Rasmussen Catalog Kelulut Indo Malayan PDF

TERMS OF USE
This pdf is provided by Magnolia Press for private/research use.

Commercial sale or deposition in a public library or website is prohibited.
ZOOTAXA
1935
Catalog of the Indo-Malayan/Australasian stingless bees

(Hymenoptera: Apidae: Meliponini)
CLAUS RASMUSSEN
Magnolia Press
Auckland, New Zealand
TERMS OF USE
Claus Rasmussen
Catalog of the Indo-Malayan/Australasian stingless bees (Hymenoptera: Apidae: Meliponini)
(Zootaxa 1935)
80 pp.; 30 cm.
17 Nov. 2008
ISBN 978-1-86977-295-6 (paperback)
ISBN 978-1-86977-296-3 (Online edition)
FIRST PUBLISHED IN 2008 BY

Magnolia Press
P.O. Box 41-383
Auckland 1346
New Zealand
e-mail: zootaxa@mapress.com
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)
2 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

TERMS OF USE
Zootaxa 1935: 180 (2008) ISSN 1175-5326 (print edition)

www.mapress.com / zootaxa/
Copyright 2008 Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
Catalog of the Indo-Malayan/Australasian stingless bees

(Hymenoptera: Apidae: Meliponini)
CLAUS RASMUSSEN
Department of Entomology, University of Illinois, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, IL 61801, USA.
E-mail: alrunen@yahoo.com
Table of contents
Abstract ............................................................................................................................................................................... 4
Catalog format ..................................................................................................................................................................... 4
Taxonomy and classification ............................................................................................................................................... 5
Distribution and biology ..................................................................................................................................................... 6
Acknowledgments ............................................................................................................................................................... 7
Catalog of the Indo-Malayan/Australasian stingless bees .................................................................................................. 8
Austroplebeia ............................................................................................................................................................... 8
Geniotrigona .............................................................................................................................................................. 11
Heterotrigona ............................................................................................................................................................. 13
Homotrigona .............................................................................................................................................................. 15
Lepidotrigona ............................................................................................................................................................. 17
Lisotrigona ................................................................................................................................................................. 21
Lophotrigona .............................................................................................................................................................. 23
Odontotrigona ............................................................................................................................................................ 24
Papuatrigona ............................................................................................................................................................. 24
Pariotrigona ............................................................................................................................................................... 25
Platytrigona ............................................................................................................................................................... 26
Sundatrigona .............................................................................................................................................................. 27
Tetragonilla ................................................................................................................................................................ 28
Tetragonula ................................................................................................................................................................ 31
Tetrigona .................................................................................................................................................................... 50
Meliponini indet ......................................................................................................................................................... 53
Incertae sedis ............................................................................................................................................................ 53
Nomina nuda .............................................................................................................................................................. 54
Appendix 1. Plants visited by stingless bees ..................................................................................................................... 55
Appendix 2. Stingless bee taxon names ............................................................................................................................ 59
Appendix 3. ...................................................................................................................................................................... 61
References ......................................................................................................................................................................... 62
Accepted by E. Almeida: 16 Oct 2008; published: 17 Nov. 2008 3

TERMS OF USE
Abstract
This catalog of the published literature on stingless bees from the Indo-Malayan/Australasian region was compiled from
520 references and deals with 129 proposed names for species-group taxa of which 89 are considered currently valid.
Information is summarized on organisms associated with stingless bee colonies as well as the 225 plants from which the
bees have been reported to forage. Lectotypes are designated for Trigona ornata described by Rayment and the following
species described by Friese: Trigona anamitica, T. australis, T. bornensis, T. flaviventris, T. keyensis, T. luteiventris, T.
pygmaea, T. reepeni, T. sericea, and T. versicolor.
Key words: Austroplebeia, Geniotrigona, Heterotrigona, Homotrigona, Lepidotrigona, Lisotrigona, Lophotrigona,

Odontotrigona, Papuatrigona, Pariotrigona, Platytrigona, Sundatrigona, Tetragonilla, Tetragonula, Tetrigona, Trigona,
bibliography
Stingless bees (Hymenoptera: Apidae: Meliponini) are important pollinators of native plants in tropical and
subtropical parts of the world (Heard 1999a). They form large perennial colonies with complex social 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 & Camargo
2008), 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 an
index to previous studies in taxonomy, behavioral research, and pollination ecology, thus consolidating the
existing knowledge in an accessible format. The extensive Neotropical fauna was recently cataloged by
Camargo & Pedro (2007: 391 spp.) and a systematic revision of the Afrotropical stingless bees included a
nearly complete bibliography (Eardley 2004: 26 spp. incl. Madagascar). The present catalog treats the remain-
ing Indo-Malayan/Australasian stingless bees (129 proposed species-group names, 89 currently accepted
valid species) and completes efforts to catalog the global stingless bee fauna. Together with recent molecular
phylogenies of the stingless bees (Rasmussen & Cameron 2007; Rasmussen 2008), it is expected that these
tools will spark renewed scientific interest in this diverse group of bees.
Catalog format
The catalog is sorted alphabetically by genus-group and currently valid species-group names; subspecies are
not recognized in the catalog. Genus-group headings are in boldface type. Following the accepted species-
group names are listed synonyms, including validly proposed junior synonyms and nomina nuda. References
are sorted in chronological order and include citation, page numbers from publications, type-specimen infor-
mation from both type-specimen labels, and the original publication where the species was first described, key
words, distribution, and floral records. Incertae sedis and nomina nuda are listed at the end of the catalog,
except when the identity is known for a nomen nudum. Page numbers are not listed for brief works or if the
particular species is treated throughout the publication. Publication dates were determined by the publishers
imprint date on the wrapper of the fascicle, when available, or information contained within the issue. When
the imprint year of the issue differed from the stated year of the volume, the latter are placed in parentheses.
Asterisks (*) following page numbers indicate where the species discussion is located in major works. Key
words in parenthesis after a reference indicate the kind of information provided and associated organisms
recorded. Misidentifications are corrected in the key word section whenever necessary or known, and 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 the
original publication. Distributional records are not listed comprehensively, but rather are limited to records

TERMS OF USE
cