.K. Houston & G.V. Maynard (Eds.), Zoological catalogue of Aus-

tralia. Vol. 10, AGPS, Canberra, pp. x, 406.Castets, J. (1893) Les abeilles du sud de l'Inde. Revue des Questions Scientifiques, 34, 465–488, plate.Cepeda, O.I. (2006) Division of labor during brood production in stingless bees with special reference to individual par-

ticipation. Apidologie, 37, 175–190.Cervancia, C.R. & Barile, G.E. (1993). Foraging behaviour of Trigona biroi Friese (Apidae: Hymenoptera. In: G.K.

Veeresh, R.U. Shaanker & K.N. Ganashaiah (Eds.), Pollination in tropics. Proceedings of the international sympo-sium on pollination in tropics, IUSSI, Bangalore, pp. 78–80.

Chaudhary, O.P. & Kumar, R. (2000) Studies on honeybee foraging and pollination in cardamom (Elettaria cardamo-mum Maton). Journal of Spices and Aromatic Crops, 9, 37–42.

Chinajariyawong, A. & Saiboon, S. (2000) Stingless bees in Wongkhla province. In Proceedings of the 38th KasetsartUniversity Annual Conference: Plants, Agricultural Extension and Communication, pp. 445–449, Bangkok.

Chinh, T.X. & Sommeijer, M.J. (2005) Production of sexuals in the stingless bee Trigona (Lepidotrigona) ventralis fla-vibasis Cockerell (Apidae, Meliponini) in northern Vietnam. Apidologie, 36, 493–503.

Chinh, T.X., Sommeijer, M.J., Boot, W.J., & Michener, C.D. (2005) Nest and colony characteristics of three stingless beespecies in Vietnam with the first description of the nest of Lisotrigona carpenteri (Hymenoptera: Apidae: Melipo-nini). Journal of the Kansas Entomological Society, 78, 363–372.

Chourykaew, B., Khunwasi, C., Boonkerd, T., & Seelanan, T. (2002) Floral visitors and fruit set in Afgekia sericea Craib(Fabaceae). Natural History Journal of Chulalongkorn University, 4, 31–44.

Claeys, J.Y. (1954) Deux Français à la conquête de Singapour. Les naturalistes Alfred Duvaucel et Pierre–Médard Diard.France–Asie (Association pour une Meilleure Connaissance de l'Asie), 99, 1001–1007.

Cobara, B. & Dejan, A. (1998) A stingless bee nesting inside ant–gardens in French Guiana (Hymenoptera: Apidae).Sociobiology, 32, 489–492.

Cockerell, T.D.A. (1905) Descriptions and records of bees. —I. Annals and Magazine of Natural History, [7]16, 216–225.

Cockerell, T.D.A. (1910) New and little known bees. Transactions of the American Entomological Society, 36, 199–249.Cockerell, T.D.A. (1911) The bees of the Solomon Islands. Proceedings of the Linnean Society of New South Wales, 36,

160–178.Cockerell, T.D.A. (1912) Descriptions and records of bees. —XLII. Annals and Magazine of Natural History, [8]9, 220–

229.Cockerell, T.D.A. (1914) Bees visiting Helianthus. Canadian Entomologist, 46, 409–415.Cockerell, T.D.A. (1915) Descriptions and records of bees. —LXVIII. Annals and Magazine of Natural History, [8]16,

1–9.Cockerell, T.D.A. (1918) Descriptions and records of bees. —LXXX. Annals and Magazine of Natural History, [9]2,

384–390.Cockerell, T.D.A. (1919a) Descriptions and records of bees. —LXXXIII. Annals and Magazine of Natural History, [9]3,

118–125.Cockerell, T.D.A. (1919b) Descriptions and records of bees. —LXXXV. Annals and Magazine of Natural History, [9]3,

240–250.Cockerell, T.D.A. (1919c) The social bees of the Philippine Islands. Philippine Journal of Science, 14, 77–81.Cockerell, T.D.A. (1920a) Descriptions and records of bees. —LXXXVIII. Annals and Magazine of Natural History,

[9]5, 113–119.Cockerell, T.D.A. (1920b) Some bees from Sandakan, Borneo. Philippine Journal of Science, 17, 221–229.Cockerell, T.D.A. (1921) Fossil arthropods in the British Museum. —VII. Annals and Magazine of Natural History, [9]8,

541–545.Cockerell, T.D.A. (1922a) Australian bees in the Queensland museum. Memoirs of the Queensland Museum, 7, 257–279.Cockerell, T.D.A. (1922b) Fossils in burmese amber. Nature, 109, 713–714.Cockerell, T.D.A. (1923) Descriptions and records of bees. —XCVIII. Annals and Magazine of Natural History, [9]12,

238–247.Cockerell, T.D.A. (1925a) Descriptions and records of bees. —CIV. Annals and Magazine of Natural History, [9]15,



489–496.Cockerell, T.D.A. (1925b) Trigona laeviceps as a pest. Journal of Economic Entomology, 18, 836.Cockerell, T.D.A. (1925c) Some Philippine bees. Philippine Journal of Science, 26, 53–55.Cockerell, T.D.A. (1926a) Descriptions and records of bees. —CXII. Annals and Magazine of Natural History, [9]18,

216–227.Cockerell, T.D.A. (1926b) Trigona laeviceps not a pest. Journal of Economic Entomology, 19, 188–189.Cockerell, T.D.A. (1927) Bees from the Malay Peninsula. Annals and Magazine of Natural History, [9]20, 530–541.Cockerell, T.D.A. (1929a) Bees from the Australian region. American Museum Novitates, 346, 1–18.Cockerell, T.D.A. (1929b) Descriptions and records of bees. —CXVII. Annals and Magazine of Natural History, [10]4,

132–141.Cockerell, T.D.A. (1929c) Descriptions and records of bees. —CXX. Annals and Magazine of Natural History, [10]4,

584–594.Cockerell, T.D.A. (1929d) Bees in the Queensland Museum. Memoirs of the Queensland Museum, 9, 298–323 [stingless

bee section should be read; 299,301,300].Cockerell, T.D.A. (1929e) Bees in the Australian Museum collection. Records of the Australian Museum, 17, 199–243.Cockerell, T.D.A. (1930a) Descriptions and records of bees. —CXXII. Annals and Magazine of Natural History, [10]5,

156–163.Cockerell, T.D.A. (1930b) The bees of Australia. Australian Zoologist, 6, 137–156, 205–238.Cockerell, T.D.A. (1930c) Australian bees in the Museum of Comparative Zoology. Psyche, 37, 141–154.Cockerell, T.D.A. (1934) Some African meliponine bees. Revue de Zoologie et de Botanique Africaines, 26, 46–62.Cockerell, T.D.A. (1936) Bees from the Solomon Islands. Proceedings of the Royal Entomological Society of London,

Series B, Taxonomy, 5, 225–226.Cockerell, T.D.A. (1939a) Bees from the Caroline and Palau islands and Yap (Hymenoptera, Apoidea). Occasional

Papers of the Bernice P. Bishop Museum, 15, 61–66.Cockerell, T.D.A. (1939b) Studies of the Pacific bees in the collection of Bishop Museum (Hymenoptera, Apoidea).

Occasional Papers of the Bernice P. Bishop Museum, 15, 133–140.Cockerell, T.D.A. & LeVeque, N. (1925) Bees from Samar, Philippine Islands. Philippine Journal of Science, 27, 169–

175.Corlett, R.T. (2004) Flower visitors and pollination in the Oriental (Indomalayan) Region. Biological Reviews, 79, 497–

532.Cortopassi–Laurino, M., Imperatriz-Fonseca, V.L., Roubik, D.W., Dollin, A.E., Heard, T.A., Aguilar, I., Venturieri, G.C.,

Eardley, C.D., & Nogueira–Neto, P. (2006) Global meliponiculture: challenges and opportunities. Apidologie, 37,275–292.

Costa, K.F., Brito, R.M., & Miyazawa, C.S. (2004) Karyotypic description of four species of Trigona (Jurine, 1807)(Hymenoptera, Apidae, Meliponini) from the State of Mato Grosso, Brazil. Genetics and Molecular Biology, 27,187–190.

Costa, M.A., Del Lama, M.A., Melo, G.A.R., & Sheppard, W.S. (2003) Molecular phylogeny of the stingless bees (Api-dae, Apinae, Meliponini) inferred from mitochondrial 16S rDNA sequences. Apidologie, 34, 73–84.

Crane, E. (1999) The World History of Beekeeping and Honey Hunting. Routledge, New York, 682 pp.Cunha, R.A.d. (1994) Estudo fenético supra–específico de Trigonini do Velho Mundo (Hymenoptera, Apidae). Revista

Brasileira de Entomologia, 38, 613–626.Cunningham, S.A., Fitzgibbon, F., & Heard, T.A. (2002) The future of pollination in Australia. Australian Journal of

Agricultural Research, 53, 893–900.Dalla Torre, K.W.v. (1896) Catalogus hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 10,

Apidae (Anthophila). G. Engelmann, Leipzig, 643.Darchen, R. (1969) La biologie des trigones et des Mélipones (Hyménoptères Apidæ) a la lumière des travaux récents.

L'Année Biologique (Paris), 8, 455–490.Darchen, R. & Louis, J. (1961) Les mélipones et leur élevage. Melipona – Trigona – Lestrimelitta. Annales de l'Institut

National de la Recherche Agronomique. Série C bis, Annales de l'abeille, 4, 5–39.Devanesan, S., Nisha, M.M., Shailaja, K.K., & Bennet, R. (2003) Natural enemies of stingless bee Trigona iridipennis

Smith in Kerala. Insect Environment, 9, 30.Disney, R.H.L. & Bartareau, T. (1995) A new species of Dohrniphora (Diptera: Phoridae) associated with a stingless bee

(Hymenoptera: Apidae) in Australia. Sociobiology, 26, 229–239.Divan, V.V. (1981("1980")) Honey bee behaviour during total solar eclipse. Indian Bee Journal, 42, 97–99.Dollin, A.E. (1996a) Introduction to Australian native bees. Australian native bee research centre, North Richmond,

NSW, 10 pp.Dollin, A.E. (1996b) Nests of Australian stingless bees. Australian native bee research centre, North Richmond, NSW, 14

pp.Dollin, A.E. (1996c) Behaviour of Australian stingless bees. Australian native bee research centre, North Richmond,



NSW, 13 pp.Dollin, A.E. (1997) How to recognize the different types of Australian stingless bees (covering the social native bee spe-

cies). Australian native bee research centre, North Richmond, NSW, 13 pp.Dollin, A.E. (2002a) Tips on stingless beekeeping by Australian beekeepers. Volume 2. Australian native bee research

centre, North Richmond, NSW, 15 pp.Dollin, A.E. (2002b) Tips on stingless beekeeping by Australian beekeepers. Volume 3. Australian native bee research

centre, North Richmond, NSW, 15 pp.Dollin, A.E. & Dollin, L.J. (1988) Australian native bees give the bicentary a buzz in Arnhem land, NT. The Australasian

beekeeper, 89, 219–223.Dollin, A.E. & Dollin, L.J. (1994) An elusive Australian native bee drone – the key to renaming a species. The Australa-

sian beekeeper, 95, 291–292.Dollin, A.E., Dollin, L.J., & Sakagami, S.F. (1997) Australian stingless bees of the genus Trigona (Hymenoptera: Api-

dae). Invertebrate Taxonomy, 11, 861–896.Dollin, A.E. & Heard, T.A. (1997) Keeping Australian stingless bees in a log or box. Australian native bee research cen-

tre, North Richmond, NSW, 14 pp.Dollin, A.E. & Heard, T.A. (1999) Tips on stingless beekeeping by Australian beekeepers. Volume 1. Australian native

bee research centre, North Richmond, NSW, 17 pp.Dollin, A.E., Zabel, R., & Zabel, J. (2001) "Do it yourself" boxing and splitting hives. Australian native bee research cen-

tre, North Richmond, NSW, 18 pp.Dollin, L.J. & Dollin, A.E. (1983) Honeymooning in far north Queensland with Australian native bees. The Australasian

Beekeeper, 85, 104–107.Dover, C. (1929) Wasps and bees in the Raffles museum, Singapore. Bulletin of the Raffles Museum (Singapore), 2, 43–

70.Drumond, P.M., Bego, L.R., & Melo, G.A.R. (1995) Nest architecture of the stingless bee Plebeia poecilochroa Moure &

Camargo, 1993 and related considerations (Hymenoptera, Apidae, Meliponinae). Iheringia. Série Zoologia (PortoAlegre), 79, 39–45.

Drumond, P.M., Oldroyd, B.P., Dollin, A.E., & Dollin, L.J. (1999) Oviposition behaviour of two Australian stinglessbees, Austroplebeia symei Rayment and Austroplebeia australis Friese (Hymenoptera: Apidae: Meliponini). Austra-lian Journal of Entomology, 38, 234–241.

Drumond, P.M., Oldroyd, B.P., & Osborne, K. (2000) Worker reproduction in Austroplebeia australia Friese(Hymenoptera, Apidae, Meliponini). Insectes Sociaux, 47, 333–336.

Drumond, P.M., Zucchi, R., Mateus, S., Bego, L.R., Yamane, S., & Sakagami, S.F. (1996a) Oviposition behavior of thestingless bees. XVII. Plebeia (Plebeia) droryana and an ethological comparison with other Meliponine taxa(Hymenoptera, Apidae). Japanese Journal of Entomology, 64, 385–400.

Drumond, P.M., Zucchi, R., Yamane, S., & Sakagami, S.F. (1996b) Oviposition behavior of the stingless bees. XVIII.Plebeia (Plebeia) emerina and P. (P.) remota with a preliminary ethological comparison of some Plebeia taxa (Api-dae, Meliponinae). Bulletin of the Faculty of Education, Ibaraki University (Natural Sciences), 45, 31–55.

Eardley, C.D. (2004) Taxonomic revision of the African stingless bees (Apoidea: Apidae: Apinae: Meliponini). AfricanPlant Protection, 10, 63–96.

Eltz, T. (2004) Spatio-temporal variation of apine bee attraction to honeybaits in Bornean forests. Journal of TropicalEcology, 20, 317–324.

Eltz, T., Brühl, C., van der Kaars, S., & Linsenmair, K.E. (2002a) Determinants of stingless bee nest density in lowlanddipterocarp forests of Sabah, Malaysia. Oecologia, 131, 27–34.

Eltz, T., Brühl, C.A., & Görke, C. (2002b) Collection of mold (Rhizopus sp.) spores in lieu of pollen by the stingless beeTrigona collina. Insectes Sociaux, 49, 28–30.

Eltz, T., Brühl, C.A., Imiyabir, Z., & Linsenmair, K.E. (2003) Nesting and nest trees of stingless bees (Apidae: Melipo-nini) in lowland dipterocarp forests in Sabah, Malaysia, with implications for forest management. Forest Ecologyand Management, 172, 301–313.

Eltz, T., Brühl, C.A., van der Kaards, S., Chey, V.K., & Linsenmair, K.E. (2001a) Pollen foraging and resource partition-ing of stingless bees in relation to flowering dynamics in a Southeast Asian tropical rainforest. Insectes Sociaux, 48,273–279.

Eltz, T., Brühl, C.A., van der Kaars, S., & Linsenmair, K.E. (2001b) Assessing stingless bee pollen diet by analysis ofgarbage pellets: a new method. Apidologie, 32, 341–353.

Engel, M.S. (2000) A review of the Indo-Malayan Meliponine genus Lisotrigona, with two new species (Hymenoptera:Apidae). Oriental Insects, 34, 229–237.

Engel, M.S. (2001) A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of theAmerican Museum of Natural History, 259, 1–192.

Engels, W. & Imperatriz-Fonseca, V.L. (1990). Caste development, reproductive strategies, and control of fertility inhoney bees and stingless bees. In: W. Engels (Ed.), Social insects, an evolutionary approach to castes and reproduc-



tion, Springer, Berlin, pp. 167–230.Esaki, T. (1936) Einige biologische Beobachtungen über die Bienen und Wespen Mikronesiens. Mushi, 9, 44–47.Eswarappa, G., Kuberappa, G.C., Sreenath, T.N., Gowda, G., & Sreeramulu, K.R. (2005a) Relative abundance, foraging

activity and pollinating efficiency of different pollinators visiting chow–chow (Sechium edule). Journal of Ecobiol-ogy (India), 17, 509–516.

Eswarappa, G., Kuberappa, G.C., Sreenath, T.N., Sreeramulu, K.R., & Gowda, G. (2005b) Role of different species ofhoneybees in cross pollination of chow–chow (Sechium edule). Journal of Ecobiology (India), 17, 101–107.

Eswarappa, G., Kuberappa, G.C., Sreenath, T.N., Sreeramulu, K.R., & Gowda, G. (2005c) Nectar secretion and its sugarconcentration on foraging activity of different species of honey bees in chow–chow (Sechium edule). Journal ofEcobiology (India), 17, 337–343.

Evans, H.E., Evans, M.A., & Hook, A. (1982) Observations on the nests and prey of Australian Bembix sand wasps(Hymenoptera: Sphecidae). Australian Journal of Zoology, 30, 71–80.

Evans, H.E. & Matthews, R.W. (1973) Systematics and nesting behavior of Australian Bembix sand wasps(Hymenoptera, Sphecidae). Memoirs of the American Entomological Institute, 20, 1–387.

Evenhuis, N.L. (2007) The insect and spider collections of the world. Available from http://hbs.bishopmuseum.org/codens/codens-r-us.html.

Fain, A. & Heard, T.A. (1987) Description and life cycle of Cerophagus trigona spec. nov. (Acari, Acaridae), associatedwith the stingless bee Trigona carbonaria Smith in Australia. Bulletin de l'Institut Royal des Sciences Naturelles deBelgique, Entomologie, 57, 197–202.

Fidalgo, A.d.O. & Kleinert, A.d.M.P. (2007) Foraging behavior of Melipona rufiventris Lepeletier (Apinae; Meliponini)in Ubatuba, SP, Brazil. Brazilian Journal of Biology, 67, 133–140.

Forster, P.I., Machin, P.J., Mound, L., & Wilson, G.W. (1994) Insects associated with reproductive structures of cycads inQueensland and northeast New South Wales, Australia. Biotropica, 26, 217–222.

Franck, P., Cameron, E.C., Good, G., Rasplus, J.–Y., & Oldroyd, B.P. (2004) Nest architecture and genetic differentiationin a species complex of Australian stingless bees. Molecular Ecology, 13, 2317–2331.

Friese, H. (1898) Die Trigona–Arten Australiens. Természetrajzi Füzetek, 21, 427–431.Friese, H. (1900) Neue Arten der Bienengattungen Melipona Ill., und Trigona Jur. Természetrajzi Füzetek, 23, 381–394.Friese, H. (1901) Neue Arten der Bienengattung Trigona Jur. (Hym.). Zeitschrift für systematische Hymenopterologie

und Dipterologie, 1, 265–271.Friese, H. (1904) Eine Bienenausbeute von Java (Hym.). Allgemeine Zeitschrift für Entomologie, 9, 138–140.Friese, H. (1905) Ein Bienennest mit Vorratskammern (Lithurgus dentipes Sm.). Zeitschrift für wissenschaftliche Insek-

tenbiologie, 1, 118–119.Friese, H. (1909) Die Bienenfauna von Neu–Guinea. Annales Historico–Naturales Musei Nationalis Hungarici (Ter-

mészettudományi Múzeum évkönyve), 7, 179–288.Friese, H. (1909("1908")). Hymenoptera II. Apidae. In: A. Wichmann (Ed.), Nova Guinea. Résultats de l'expédition sci-

entifique néerlandaise à la Nouvelle–Guinée en 1903. Vol. 5 (Zoologie), E.J. Brill, Leiden, pp. 353–359.Friese, H. (1912) Zur Bienenfauna von Neu-Guinea und Oceanien. Übersicht des von den Herren Dahl, Finsch, Krämer,

Moszkowski, Neuhaus, Schoede und L. Schultze in den letzten Jahren dem Zoologischen Museum in Berlin über-wiesenen Forschungsmaterials. Mitteilungen aus dem Zoologischen Museum in Berlin, 6, 91–96.

Friese, H. (1914a) Die Bienenfauna von Java. Tijdschrift voor Entomologie, 57, 1–13.Friese, H. (1914b) Anhang zur: "Bienenfauna von Java". Tijdschrift voor Entomologie, 57, 14–61.Friese, H. (1915) Apiden aus Nord-Neu-Guinea gesammelt von Dr. P. N. van Kampen und K. Gjellerup, in den Jahren

1910 und 1911. Tijdschrift voor Entomologie, 58, 1–4.Friese, H. (1917) Results of Dr. E. Mjöberg's Swedish scientific expeditions to Australia 1910–1913. 13. Apidae. Arkiv

för Zoologi (Stockholm), 11(2), 1–9.Friese, H. (1918) Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien, ausgeführt im Auftrage der Kgl.

Preuß. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen. VII. Bienen aus Sumatra, Java, Malakkaund Ceylon. Gesammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren 1911–1912. Zoologische Jahrbücher,Abteilung für Systematik, Geographie und Biologie der Tiere, 41, 489–520.

Friese, H. (1925) Neue neotropischen Bienenarten, zugleich II. Nachtrag zur Bienenfauna von Costa Rica (Hym.). Stet-tiner Entomologische Zeitung, 86, 1–41.

Friese, H. (1933a) Neue Trigona–Arten von Borneo (Hym.). Natuurhistorisch Maandblad, 22, 45–46.Friese, H. (1933b) Eine neue Trigona von Sumatra (Hymenoptera, Apidae). Natuurhistorisch Maandblad, 22, 147.Frith, C.B. (1978) Weaver ants, Oecophylla smeragadina [sic], systematically predating worker sweat bees, Trigona sp.,

by snatching them from flight. Natural History Bulletin of the Siam Society, 27, 194–195.Gardener, M.C., Rowe, R.J., & Gillman, M.P. (2003) Tropical bees (Trigona hockingsi) show no preference for nectar

with amino acids. Biotropica, 35, 119–125.George, C.J. (1933) Excretion in Melipona during metamorphosis. Journal of the University of Bombay, 1, 58–61.George, C.J. (1934) The bionomics, morphology, and metamorphosis of Melipona irridipennis (sic). Journal of the Uni-



versity of Bombay, 2, 1–16.Gibbons, B. (1992) Stalking the bush bee. Wildlife Australia (Wildlife Preservation Society of Queensland), 29, 3.Gloag, R.S., Beekman, M., Heard, T.A., & Oldroyd, B.P. (2007) No worker reproduction in the Australian stingless bee

Trigona carbonaria Smith (Hymenoptera, Apidae). Insectes Sociaux, 54, 412–417.Goebel, R.L. (1986) Australian native bees. Queensland Agricultural Journal, 112, 285–286.Goel, S.C. & Kumar, A. (1990) Insect pests and predators associated to sunflower in winters of northern India. Indian

Journal of Entomology, 52, 39–45.Graur, D. (1985) Gene diversity in Hymenoptera. Evolution, 39, 190–199.Greco, M., Spooner–Hart, R., & Holford, P. (2005) A new technique for monitoring Trigona carbonaria nest contents,

brood and activity using X-ray computerized tomography. Journal of Apicultural Research, 44, 97–100.Green, C.L., Franck, P., & Oldroyd, B.P. (2001) Characterization of microsatellite loci for Trigona carbonaria, a sting-

less bee endemic to Australia. Molecular Ecology Notes, 1, 89–92.Green, C.L. & Oldroyd, B.P. (2002) Queen mating frequency and maternity of males in the stingless bee Trigona carbon-

aria Smith. Insectes Sociaux, 49, 196–202.Gribodo, G. (1891) Contribuzioni imenotterologiche sopra alcune specie nuove o poco conosciute di imenotteri antofili

(Generi Ctenoplectra, Xylocopa, Centris, Trigona e Bombus). Bollettino della Societá Entomologica Italiana, Gen-ova, 23, 102–119.

Gribodo, G. (1893) Note imenotterologische. Nota II. Nuovi generi e nuove specie di imenotteri antofili ed osservazionisopra alcune specie già conosciute. Bollettino della Societá Entomologica Italiana, Genova, 25, 248–287.

Grimaldi, D.A., Shedrinsky, A., Ross, A.J., & Baer, N.S. (1995) Forgeries of fossils in “amber”: history, identificationand case studies. Curator (New York), 37, 251–274.

Gross, C.L. (1993) The breeding system and pollinators of Melastoma affine (Melastomataceae); a pioneer shrub in trop-ical Australia. Biotropica, 25, 468–474.

Haddon, A.C. (1901) The omen animals of Sarawak. Popular Science Monthly, 60, 80–88.Harriss, F.C.L. & Beatie, A.J. (1991) Viability of pollen carried by Apis mellifera L., Trigona carbonaria Smith and

Vespula germanica (F.) (Hymenoptera: Apidae, Vespidae). Journal of the Australian Entomological Society, 30, 45–47.

Hartfelder, K., Makert, G.R., Judice, C.C., Pereira, G.A.G., Santana, W.C., Dallacqua, R., & Bitondi, M.M.G. (2006)Physiological and genetic mechanisms underlying caste development, reproduction and division of labor in stinglessbees. Apidologie, 37, 144–163.

Hawkeswood, T.J. (1983) Pollination and fruit production of Cupaniopsis anacardioides (A. Rich.) Radlkf. (Sapin-daceae) at Townsville, north Queensland. 1. Pollination and floral biology. Victorian Naturalist, 100, 12–20.

Heard, T.A. (1987) Nest architecture of Trigona bees. Entomological Society of Queensland News Bulletin, 14, 132.Heard, T.A. (1988) Propagation of hives of Trigona carbonaria Smith (Hymenoptera: Apidae). Journal of the Australian

Entomological Society, 27, 303–304.Heard, T.A. (1993) Pollinator requirements and flowering patterns of Macademia integrifolia. Australian Journal of Bot-

any, 41, 491–497.Heard, T.A. (1994) Behaviour and pollinator efficiency of stingless bees and honey bees on macadamia flowers. Journal

of Apicultural Research, 33, 191–198.Heard, T.A. (1996) Stingless bees. Nature Australia, 1996(spring), 50–55.Heard, T.A. (1998) Stingless bees as pollinators of macadamia. Aussie Bee, 7, 8.Heard, T.A. (1999a) The role of stingless bees in crop pollination. Annual Review of Entomology, 44, 183–206.Heard, T.A. (1999b) Pollination of mangoes: flies or stingless bees. Aussie Bee, 11, 11.Heard, T.A. (2000) Could Aussie stingless bees pollinate avocado? Secrets from the jungles of Central America. Aussie

Bee, 15, 20–21.Heard, T.A. (2003) Native stingless bees: biology, conservation and economic utilisation. News Bulletin of the Entomo-

logical Society of Queensland, 31, 134–141.Heard, T.A. & Dollin, A.E. (1998) Crop pollination with australian stingless bees. Australian native bee research centre,

North Richmond, NSW, 17.Heard, T.A. & Dollin, A.E. (2000) Stingless bee keeping in Australia: snapshot of an infant industry. Bee World, 81, 116–

125.Heard, T.A. & Exley, E.M. (1994) Diversity, abundance, and distribution of insect visitors to Macadamia flowers. Envi-

ronmental Entomology, 23, 91–100.Heard, T.A. & Hendrikz, J.K. (1993) Factors influencing flight activity of colonies of the stingless bee Trigona carbon-

aria (Hymenoptera: Apidae). Australian Journal of Zoology, 41, 343–353.Heard, T.A., Vithanage, V., & Chacko, E.K. (1990) Pollination biology of cashew in the Northern Territory of Australia.

Australian Journal of Agricultural Research, 41, 1101–1114.Hedicke, H. (1926) Beiträge zur Apidenfauna der Philippinen (Hym.). (2. Beitrag zur Kenntnis orientalischer Apiden).

Deutsche Entomologische Zeitschrift, 1926, 413–423.



Heiling, A.M. & Herberstein, M.E. (2004) Predator-prey coevolution: Australian native bees avoid their spider predators.Proceedings of the Royal Society of London, Series B (Supplement), 271, S196–S198.

Hilário, S.D., Imperatriz-Fonseca, V.L., & Leinert, A.d.M.P. (2000) Flight activity and colony strength in the stinglessbee Melipona bicolor bicolor (Apidae, Meliponinae). Revista Brasileira de Biologia, 60, 299–306.

Hilário, S.D., Imperatriz-Fonseca, V.L., & Leinert, A.d.M.P. (2001) Responses to climatic factors by foragers of Plebeiapugnax Moure (in litt.) (Apidae, Meliponinae). Revista Brasileira de Biologia, 61, 191–196.

Hockings, H.J. (1884) Notes on two Australian species of Trigona. Transactions of the Entomological Society of London,32, 149–157.

Hoese, F.J., Law, E.A.J., Rao, D., & Heberstein, M.E. (2006) Distinctive yellow bands on a sit-and-wait predator: preyattractant or camouflage? Behaviour, 143, 763–781.

Hofstede, F.E. & Sommeijer, M.J. (2006) Effect of food availability on individual foraging specialisation in the stinglessbee Plebeia tobagoensis (Hymenoptera, Meliponini). Apidologie, 37, 387–397.

Hohmann, H. (2003) Zur Kommunikation stachelloser Bienen in Neuguinea. TenDenZen (Jahrbuch ÜberseemuseumBremen), 11, 9–26.

Horne, C. & Smith, F. (1870) Notes on the habits of some hymenopterous insects from the North-West provinces ofIndia. With an appendix, containing descriptions of some new species of Apidae and Vespidae collected by Mr.Horne by Frederick Smith, of the British Museum. Illustrated by plates from drawings by the author of the notes.Transactions of the Zoological Society of London, 7, 161–196.

Hoshiba, H. & Imai, H.T. (1993) Chromosome evolution of bees and wasps (Hymenoptera, Apocrita) on the basis of C-banding pattern analyses. Japanese Journal of Entomology, 61, 465–492.

Ihering, H.v. (1912) Zur Biologie der brasilianischen Meliponiden. Zeitschrift für wissenschaftliche Insektenbiologie, 8,1–5, 43–46.

Ikudome, S. & Kusigemati, K. (1996) Notes on some bees from the Palau Islands (Hymenoptera, Apoidea). KagoshimaUniversity Research Center for the South Pacific Occasional Papers, 30, 17–21.

Illiger, K. (1806) William Kirby's Familien der Bienenartigen Insekten mit Zusätzen, Nachweisungen und Bemerkungen.Magazin für Insektenkunde, 5, 28–175.

Imperatriz-Fonseca, V.L. & Zucchi, R. (1995) Virgin queens in stingless bee (Apidae, Meliponinae) colonies: A review.Apidologie, 26, 231–244.

Inoue, K., Kato, M., & Inoue, T. (1995) Pollination ecology of Dendrobium setifolium, Neuwiedia borneensis, andLecanorchis multiflora (Orchidaceae) in Sarawak. Tropics, 5, 95–100.

Inoue, K., Maki, M., & Masuda, M. (1996a). Evolution of Campanula flowers in relation to insect pollinators on islands.In: D.G. Lloyd & S.C.H. Barrett (Eds.), Floral biology, Chapman & Hall, New York, pp. 377–400.

Inoue, T. (1990). An analysis of apid bee richness in Sumatra. In: G.K. Veeresh, B. Mallik & C.A. Viraktamath (Eds.),Social insects and the environment, proceedings of the 11th international congress of IUSSI, 1990 (Internationalunion for the study of social insects), Oxford, New Delhi, pp. 581–582.

Inoue, T., Adri, & Salmah, S. (1990a). Nest site selection and reproductive ecology of the Asian honey bee, Apis ceranaindica, in central Sumatra. In: S.F. Sakagami, R. Ohgushi & D.W. Roubik (Eds.), Natural History of social waspsand bees in equatorial Sumatra, Hokkaido University Press, Sapporo, pp. 219–232.

Inoue, T., Nakamura, K., Salmah, S., & Abbas, I. (1993) Population dynamics of animals in unpredictably-changingtropical environments. Journal of Bioscience, 18, 425–455.

Inoue, T., Roubik, D.W., & Suka, T. (1999) Nestmate recognition in the stingless bee Melipona panamica (Apidae, Meli-ponini). Insectes Sociaux, 46, 208–218.

Inoue, T. & Sakagami, S.F. (1993) A new name of Trigona (Hymenoptera, Apidae). Japanese Journal of Entomology,61, 769.

Inoue, T., Sakagami, S.F., Salmah, S., & Nukmal, N. (1984a) Discovery of successful absconding in the stingless beeTrigona (Tetragonula) laeviceps. Journal of Apicultural Research, 23, 136–142.

Inoue, T., Sakagami, S.F., Salmah, S., & Yamane, S. (1984b) The process of colony multiplication in the Sumatran sting-less bee Trigona (Tetragonula) laeviceps. Biotropica, 16, 100–111.

Inoue, T., Salmah, S., Abbas, I., & Yusuf, E. (1985a) Foraging behavior of individual workers and foraging dynamics ofcolonies of three sumatran stingless bees. Researches on Population Ecology (Kyoto), 27, 373–392.

Inoue, T., Salmah, S., & Sakagami, S.F. (1996b) Individual variations in worker polyethism of the Sumatran stinglessbee, Trigona (Tetragonula) minangkabau (Apidae, Meliponinae). Japanese Journal of Entomology, 64, 641–668.

Inoue, T., Salmah, S., Sakagami, S.F., Yamane, S., & Kato, M. (1990b). An analysis of anthophilous insects in centralSumatra. In: S.F. Sakagami, R. Ohgushi & D.W. Roubik (Eds.), Natural History of social wasps and bees in equato-rial Sumatra, Hokkaido University Press, Sapporo, pp. 175–200.

Inoue, T., Salmah, S., & Yusuf, E. (1985b). Foraging behavior of individual workers and foraging dynamics of coloniesof three Sumatran stingless bees [summary of Researches on Population Ecology (Kyoto), 27: 373–392]. In:Sumatra nature study (Entomology) (Ed.), Evolutionary ecology of insects in humid tropics, especially in centralSumatra, Kanazawa University, Kanazawa (Japan), pp. 44–49.



Inson, C. & Malaipan, S. (2006) [original title in Thai=Diversity of stingless bees (Apidae: Trigona spp. and Hypotrig-ona spp.) and their resin and gum collecting behavior from nature in golden jubilee Thong Pha Phum project, ThongPha Phum district, Kanchanaburi Province]. BRT (The Thai Response to Biodiversity), 2549, 20–31.

International Commission on Zoological Nomenclature (1999) International code of zoological nomenclature, fourthedition. International Trust for Zoological Nomenclature, London, xxix+1+306.

Irish, J., Carter, D.A., Blair, S.E., & Heard, T.A. (2008) Antibacterial activity of honey from the Australian stingless beeTrigona carbonaria. International Journal of Antimicrobial Agents, 32, 89–98.

Jagadish, K.S., Venkatesh-Gowda, T.N., Lakshmikantha, B.P., & Gowda, A.N.S. (2002) Foraging behaviour of honeybees and other insect visitors on the bloom of two medicinal herbs, Hygrophila auriculata (Schum.) Heine and Leu-cas aspera (Willd.) Link. Pest Management and Economic Zoology, 10, 67–70.

Jain, P.C. & Kushwaha, K.S. (1972) Morphometrics of bees, Apis dorsata F., A. indica F., A. florea F., Megachile lanataF., Trigona smithii B. (Hymenoptera: Apidae) and their foraging habits at Udaipur. University of Udaipur ResearchJournal, [5], 38–40 [reference not seen].

Jander, R. & Jander, U. (1970) Über die Phylogenie der Geotaxis innerhalb der Bienen (Apoidea). Zeitschrift für vergle-ichende Physiologie, 66, 355–368.

Jarau, S., Hrncir, M., Zucchi, R., & Barth, F.G. (2000) Recruitment behavior in stingless bees, Melipona scutellaris andM. quadrifasciata. I. Foraging at food sources differing in direction and distance. Apidologie, 31, 81–91.

Jobiraj, T. & Narendran, T.C. (2004) A revised key to the world species of Lisotrigona Moure (Hymenoptera: Apoidea:Apidae) with description of a new species from India. Entomon, 29, 39–43.

Johnson, L.K. & Appanah, S. (1990). Defenses against ants in nine species of stingless bees. In: G.K. Veeresh, B. Mallik& C.A. Viraktamath (Eds.), Social insects and the environment, proceedings of the 11th international congress ofIUSSI, 1990 (International union for the study of social insects), Oxford, New Delhi, pp. 585–586.

Jongjitvimol, T., Boontawon, K., Wattanachaiyingcharoen, W., & Deowanish, S. (2005) Nest dispersion of a stinglessbee species, Trigona collina Smith, 1857 (Apidae, Meliponinae) in a mixed deciduous forest in Thailand. NaturalHistory Journal of Chulalongkorn University, 5, 69–71.

Jongjitvimol, T. & Wattanachaiyingcharoen, W. (2006) Pollen food sources of the stingless bees Trigona apicalis Smith,1857, Trigona collina Smith, 1857 and Trigona fimbriata Smith, 1857 (Apidae, Meliponinae) in Thailand. NaturalHistory Journal of Chulalongkorn University, 6, 75–82.

Jongjitvimol, T. & Wattanachaiyingcharoen, W. (2007) Distribution, nesting sites and nest structures of the stingless beespecies, Trigona collina Smith, 1857 (Apidae, Meliponinae) in Thailand. Natural History Journal of ChulalongkornUniversity, 7, 25–34.

Jongjitvimol, T., Wattanachaiyingcharoen, W., Deowanish, S., & Wattanachaiyingcharoen, D. (2006) [Original title inThai=Factors affecting foraging behaviors and food resource partitioning of three Trigona species (Trigona apicalisSmith, T. collina Smith and T. fimbriata Smith) in the Phitsanulok area]. BRT (The Thai Response to Biodiversity),2549, 246–252.

Joshi, M.A., Lakshmi, K., & Suryanarayana, M.C. (1998) Melittopalynological investigations on Apis and Trigona hon-eys collected in and around Pune, Maharashtra. Indian Bee Journal, 60, 90–98.

Kajobe, R. & Echazarreta, C.M. (2005) Temporal resource partitioning and climatological influences on colony flightand foraging of stingless bees (Apidae; Meliponini) in Ugandan Tropical Forests. African Journal of Ecology, 43,267–275.

Kakutani, T., Inoue, T., Tezuka, T., & Maeta, Y. (1993) Pollination of strawberry by the stingless bee, Trigona minangka-bau, and the honey bee, Apis mellifera: an experimental study of fertilization efficiency. Researches on PopulationEcology (Kyoto), 35, 95–111.

Kerr, W.E. (1960) Evolution of communication in bees and its role in speciation. Evolution, 14, 386–387.Kerr, W.E. (1969) Some aspects of the evolution of social bees (Apidae). Evolutionary Biology, 3, 119–175.Kerr, W.E., Carvalho, G.A., & Nascimento, V.A. (1996) Abelha Uruçu — biologia manejo e conservação. Fundação

Acangaú, Belo horizonte, 144 pp.Kerr, W.E. & Cunha, R.A.d. (1976) Taxonomic position of two social bees (Apidae). Revista de Biología Tropical, 24,

35–43.Kerr, W.E. & Esch, H. (1965) Comunicação entre as abelhas sociais brasileiras e sua contribuição para o entendimento da

sua evolução. Ciência e Cultura, 17, 529–538.Kerr, W.E., Ferreira, A., & Mattos, N.S.d. (1963) Communication among stingless bees — additional data

(Hymenoptera: Apidae). Journal of the New York Entomological Society, 71, 80–90.Kerr, W.E. & Maule, V. (1964) Geographic distribution of stingless bees and its implications (Hymenoptera: Apidae).

Journal of the New York Entomological Society, 72, 2–18.Kerr, W.E., Sakagami, S.F., Zucchi, R., Portugal-Araújo, V.d., & Camargo, J.M.F. (1967) Observações sôbre a

arquitetura dos ninhos e comportamento de algumas espécies de abelhas sem ferrão das vizinhaças de Manaus,Amazonas (Hymenoptera, Apoidea). Atas do Simpósio Sôbre a Biota Amazônica, Zoologia, 5, 255–309.

Kiew, B.H. (1985) Effective and economical lure for catching sweat bees. Malayan Naturalist, 38, 30.



Kistner, D.H. (1982). The social insects' bestiary. In: H.R. Hermann (Ed.), Social insects. Vol. 3, Academic Press, NewYork, pp. 1–244.

Klakasikorn, A., Wongsiri, S., Deowanish, S., & Duangphakdee, O. (2005) New record of stingless bees (Meliponini:Trigona) in Thailand. Natural History Journal of Chulalongkorn University, 5, 1–7.

Klein, A.-M., Steffan-Dewenter, I., Buchori, D., & Tscharntke, T. (2002) Effects of land-use intensity in tropical agrofor-estry systems on coffee flower-visiting and trap-nesting bees and wasps. Conservation Biology, 16, 1003–1014.

Klein, A.-M., Steffan-Dewenter, I., & Tscharntke, T. (2003a) Pollination of Coffea canephora in relation to local andregional agroforestry management. Journal of Applied Ecology, 40, 837–845.

Klein, A.-M., Steffan-Dewenter, I., & Tscharntke, T. (2003b) Fruit set of highland coffee increases with the diversity ofpollinating bees. Proceedings of the Royal Society of London, Series B, 270, 955–961.

Klumpp, J. (2007) Australian stingless bees, a guide to sugarbag beekeeping. Earthling Enterprises, West End, Australia,110.

Kocyan, A. & Endress, P.K. (2001) Floral structure and development of Apostasia and Neuwiedia (Apostasioideae) andtheir relationships to other Orchidaceae. International Journal of Plant Sciences, 162, 847–867.

Koedam, D., Jungnickel, H., Tentschert, J., Jones, G.R., & Morgan, E.D. (2002) Production of wax by virgin queens ofthe stingless bee Melipona bicolor (Apidae, Meliponinae). Insectes Sociaux, 49, 229–233.

Koeniger, N. & Vorvohl, G. (1979) Competition for food among four sympatric species of Apini in Sri Lanka (Apis dor-sata, Apis cerana, Apis florea and Trigona iridipennis). Journal of Apicultural Research, 18, 95–109.

Koshy, K.C., Harikumar, D., & Narendran, T.C. (2001) Insect visits to some bamboos of the Western Ghats, India. Cur-rent Science (India), 81, 833–838.

Krombein, K.V. (1949) Records of bees from the Solomon islands with descriptions of new subspecies (Hymenoptera,Apoidea). Bulletin of the Brooklyn Entomological Society, 44, 10–14.

Krombein, K.V. (1950) The aculeata Hymenoptera of Micronesia, II. Colletidae, Halictidae, Megachilidae, and Apidae.Proceedings of the Hawaiian Entomological Society, 14, 101–142.

Krombein, K.V. (1951) Additional notes on the bees of the Solomon islands (Hymenoptera: Apoidea). Proceedings of theHawaiian Entomological Society, 14, 277–295.

Kshirsagar, K.K. & Chauhan, R.M. (1977) Preliminary observations on oviposition behaviour in Trigona (Tetragona) iri-dipennis Smith. Indian Bee Journal, 39, 22–23.

Kuberappa, G.C., Eswarappa, G., Vishweshwaraiah, C.H., & Sajjanar, S.M. (2002) Activity and abundance of insects vis-iting guava (Psidium guajava L.) flowers. Pest Management and Economic Zoology, 11, 59–63.

Kumar, S. & Kumar, P. (1989) A study of the pollen carrying capacity of honey bees found in Lucknow in relation totheir body weight. Bulletin of Entomology (Madras), 30, 172–177.

Lakshmi, K. & Suryanarayana, M.C. (1999("1997")) A contribution to the study of the kapok tree, Ceiba pentandra (L.)Gaertn., as a source of forage to honey bees. Indian Bee Journal, 59, 165–169.

Lea, A.M. (1910) Australian and Tasmanian Coleoptera inhabiting or resorting to the nests of ants, bees, and termites.Proceedings of the Royal Society of Victoria, 23, 116–230.

Lea, A.M. (1912) Australian and Tasmanian Coleoptera inhabiting or resorting to the nests of ants, bees, and termites,supplement. Proceedings of the Royal Society of Victoria, 25, 31–78.

Leonhardt, S.D., Dworschak, K., Eltz, T., & Blüthgen, N. (2007) Foraging loads of stingless bees and utilisation of storednectar for pollen harvesting. Apidologie, 38, 125–135.

Lepeletier de Saint Fargeau, A.L.M. (1836). Histoire naturelle des insectes. Hyménoptères. In. Vol. 1, Librairie encyclo-pédique de Roret, Paris, pp. 1–547.

Lindauer, M. (1956) Über die Verständigung bei indischen Bienen. Zeitschrift für vergleichende Physiologie, 38, 521–557.

Lindauer, M. & Kerr, W.E. (1958) Die gegenseitige Verständigung bei den stachellosen Bienen. Zeitschrift für vergle-ichende Physiologie, 41, 405–434.

Lindauer, M. & Kerr, W.E. (1960) Communication between the workers of stingless bees. Bee World, 41, 29–41, 65–71.Liow, L.H. (1999) Bee diversity along a gradient of disturbance in tropical lowland forests of Southeast Asia. CBS:s

Skriftserie (Sweden), 3, 101–130.Liow, L.H., Sodhi, N.S., & Elmqvist, T. (2001) Bee diversity along a gradient of disturbance in tropical lowland forests

of South-east Asia. Journal of Applied Ecology, 38, 180–192.MacPherson, K. & Rupp, H.M.R. (1936) Further notes on orchid pollination. The North Queensland Naturalist (The

Journal and Magazine of the North Queensland Naturalists' Club), 4, 25–26.Maynard, G.V., Hamilton, J.G., & Grimshaw, J.F. (2004) Quarantine — Phytosanitary, sanitary and incursion manage-

ment: an Australian entomological perspective. Australian Journal of Entomology, 43, 318–328.McKenzie, E. (1975) Growing up with Aborigines. The Queensland Naturalist, 21, 46–51.Michener, C.D. (1961) Observations on the nests and behavior of Trigona in Australia and New Guinea (Hymenoptera,

Apidae). American Museum Novitates, 2026, 1–46.Michener, C.D. (1964) Evolution of the nests of bees. American Zoologist, 4, 227–239.



Michener, C.D. (1965) A classification of the bees of the Australian and south Pacific regions. Bulletin of the AmericanMuseum of Natural History, 130, 1–362.

Michener, C.D. (1974) The social behavior of the bees. Harvard University Press, Cambridge, MA, xii+404.Michener, C.D. (1990) Classification of the Apidae (Hymenoptera). Appendix: Trigona genalis Friese, a hitherto

unplaced New Guinea species. University of Kansas Science Bulletin, 54, 75–163.Michener, C.D. (1997) Genus-group names of bees and supplemental family-group names. Scientific Papers, Natural

History Museum, University of Kansas, 1–81.Michener, C.D. (2000) The bees of the world. Johns Hopkins University Press, Baltimore, xiv+[1]+913.Michener, C.D. (2001) Comments on minute Meliponini and the male of the genus Pariotrigona (Hymenoptera: Api-

dae). Journal of the Kansas Entomological Society, 74, 231–236.Michener, C.D. (2007a) The bees of the world, second edition. Johns Hopkins University Press, Baltimore, xvi+953.Michener, C.D. (2007b) Lisotrigona in Thailand, and the male of the genus (Hymenoptera: Apidae: Meliponini). Journal

of the Kansas Entomological Society, 80, 130–135.Michener, C.D. & Boongird, S. (2004) A new species of Trigona from Peninsular Thailand (Hymenoptera: Apidae:

Meliponini). Journal of the Kansas Entomological Society, 77, 143–146.Michener, C.D. & Roubik, D.W. (1993). Observations on the labium of Meliponine bees (Hymenoptera: Apidae). In: T.

Inoue & S. Yamane (Eds.), Evolution of insect societies, Hakuhinsha, Tokyo, pp. 251–266.Milborrow, B.V., Kennedy, J.M., & Dollin, A.E. (1987) Composition of wax made by the Australian stingless bee Trig-

ona australis. Australian Journal of Biological Sciences, 40, 15–25.Mohan, R. & Devanesan, S. (2000) Studies on the behaviour of the stingless bee, Trigona iridipennis Smith (Apidae:

Meliponinae). Indian Bee Journal, 62, 59–62.Mohana Rao, G. (1988("1987")) Scented methi — a promising bee forage crop. Indian Bee Journal, 49, 14–15.Mohana Rao, G. & Suryanarayana, M.C. (1989("1988")) Studies on pollination of watermelon (Citrullus lanatus

(Thunb.) Mansf.). Indian Bee Journal, 50, 5–8.Mohana Rao, G. & Suryanarayana, M.C. (1992("1990")) Studies on the foraging behaviour of honey bees and its effect

on the seed yield in niger. Indian Bee Journal, 52, 31–33.Momose, K. & Hamid, A.A. (2005). The plant-pollinator community in a lowland Dipterocarp forest. In: D.W. Roubik,

S. Sakai & A.A. Hamid (Eds.), Pollination ecology and the rain forest, Sarawak studies. Vol. 174, Springer, pp. 65–72.

Momose, K., Yumoto, T., Nagamitsu, T., Kato, M., Nagamasu, H., Sakai, S., Harrison, R.D., Itioka, T., Hamid, A.A., &Inoue, T. (1998) Pollination biology in a lowland dipterocarp forest in Sarawak, Malaysia. I. Characteristics of theplant-pollinator community in a lowland dipterocarp forest. American Journal of Botany, 85, 1477–1501.

Moure, J.S. (1961) A preliminary supra-specific classification of the old world meliponine bees (Hymenoptera,Apoidea). Studia Entomologica, 4, 181–242.

Moure, J.S. & Camargo, J.M.F. (1978) A fossil stingless bee from copal (Hymenoptera: Apidae). Journal of the KansasEntomological Society, 51, 560–566.

Moure, J.S., Nogueira-Neto, P., & Kerr, W.E. (1958) Evolutionary problems among the Meliponinae (Hymenoptera, Api-dae). Proceedings of the tenth international congress of entomology, 2, 481–493.

Nagamitsu, T. & Inoue, T. (1997) Aggressive foraging of social bees a mechanism of floral resource partitioning in anAsian tropical rainforest. Oecologia, 110, 432–439.

Nagamitsu, T. & Inoue, T. (1998) Interspecific morphological variation in stingless bees (Hymenoptera: Apidae, Meli-poninae) associated with floral shape and location in an Asian tropical rainforest. Entomological Science, 1, 189–194.

Nagamitsu, T. & Inoue, T. (2002) Foraging activity and pollen diets of subterranean stingless bee colonies in response togeneral flowering in Sarawak, Malaysia. Apidologie, 33, 303–314.

Nagamitsu, T. & Inoue, T. (2005). Floral resource utilization by stingless bees (Apidae, Meliponini). In: D.W. Roubik, S.Sakai & A.A. Hamid (Eds.), Pollination ecology and the rain forest, Sarawak studies. Vol. 174, Springer, pp. 73–88.

Nagamitsu, T., Momose, K., Inoue, T., & Roubik, D.W. (1999) Preference in flower visits and partitioning in pollen dietsof stingless bees in an Asian tropical rain forest. Researches on Population Ecology (Kyoto), 41, 195–202.

Nieh, J.C. (2004) Recruitment communication in stingless bees (Hymenoptera, Apidae, Meliponini). Apidologie, 35,159–182.

Nieh, J.C., Tautz, J., Spaethe, J., & Bartareau, T. (2000) The communication of food location by a primitive stingless bee,Trigona carbonaria. Zoology (Analysis of Complex Systems), 102, 238–246.

Nogueira-Neto, P. (1949) India and stingless bees. Indian Bee Journal, 11, 121–122.Nogueira-Neto, P. (1951) A stingless bee variety of upper India recognized. Indian Bee Journal, 13, 132.Nogueira-Neto, P. (1954) Notas bionomicas sobre Meliponineos. III. Sobre a enxameagem (Hymenoptera, Apoidea).

Arquivos do Museu Nacional, 42, 419–451.Nogueira-Neto, P. (1970) A criação de abelhas indígenas sem ferrão, 2. ed. Tecnapis, São Paulo, 365.Nogueira-Neto, P. (1997) Vida e criação de abelhas indígenas sem ferrão. Edição Nogueirapis, São Paulo, 446.



Nurse, C.G. (1907) A new species of Indian wax-producing bee. Journal of the Bombay Natural History Society, 17, 619.Okada, H., Kubo, S., & Mori, Y. (1996) Pollination system of Neuwiedia veratrifolia Blume (Orchidaceae, Apostasio-

ideae) in the Malesian wet tropics. Acta Phytotaxonomica et Geobotanica, 47, 173–181.Oldroyd, B.P., Rinderer, T., & Wongsiri, S. (1992) Pollen resource partitioning by Apis dorsata, A. cerana, A. andrenifor-

mis and A. florea in Thailand. Journal of Apicultural Research, 31, 3–7.Oliveira, M.L., Morato, E.F., & Garcia, M.V.B. (1995) Diversidade de espécies e densidade de ninhos de abelhas sociais

sem ferrão (Hymenoptera, Apidae, Meliponinae) em floresta de terra firme na Amazonia Central. Revista Brasileirade Zoologia, 12, 13–24.

Ornduff, R. (1991) Size classes, reproductive behavior, and insect associates of Cycas media (Cycadaceae) in Australia.Botanical Gazette, 152, 203–207.

Oshy, K.C.K. & Arikumar, D.H. (2000) Flowering incidences and breeding system in Bambusa vulgaris. Current Sci-ence (India), 79, 1650–1652.

Packard, A.S. (1867) The home of the bees. American Naturalist, 1, 364–378.Pagden, H.T. (1957) Notes of communal flight activites of stingless bees. Malayan Nature Journal, 12, 15–19.Palmer, K.A., Oldroyd, B.P., Quezada-Euán, J.G., Paxton, R.J., & May-Itza, W.d.J. (2002) Paternity frequency and

maternity of males in some stingless bee species. Molecular Ecology, 11, 2107–2113.Pande, Y.D. & Sumita, B. (1987("1985")a) The foraging behaviour of honey bees on flowers of pigeon pea (Cajanus

cajan) in Agartala, Tripura. Indian Bee Journal, 47, 13–15.Pande, Y.D. & Sumita, B. (1987("1985")b) Effect of fenitrothion on the foraging activity of honey bees on Cajanus cajan

in Tripura. Indian Bee Journal, 47, 42–43.Parish, C.S.P. (1866) Pwai-ngyet. Science-Gossip, 2, 198–200.Pascoe, F.P. (1863) (F.P. Pascoe, Esq., V.P., in chair) This very interesting insect was obtained by Mr. T. W. Woodbury ...

Transactions of the Entomological Society of London, 11, 171.Pedro, S.R.M. & Camargo, J.M.F. (2003) Meliponini neotropicais: o gênero Partamona Schwarz, 1939 (Hymenoptera,

Apidae). Revista Brasileira de Entomologia, 47, 1–117.Peeters, C. & Ito, F. (2001) Colony dispersal and the evolution of queen morphology in social Hymenoptera. Annual

Review of Entomology, 46, 601–630.Percy, A.P. (1989) Division of Trigona iridipennis Smith colony. Indian Bee Journal, 51, 149.Peters, J.M., Queller, D.C., Imperatriz-Fonesca, V.L., Roubik, D.W., & Strassmann, J.E. (1999) Mate number, kin selec-

tion and social conflicts in stingless bees and honeybees. Proceedings of the Royal Society of London, Series B, 266,379–384.

Phadke, R.P. (1968) Studies on Indian honeys. 3. Proximate composition and physico-chemical characterization of hon-eys from the wild honey bees, Apis dorsata, Apis florea and Trigona. Indian Bee Journal, 30, 3–8.

Pick, R.A. & Blochtein, B. (2002) Atividades de vôo de Plebeia saiqui (Holmberg) (Hymenoptera, Apidae, Meliponini)durante o período de postura da rainha e em diapausa. Revista Brasileira de Zoologia, 19, 827–839.

Pooley, A.C. & Michener, C.D. (1969) Observations on nest of stingless honeybees in Natal (Hymenoptera: Apidae).Journal of the Entomological Society of Southern Africa, 32, 423–430.

Portugal-Araújo, V.d. & Kerr, W.E. (1959) A case of sibling species among social bees. Revista Brasileira de Biologia,19, 224–228.

Prasad, J.N. & Hemanth, J. (1992) Pond heron Ardeola grayii (Sykes) feeding on bees. Journal of the Bombay NaturalHistory Society, 89, 246.

Prell, H. (1919) Menschenschädel als Bienenwohnung (Hym.). Entomologische Mitteilungen, 8, 157–162.Pugh, B.M. (1947) Varieties of bees found in Assam (Extract from the report of the Assistant Agriculture Officer, North

East Frontier Agency, Assam, on "Agriculturein the Mynner (Jirang) state, Khasi Hills, Assam"). Indian Bee Journal, 9, 62.

Raffudin, R. & Crozier, R.H. (2007) Phylogenetic analysis of honey bee behavioral evolution. Molecular Phylogeneticsand Evolution, 43, 543–552.

Rajitparinya, T. (2000) Ecology and diversity of stingless bees (Hymenoptera: Apidae) in Chiang Mai.Raju, A.J.S. (2005) Passerine bird pollination and seed dispersal in Woodfordia floribunda Salisb. (Lythraceae), a com-

mon low altitude woody shrub in the eastern Ghats forests of India. Ornithological Science (Japan), 4, 103–108.Raju, A.J.S. & Esradanam, V. (2002) Pollination ecology and fruiting behaviour in a monoecious species, Jatropha cur-

cas L. (Euphorbiaceae). Current Science (India), 83, 1395–.Raju, A.J.S. & Rao, S.P. (2003) Decline of bee pollinators in Visakhaptnam region. Insect Environment, 9, 2–3.Raju, A.J.S. & Rao, S.P. (2006) Explosive pollen release and pollination as a function of nectar-feeding activity of cer-

tain bees in the biodiesel plant, Pongamia pinnata (L.) Pierre (Fabaceae). Current Science (India), 90, 960–967.Raju, A.J.S., Rao, S.P., & Jonathan, K.H. (2006) Andromonoecy, insect pollination and fruiting behaviour in Acacia cae-

sia (L.) Willd. (Mimosaceae) in the Eastern Ghats. Current Science (India), 91, 939–943.Raju, A.J.S. & Reddi, C.S. (1989) Pollination biology of Anisomeles indica and A. malabarica (Lamiaceae). Plant Spe-

cies Biology, 4, 157–167.



Raju, A.J.S., Zafar, R., & Rao, S.P. (2005) Floral device for obligate selfing by remote insect activity and anemochory inWrightia tinctoria (Roxb.) R.Br. (Apocynaceae). Current Science (India), 88, 1378–1380.

Ramanujam, C.G.K., Fatima, K., & Kalpana, T.P. (1993("1992")) Nectar and pollen sources for dammer bee (Trigona iri-dipennis Smith) in Hyderabad (India). Indian Bee Journal, 55, 25–28.

Ranger, S. & O'Donnell (1999) Genotypic effects on forager behavior in the neotropical stingless bee Partamona bilin-eata (Hymenoptera: Meliponidae). Naturwissenschaften, 86, 187–190.

Rao, S.P., Atluri, J.B., & Subba Reddi, C. (1999("1997")) Crataeva magna (Lour.) D.C. as a pollen source for the honeybees. Indian Bee Journal, 59, 85–87.

Rasmussen, C. (2004) A stingless bee nesting with a paper wasp (Hymenoptera: Apidae, Vespidae). Journal of the Kan-sas Entomological Society, 77, 593–601.

Rasmussen, C. (2008). Molecular phylogeny of stingless bees: Insights into divergence times, biogeography, and nestarchitecture evolution (Hymenoptera: Apidae: Meliponini). Ph.D. Dissertation, University of Illinois at Urbana-Champaign, USA, iii + 301 pp.

Rasmussen, C. & Ascher, J.S. (2008) Heinrich Friese (1860–1948): Names proposed and notes on a pioneer melittologist(Hymenoptera, Anthophila). Zootaxa, 1833, 1–118.

Rasmussen, C. & Camargo, J.M.F. (2008) A molecular phylogeny and the evolution of nest architecture and behavior inTrigona s.s. (Hymenoptera: Apidae: Meliponini). Apidologie, 39, 102–118.

Rasmussen, C. & Cameron, S.A. (2007) A molecular phylogeny of the Old World stingless bees (Hymenoptera: Apidae:Meliponini) and the non-monophyly of the large genus Trigona. Systematic Entomology, 32, 26–39.

Rayment, P.T. (1930) New and remarkable bees. Proceedings of the Royal Society of Victoria, 43, 42–61.Rayment, P.T. (1932a) The stingless bees of Australia. 1. Introduction. Victorian Naturalist, 48, 183–189.Rayment, P.T. (1932b) The stingless bees of Australia. 2. The architecture. Victorian Naturalist, 48, 203–212.Rayment, P.T. (1932c) The stingless bees of Australia. 3. Behaviour of the individual. Victorian Naturalist, 48, 246–254.Rayment, P.T. (1932d) The stingless bees of Australia. 4. Food of the larval bee. Victorian Naturalist, 49, 9–14.Rayment, P.T. (1932e) The stingless bees of Australia. 5. Is there a symbiotic association? Victorian Naturalist, 49, 39–

43.Rayment, P.T. (1932f) The stingless bees of Australia. 6. The finding of a new species. Victorian Naturalist, 49, 104–107.Rayment, P.T. (1935) A cluster of bees; sixty essays on the life-histories of Australian bees, with specific descriptions of

over 100 new species, and an introduction by E.F. Phillips. Endeavour, Sydney, 752.Rayment, P.T. (1939) Bees from the high lands of New South Wales and Victoria. Australian Zoologist, 9, 263–294.Rayment, P.T. (1953) The distribution of the bee genus, Trigona. Western Australian Naturalist, 4, 22–23.Rayment, P.T. (1961) A tiny bee and a very large flower. Proceedings of the Royal Zoological Society of New South

Wales, 1958–1959, 96–97.Rinderer, T.E., Marx, B.D., Gries, M., & Tingek, S. (1996) A scientific note on stratified foraging by Sabahan bees on the

yellow flame tree (Peltophorum pterocarpum). Apidologie, 27, 423–425.Rocha, M.P., Pompolo, S.d.G., & Campos, L.A.O. (2003). Citogenética da tribo Meliponini (Hymenoptera, Apidae). In:

G.A.R. Melo & I. Alves-dos-Santos (Eds.), Apoidea neotropica: Homenagem aos 90 anos de Jesus Santiago Moure,Editora UNESC, Criciúma, pp. 311–320.

Roubik, D.W. (1989) Ecology and natural history of tropical bees. Cambridge University Press, New York, 514.Roubik, D.W. (1990a) Mate location and mate competition in males of stingless bees (Hymenoptera: Apidae: Meliponi-

nae). Entomologia Generalis, 15, 115–120.Roubik, D.W. (1990b). Niche preemption in tropical bee communities: a comparison of neotropical and Malesian faunas.

In: S.F. Sakagami, R. Ohgushi & D.W. Roubik (Eds.), Natural History of social wasps and bees in equatorialSumatra, Hokkaido University Press, Sapporo, pp. 245–257.

Roubik, D.W. (1990c). Nesting and defence in tropical honeybees and stingless bees: you are what eats you? In: G.K.Veeresh, B. Mallik & C.A. Viraktamath (Eds.), Social insects and the environment, proceedings of the 11th interna-tional congress of IUSSI, 1990 (International union for the study of social insects), Oxford, New Delhi, pp. 662–663.

Roubik, D.W. (1992). Loose niches in tropical communities: why are there so few bees and so many trees? In: M.D.Hunter, P.W. Price & T. Ohgushi (Eds.), Effects of resource distribution on animal-plant interactions, AcademicPress, San Diego, pp. 327–354.

Roubik, D.W. (1993) Direct costs of forest reproduction, bee-cycling and the efficiency of pollination modes. Journal ofBioscience, 18, 537–552.

Roubik, D.W., ed. (1995) Pollination of cultivated plants in the tropics. Vol. 118, pp x+1–196. FAO, Rome.Roubik, D.W. (1996). Wild bees of Brunei Darussalam. In: D.S. Edwards, W.E. Booth & S.C. Choy (Eds.), Tropical

rainforest research — current issues: proceedings of the conference held in Bandar Seri Begawan, April 1993. Vol.74, Kluwer, Dordrecht, pp. 59–66.

Roubik, D.W. (2005) Honeybees in Borneo. In Pollination ecology and the rain forest, Sarawak studies eds D.W. Roubik,S. Sakai & A.A. Hamid, Vol. 174, pp. 89–103. Springer.



Roubik, D.W. (2006) Stingless bee nesting biology. Apidologie, 37, 124–143.Sakagami, S.F. (1959) Stingless bees collected by Prof. S. Matsumura from Singapore (Hymenoptera, Apidae). Insecta

Matsumurana, 22, 119–121.Sakagami, S.F. (1960) Some bees of Apinae and Xylocopinae collected in Cambodia. Kontyû, 28, 146–147.Sakagami, S.F. (1963) On the male of Trigona (Tetragona) fimbriata Smith (Hymenoptera: Apidae). Journal of the New

York Entomological Society, 71, 39–44.Sakagami, S.F. (1971) Ethosoziologischer Vergleich zwischen Honigbienen und stachellosen Bienen. Zeitschrift für

Tierpsychologie, 28, 337–350.Sakagami, S.F. (1975) Stingless bees (excl. Tetragonula) from the continental Southeast Asia in the collection of Bernice

P. Bishop museum, Honolulu (Hymenoptera, Apidae). Journal of the Faculty of Science, Hokkaido University,Series VI, Zoology, 20, 49–76.

Sakagami, S.F. (1978) Tetragonula stingless bees of the continental Asia and Sri Lanka (Hymenoptera, Apidae). Journalof the Faculty of Science, Hokkaido University, Series VI, Zoology, 21, 165–247.

Sakagami, S.F. (1982). Stingless bees. In: H.R. Hermann (Ed.), Social insects. Vol. 3, Academic Press, New York, pp.361–423.

Sakagami, S.F. (1989) [Original title in Japanese=How does the myrmecophilous stingless bee Trigona (Trigonella)moorei, invade ants' nests?]. Insectarium (Insekutaryumu: Ikita Konchu no Zasshi), 26, 164–175.

Sakagami, S.F., Camillo, C., & Zucchi, R. (1973) Oviposition behavior of a Brazilian stingless bee, Plebeia (Friesella)schrottkyi, with some remarks on the behavioral evolution in stingless bees [IIX]. Journal of the Faculty of Science,Hokkaido University, Series VI, Zoology, 19, 163–189.

Sakagami, S.F. & Inoue, T. (1985) Taxonomic notes on three bicolorous Tetragonula stingless bees in Southeast Asia.Kontyû, 53, 174–189.

Sakagami, S.F. & Inoue, T. (1987) Stinglee bees of the genus Trigona (subgenus Trigonella) with notes on the reductionof spatha in male genitalia of the subgenus Tetragonula (Hymenoptera, Apidae). Kontyû, 55, 610–627.

Sakagami, S.F. & Inoue, T. (1989) Stingless bees of the genus Trigona (subgen. Geniotrigona) (Hymenoptera, Apidae),with description of T. (G.) incisa sp.nov. from Sulawesi. Japanese Journal of Entomology, 57, 605–620.

Sakagami, S.F. & Inoue, T. (1990). Oviposition behavior of two Sumatran stingless bees, Trigona (Tetragonula) laevi-ceps and T. (T.) fuscobalteata [XIII]. In: S.F. Sakagami, R. Ohgushi & D.W. Roubik (Eds.), Natural History of socialwasps and bees in equatorial Sumatra, Hokkaido University Press, Sapporo, pp. 201–217.

Sakagami, S.F., Inoue, T., & Salmah, S. (1985). Key to the stingless bee species found or expected from Sumatra. In:Sumatra Nature Study (Entomology) group (Ed.), Evolutionary ecology of insects in humid tropics, especially incentral Sumatra, Kanazawa University, Kanazawa (Japan), pp. 37–43.

Sakagami, S.F., Inoue, T., & Salmah, S. (1990). Stingless bees of central Sumatra. In: S.F. Sakagami, R. Ohgushi & D.W.Roubik (Eds.), Natural History of social wasps and bees in equatorial Sumatra, Hokkaido University Press, Sap-poro, pp. 125–137.

Sakagami, S.F., Inoue, T., Yamane, S., & Salmah, S. (1983a) Nest architecture and colony composition of the Sumatranstingless bee Trigona (Tetragonula) laeviceps. Kontyû, 51, 100–111.

Sakagami, S.F., Inoue, T., Yamane, S., & Salmah, S. (1989) Nests of the myrmecophilous stingless bee, Trigona moorei:how do bees initiate their nest within an arboreal ant nest. Biotropica, 21, 265–274.

Sakagami, S.F. & Ito, M. (1981) Specific and subgeneric variations in tibial corbiculation of male bumblebees(Hymenoptera: Apidae), an apparently functionless character. Entomologica Scandinavica, 15, 365–376.

Sakagami, S.F. & Khoo, S.G. (1987) Taxonomic status of the Malesian stingless bee Trigona reepeni, with discovery ofTrigona pagdeni from northern Malaya. Kontyû, 55, 207–214.

Sakagami, S.F. & Yamane, S. (1984) Notes on the taxonomy and nest architecture of the Taiwanese stingless bee Trigona(Lepidotrigona) ventralis hoozana. Bulletin of the Faculty of Education, Ibaraki University (Natural Sciences), 33,37–48.

Sakagami, S.F. & Yamane, S. (1987) Oviposition behavior and related notes of the Taiwanese stingless bee Trigona (Lep-idotrigona) ventralis hoozana [XII]. Journal of Ethology, 5, 17–27.

Sakagami, S.F., Yamane, S., & Hambali, G.G. (1983b) Nests of some southeast Asian stingless bees. Bulletin of the Fac-ulty of Education, Ibaraki University (Natural Sciences), 32, 1–21.

Sakagami, S.F., Yamane, S., & Inoue, T. (1983c). Intranidal behaviors of Trigona (Tetragonula) laeviceps and T. (Trigo-nella) moorei. In: Sumatra Nature Study (Entomology) group (Ed.), Ecological study on social insects in CentralSumatra with special reference to wasps and bees, pp. 46–51.

Sakagami, S.F., Yamane, S., & Inoue, T. (1983d) Oviposition behavior of two southeast Asian stingless bees, Trigona(Tetragonula) laevices [=minangkabau] and T. (T.) pagdeni [XI]. Kontyû, 51, 441–457.

Sakagami, S.F. & Yoshikawa, K. (1961) Bees of Xylocopinae and Apinae collected by the Osaka city University biolog-ical expedition to Southeast Asia 1957–58, with some biological notes. Nature and Life in Southeast Asia, 1, 409–444.

Salmah, S., Inoue, T., Mardius, P., & Sakagami, S.F. (1987) Incubation period and post–emergence pigmentation in the



sumatran stingless bee, Trigona (Trigonella) moorei. Kontyû, 55, 383–390.Salmah, S., Inoue, T., & Sakagami, S.F. (1984) Relationship between age sequence and pigmentation in the stingless bee

Trigona (Tetragonula) laeviceps. Journal of Apicultural Research, 23, 55–58.Salmah, S., Inoue, T., & Sakagami, S.F. (1990). An analysis of apid bee richness (Apidae) in central Sumatra. In: S.F.

Sakagami, R. Ohgushi & D.W. Roubik (Eds.), Natural History of social wasps and bees in equatorial Sumatra,Hokkaido University Press, Sapporo, pp. 139–174.

Salmah, S., Inoue, T., & Sakagami, S.F. (1991) Incubation period in the sumatran stingless bee Trigona (Tetragonula)minangkabau. Treubia, 30, 195–201.

Salmah, S., Inoue, T., & Sakagami, S.F. (1996) Incubation period and post-emergence pigmentation in the Sumatranstingless bee Trigona (Heterotrigona) itama (Apidae, Meliponinae). Japanese Journal of Entomology, 64, 401–411.

Salt, G. (1929) A contribution to the ethology of the Meliponinae. Transactions of the Entomological Society of London,77, 431–470.

Samejima, H., Marzuki, M., Nagamitsu, T., & Nakasizuka, T. (2004) The effects of human disturbance on a stingless beecommunity in a tropical rainforest. Biological Conservation, 120, 577–587.

San, B.K. & Osawa, N. (2002) Seasonal variation and community structure of tropical bees.Sandhouse, G.A. (1943) The type species of the genera and subgenera of bees. Proceedings of the United States National

Museum, 92, 519–619.Santos-Filho, P.d.S., Alves, D.d.A., Eterovic, A., Imperatriz-Fonseca, V.L., & Kleinert, A.d.M.P. (2006) Numerical

investment in sex and caste by stingless bees (Apidae: Meliponini): a comparative analysis. Apidologie, 37, 207–221.

Schmid, J. (1964) Zur Frage der Störung des Bienengedächtnisses durch Narkosemittel, zugleich ein Beitrag zur Störungder sozialen Bindung durch Narkose. Zeitschrift für vergleichende Physiologie, 47, 559–595.

Schmidt, V.M., Zucchi, R., & Barth, F.G. (2003) A stingless bee marks the feeding site in addition to the scent path(Scaptotrigona aff. depilis). Apidologie, 34, 237–248.

Schneider, E.L. (1982) Observations on the pollination biology of Nymphaea gigantea W. J. Hooker (Nymphaeaceae).Western Australian Naturalist, 15, 71–72.

Schulz, W.A. (1907) Die indoaustralische Trigona laeviceps F. Sm. und ihr Nest. Zeitschrift für wissenschaftliche Insek-tenbiologie, 3, 65–73.

Schulz, W.A. (1909) Ein javanisches Nest von Trigona canifrons F. Sm. in einem Bambusstabe. Zeitschrift für wissen-schaftliche Insektenbiologie, 5, 338–341.

Schwarz, H.F. (1932) The genus Melipona. The type genus of the Meliponidae or stingless bees. Bulletin of the AmericanMuseum of Natural History, 63, 231–460.

Schwarz, H.F. (1937) Results of the Oxford University Sarawak (Borneo) expedition: Bornean stingless bees of thegenus Trigona. Bulletin of the American Museum of Natural History, 73, 281–328.

Schwarz, H.F. (1939a) The Indo-Malayan species of Trigona. Bulletin of the American Museum of Natural History, 76,83–141.

Schwarz, H.F. (1939b) Two Trigona bees collected by Prof. Teiso Esaki on the Palau and East Caroline Islands. Mushi,12, 151–152.

Schwarz, H.F. (1945) The wax of stingless bees (Meliponidae) and the uses to which it has been put. Journal of the NewYork Entomological Society, 53, 137–144.

Schwarz, H.F. (1948) Stingless bees (Meliponidae) of the western hemisphere. Lestrimelitta and the following subgeneraof Trigona: Trigona, Paratrigona, Schwarziana, Parapartamona, Cephalotrigona, Oxytrigona, Scaura, andMourella. Bulletin of the American Museum of Natural History, 90, 1–546.

Schwarz, H.F. (1951) Stingless bees collected by R. Takahashi in Indo-China and Siam (Hymenoptera). Mushi, 22, 63.Serrão, J.E. (2001) A comparative study of the proventricular structure in corbiculate Apinae (Hymenoptera, Apidae).

Micron, 32, 379–385.Sharma, H.K. (2004). Cash crops farming in the Himalayas: the importance of pollinators and pollination in vegetable

seed production in Kullu valley of Himachal Pradesh, India. International Centre for Integrated Mountain Develop-ment, Kathmandu (Nepal).

Slaa, E.J., Chaves, L.A.S., Malagodi-Braga, K.S., & Hofstede, F.E. (2006) Stingless bees in applied pollination: practiceand perspectives. Apidologie, 37, 293–315.

Smiley, R.L., Baker, E.W., & Delfinado-Baker, M. (1996) New species of Hypoaspis (Acari: Mesostigmata: Laelapidae)from the nest of a stingless bee in Malaysia (Hymenoptera: Meliponinae, Apidae). Anales del Instituto de Biología,Universidad Nacional Autónoma de México. Serie Zoología, 67, 197–203.

Smith, F. (1854) Catalogue of the Hymenopterous Insects in the Collection of the British Museum. Part II, Apidæ. Brit-ish Museum (Natural History), London, 197.

Smith, F. (1857) Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca; and atSingapore, by A. R. Wallace. Journal of the Proceedings of the Linnean Society, Zoology, 2, 42–88, pl. 1–2.

Smith, F. (1858) Mr. Smith exhibited some specimens of a small species of Trigona from Moulmein... Transactions of



the Entomological Society of London, 9, 98–99.Smith, F. (1859) Catalogue of Hymenopterous insects collected by Mr. A. R. Wallace at the Islands of Aru and Key.

Journal of the Proceedings of the Linnean Society, Zoology, 3, 132–178.Smith, F. (1863a) Prof. Westwood also exhibited specimens of a beetle ... parasitic in the nest of Trigona carbonaria ...

Proceedings of the Entomological Society of London, 11, 171.Smith, F. (1863b) (Frederick Smith, Esq., President, in the chair) The president exhibited the nest of Trigona carbonaria

... Transactions of the Entomological Society of London, 11, 174.Smith, F. (1863c) (Frederick Smith, Esq., President, in the chair) On the nest of Trigona carbonaria. Transactions of the

Entomological Society of London, 11, 181–182.Smith, F. (1865) Descriptions of new species of Hymenopterous insects from the Islands of Sumatra, Sula, Gilolo, Sal-

watty, and New Guinea, collected by Mr. A.R. Wallace. Journal of the Proceedings of the Linnean society, Zoology,8, 61–94, plate 4.

Smith, F. (1871) A catalogue of the Aculeate Hymenoptera and Ichneumonidae of India and the eastern archipelago. ByF. Smith, with introductory remarks by A. R. Wallace. Journal of the Proceedings of the Linnean Society, Zoology,11, 285–415.

Smith, F. (1878) List of Hymenoptera obtained by Mr. Ossian Limborg east of Maulmain, Tenasserim provinces, duringthe months december 1876, january, march and april 1877, with descriptions of new species. Journal of the AsiaticSociety of Bengal (Physical Science), 47, 167–169.

Smythe, R. (1970) Pollination by the common native bee. The Orchadian (Australasian Native Orchid Society), 3, 128,149.

Snelling, R.R. (1996). Insects part 1: The social Hymenoptera; Social Hymenoptera collected during the survey; Naturalhistory notes on the social Hymenoptera. In: A.L. Mack (Ed.), A biological assessment of the Lakekamu basin,Papua New Guinea. Vol. 9, Conservation International, Washington, D.C., pp. 39–47, 131–139, 140–143.

Spinola, M. (1853) Compte rendu des Hyménoptères inédits provenant du voyagae entomologique de M. Ghiliani dansle Para en 1846. Memorie della Accademia delle Scienze di Torino, [2]13, 19–94.

Starr, C.K. & Sakagami, S.F. (1987) An extraordinary concentration of stingless bee colonies in the Philippines, withnotes on nest structure (Hymenoptera: Apidae: Trigona spp.). Insectes Sociaux, 34, 96–107.

Stow, A., Briscoe, D., Gillings, M., Holley, M., Smith, S., Leys, R., Silberbauer, T., Turnbull, C., & Beattie, A. (2007)Antimicrobial defences increase with sociality in bees. Biology Letters, 3, 422–424.

Strand, E. (1911) Zur Kenntnis papuanischer und australischer Hymenopteren, insbesondere Schlupfwespen. Internatio-nale Entomologische Zeitschrift, 5, 86–87.

Strand, E. (1912) Biologische Notiz über papuanische Trigonen. Internationale Entomologische Zeitschrift, 6, 11.Strand, E. (1913a) Apidae con Ceylon, gesammelt 1899 von Herrn Dr. W. Horn. Archiv für Naturgeschichte. Abteilung

A, 79, 135–150.Strand, E. (1913b) [H. Sauter's Formosa-Ausbeute] Apidae I (Hym.). Supplementa Entomologica, 2, 23–67.Strand, E. (1915) Beiträge zur Systematik und insbesondere zur Verbreitung der Apidae. Archiv für Naturgeschichte.

Abteilung A, 81(11), 124–139.Suka, T. & Inoue, T. (1993) Nestmate recognition of the stingless bee Trigona (Tetragonula) minangkabau (Apidae:

Meliponinae). Journal of Ethology, 11, 141–147.Suka, T., Inoue, T., & Yamaoka, R. (1990). Chemical cues for recognition of kin and nestmates in stingless bees. In: G.K.

Veeresh, B. Mallik & C.A. Viraktamath (Eds.), Social insects and the environment, proceedings of the 11th interna-tional congress of IUSSI, 1990 (International union for the study of social insects), Oxford, New Delhi, pp. 521.

Sung, I.-H., Chiang, C.-H., Ho, K.-K., & Yamane, S. (2006a) [Original title in Japanese=A review on the history of bee-keeping in Taiwan and the development of its research works (Hymenoptera, Apidae, Apinae)]. 8, 129–147.

Sung, I.-H., Lin, M.-Y., Chang, C.-H., Cheng, A.-S., & Chen, W.-S. (2006b) Pollinators and their behaviors on mangoflowers in southern Taiwan. Formosan Entomologist, 26, 161–170.

Sung, I.-H., Yamane, S., Ho, K.-K., & Chen, W.-S. (2006c) [Original title in Japanese=Geographic distribution and nest-ing sites of the Taiwanese stingless bee Trigona ventralis hoozana and an unidentified subspecies of Asian honeybeeApis cerana in Taiwan (Hymenoptera: Apidae)]. Japanese Journal of Entomology, N.S., 9, 33–45.

Sung, I.-H., Yamane, S., Ho, K.-K., Wu, W.-J., & Chen, Y.-W. (2004) Morphological caste and sex differences in the Tai-wanese stingless bee Trigona ventralis hoozana (Hymenoptera: Apidae). Entomological Science, 7, 263–269.

Tadauchi, O., Ito, M., Kojima, J.-I., & Toda, M.J. (1998) Species and subspecies described by the late professor Dr. Sh. F.Sakagami (Insecta: Hymenoptera). Natural History Bulletin of Ibaraki University, 2, 229–246.

Tangmitcharoen, S., Pianhanuruk, P., & Cunningham, R. (2002) [Original title in Thai=Pollination ecology in relation tofruit setting of teak (Tectona grandis Linn.f.): natural and plantation forests]. In Proceedings of the Forestry Confer-ence, pp. 202–214.

Tangmitcharoen, S., Takaso, T., Siripatanadilox, S., Tasen, W., & Owens, J.N. (2006) Behavior of major insect pollina-tors of teak (Tectona grandis L. f.): a comparison of clonal seed orchard versus wild trees. Forest Ecology and Man-agement, 222, 67–74.



Temaru, E., Shimura, S., Amano, K., & Karasawa, T. (2007) Antibacterial activity of honey from stingless honeybees(Hymenoptera; Apidae; Meliponinae). Polish Journal of Microbiology, 56, 281–285.

Tennent, J.E. (1859) Ceylon, an account of the island physical, historical, and topographical with notices of its naturalhistory, antiquities and productions. Longman, London, 619.

Tennent, J.E. (1861) Sketches of the natural history of Ceylon with narratives and anecdotes illustrative of the habits andinstincts of the Mammalia, birds, reptiles, fishes, insects, &c., including a monograph of the elephant and a descrip-tion of the modes of capturing and training it with engravings from original drawings. Longman, London, xxiii, 500.

Tezuka, T. & Maeta, Y. (1995) Pollen robbing behaviors observed in two species of introduced stingless bees(Hymenoptera, Apidae). Japanese Journal of Entomology, 63, 759–762.

Thomson, V.P., Nicotra, A.B., & Cunningham, S.A. (2004) Herbivory differentially affects male and female reproductivetraits of Cucumis sativus. Plant Biology, 6, 621–628.

Thummajitsakul, S., Klinbunga, S., Smith, D., & Sittipraneed, S. (2008) Genetic diversity and population structure ofTrigona pagdeni Schwarz in Thailand. Apidologie, 39, 446–455.

Tóth, E., Queller, D.C., Dollin, A.E., & Strassmann, J.E. (2004) Conflict over male parentage in stingless bees. InsectesSociaux, 51, 1–11.

Tóth, E., Strassmann, J.E., Imperatriz-Fonseca, V.L., & Queller, D.C. (2003) Queens, not workers, produce the males inthe stingless bee Schwarziana quadripunctata quadripunctata. Animal Behaviour, 66, 359–368.

Tsuji, K., Hasyim, A., Nakamura, H., & Nakamura, K. (2004) Asian weaver ants, Oecophylla smaragdina, and theirrepelling of pollinators. Ecological Research, 19, 669–673.

Uji, T. (1987) Penyerbukan pada rambutan (Nephelium lappaceum var. lappaceum) [=Pollination in Nephelium lappa-ceum var. lappaceum]. Berita Biologi, 32, 31–34.

Utami, N. (1986) Penyerbukan pada sagu (Metroxylon sagu) [=Pollination in Metroxylon sagu]. Berita Biologi, 3, 229–231.

Utami, N. & Kahono, S. (1989) Penyerbukan pada pinang yaki (Areca vestiaria) di kebun raya Bogor [=Pollination onthe pinang yaki (Areca vestiaria Giseke) in the Bogor Botanical Gardens]. Berita Biologi, 3, 470–472.

van Veen, J.W. & Sommeijer, M.J. (2000) Colony reproduction in Tetragonisca angustula (Apidae, Meliponini). InsectesSociaux, 47, 70–75.

Velthuis, H.H.W., Koedam, D., & Imperatriz-Fonseca, V.L. (2005) The males of Melipona and other stingless bees, andtheir mothers. Apidologie, 36, 169–185.

Veth, P.J. & Ritsema, C. (1892). Natuurlijke historie. Tiende afdeeling. Hymenoptera en Hemiptera. In: P.J. Veth (Ed.),Midden-Sumatra. Reizen en onderzoekingen der Sumatra-expeditie, uitgerust door het Aardrijkskundig genootsc-hap, 1877–1879, beschreven door de leden der expeditie, onder toezicht van Prof. P.J. Veth. Vol. 4 (Natuurlijke his-torie, fauna), E.J. Brill, Leiden, pp. 1–23.

Viraktamath, S., Lingappa, S., Kallesha, G.R., Holi, K.S., & Hunur, S. (1999) Comparative studies on pollen sources toTrigona sp. Apis mellifera L. and Apis cerana F. in Dharwad, Karnataka, India. Indian Bee Journal, 61, 71–73.

Wagner, A.E. & Briscoe, D.A. (1983) An absence of enzyme variability within two species of Trigona (Hymenoptera).Heredity, 50, 97–103.

Wagner, A.E. & Dollin, L.J. (1982a) Probing the mysteries of the N.S.W. native bees. The Australasian Beekeeper, 83,157–161.

Wagner, A.E. & Dollin, L.J. (1982b) Swarming in Australian native bees — help solve the mystery! The AustralasianBeekeeper, 84, 34–38.

Wagner, A.E. & Dollin, L.J. (1982c) North Queensland's native bees — the little Aussie battlers. The Australasian Bee-keeper, 84, 70–72.

Wakefield, N.A. (1962) Insect swarms. Victorian Naturalist, 79, 11.Walker, F. (1860) Characters of some apparently undescribed Ceylon insects. Annals and Magazine of Natural History,

[3]5, 304–311.Wallace, H.M., Howell, M.G., & Lee, D.J. (2008) Standard yet unusual mechanisms of long-distance dispersal: seed dis-

persal of Corymbia torelliana by bees. Diversity and Distributions, 14, 87–94.Wallace, H.M., Maynard, G.V., & Trueman, S.J. (2002) Insect flower visitors, foraging behaviour and their effectiveness

as pollinators of Persoonia virgata R. Br. (Proteaceae). Australian Journal of Entomology, 41, 55–59.Wallace, H.M. & Trueman, S.J. (1995) Dispersal of Eucalyptus torelliana seeds by the resin-collecting stingless bee,

Trigona carbonaria. Oecologia, 104, 12–16.Waterhouse, C.O. (1900) Mr. C. O. Waterhouse exhibited a tube which formed the entrance to a nest of a Trigona, sent

from Singapore by Mr. H. N. Ridley... Transactions of the Entomological Society of London, 48, v.Waterhouse, C.O. (1903) Notes on the nests of bees of the genus Trigona. Transactions of the Entomological Society of

London, 51, 133–136, pl. 6.Wattanachaiyingcharoen, W. & Jongjitvimol, T. (2007) First record of the predator, Pahabengkakia piliceps Miller, 1941

(Reduviidae, Harpactorinae) in the stingless bee, Trigona collina Smith, 1857 (Apidae, Meliponinae) in Thailand.Natural History Journal of Chulalongkorn University, 7, 71–74.



Wattanachaiyingcharoen, W., Jongjitvimol, T., Boonthavon, K., Sripromma, M., Ruangridee, M., & Chutiyarat, S. (2004)[Original title in Thai=Species diversity and nesting sites of stingless bees in Sup-Lungka wild life conservationarea, Lopburee province]. NU (Naresuan University) Science Journal, 1, 63–74.

Welch, D. (1995) Beeswax rock art in the Kimberley, Western Australia. Rock Art Research: The Journal of the Austra-lian Rock Art Research Association, 12, 23–28.

Westwood, J.O. (1840). Entomology [Vol. 6]. Bees, comprehending the uses and economical management of the honey-bee of Britain and other countries, together with descriptions of the known wild species. In: W. Jardine (Ed.), Thenaturalist's library. Vol. 26, W.H. Lizars, Edinburgh, pp. viii+17–301, 30 pls.

White, D., Cribb, B.W., & Heard, T.A. (2001) Flower constancy of the stingless bee Trigona carbonaria Smith(Hymenoptera: Apidae: Meliponini). Australian Journal of Entomology, 40, 61–64.

Wijesekara, A. (2001) An annotated list of bees (Hymenoptera: Apoidea: Apiformis) of Sri Lanka. Tijdschrift voor Ento-mologie, 144, 145–158.

Wille, A. (1961) Evolutionary trends in the ventral nerve cord of the stingless bees (Meliponini). Revista de BiologíaTropical, 9, 117–129.

Wille, A. (1962) A new species of stingless bee (Meliponini) from Costa Rica. Revista de Biología Tropical, 10, 177–181.

Wille, A. (1977) A general review of the fossil stingless bees. Revista de Biología Tropical, 25, 43–46.Wille, A. (1979a) A comparative study of the pollen press and nearby structures in the bees of the family Apidae. Revista

de Biología Tropical, 27, 217–222.Wille, A. (1979b) Phylogeny and relationships among the genera and subgenera of the stingless bees (Meliponinae) of

the world. Revista de Biología Tropical, 27, 241–277.Wille, A. (1983) Biology of the stingless bees. Annual Review of Entomology, 28, 41–64.Wille, A. & Michener, C.D. (1973) The nest architecture of stingless bees with special reference to those of Costa Rica.

Revista de Biología Tropical, 21(supl.), 9–278.Williams, F.X. (1928) The natural history of a Philippine nipa house with descriptions of new wasps. Philippine Journal

of Science, 35, 53–118.Williams, G. & Adam, P. (1993) Ballistic pollen release in Australian members of the Moraceae. Biotropica, 25, 478–

480.Williams, G. & Walker, K. (2003) Pollination of the wet forest herb Pollia crispata (Commelinaceae). Cunninghamia, 8,

141–146.Williams, G.A. & Adam, P. (1997) The composition of the bee (Apoidea: Hymenoptera) fauna visiting flowering trees in

New South Wales lowland subtropical rainforest remnants. Proceedings of the Linnean Society of New South Wales,118, 69–95.

Wittmann, D., Bego, L.R., Zucchi, R., & Sakagami, S.F. (1991) Oviposition behavior and related aspects of the stinglessbees. XIV. Plebeia (Mourella) caerulea, with comparative notes on the evolution of the oviposition patterns (Api-dae, Meliponinae). Japanese Journal of Ecology, 59, 793–809.

Wu, Y. (2000). [Original title in Chinese=Hymenoptera Melittidae Apidae]. In: Fauna Sinica, Insecta. Vol. 20, SciencePress, Beijing, pp. 1–442, pls. 1–9.

Yamane, S. (1996) [Original title in Japanese=Social wasps in Australia]. Nature and Insects, 31, 10–15.Yamane, S., Heard, T.A., & Sakagami, S.F. (1995) Oviposition behavior of the stingless bees (Apidae, Meliponinae).

XVI. Trigona (Tetragonula) carbonaria endemic to Australia, with a highly integrated oviposition process. Japa-nese Journal of Entomology, 63, 275–296.

Yasumatsu, K. (1935) Notes on two Apoidea collected by Prof. Tohru Uchida on the Mariana and West Caroline islands.Mushi, 8, 94–96.

Yong, H.-S. (1986) Allozyme variation in the stingless bee Trigona fuscobalteata (Hymenoptera, Apidae) from Peninsu-lar Malaysia. Comparative Biochemistry and Physiology, Series B: Biochemistry and Molecular Biology, 83, 627–628.

Yong, H.-S. (1991) Biochemical genetic differentiation in the Trigona atripes species group of Malaysian stingless bees(Insecta: Hymenoptera: Apidae). Comparative Biochemistry and Physiology, Series B: Biochemistry and MolecularBiology, 99, 625–628.

Yong, H.-S. (1993). Genetic diversity and relationship of Trigona stingless bees of the subgenus Lepidotrigona in penin-sular Malaysia. In: L.J. Connor, T. Rinderer, H.A. Sylvester & S. Wongsiri (Eds.), Asian apiculture: proceedings ofthe first International conference on the Asian honey bees and bee mites, Wicwas Press, Cheshire, Connecticut, pp.51–56.

Yong, H.-S., Khoo, S.G., Sarjan, R., & Tho, Y.P. (1987) Glucose phosphate isomerase, malate dehydrogenase and isoci-trate dehydrogenase allozyme variation in the stingless bee, Trigona (Tetragonula) fuscobalteata (Insecta:Hymenoptera: Apidae), from Peninsular Malaysia. Comparative Biochemistry and Physiology, Series B: Biochemis-try and Molecular Biology, 87, 465–467.

Yoshikawa, K., Ohgushi, R., & Sakagami, S.F. (1969) Preliminary report on entomology of the Osaka city University 5th



scientific expedition to Southeast Asia 1966. With descriptions of two new genera of stenogastrine waps by J. vander Vecht. Nature and Life in Southeast Asia, 6, 153–182.

Yumoto, T. (2000) Bird-pollination of three Durio species (Bombacaceae) in a tropical rainforest in Sarawak, Malaysia.American Journal of Botany, 87, 1181–1188.

Zeuner, F.E. & Manning, F.J. (1976) A monograph on fossil bees (Hymenoptera: Apoidea), edited with an appendix bySamuel Francis Morris. Bulletin of the British Musem (Natural History), series Geology, 27, 149–268.

Zherikhin, V.V. & Ross, A.J. (2000) A review of the history, geology and age of Burmese amber (Burmite). Bulletin ofthe Natural History Museum (Geology), 56, 3–10.

Zoological Society of London, ed. (1864–present) The Zoological Record [former Record of Zoological Literature]. J.V.Vorst, London.

Documents

Rasmussen2008 7750 Catalog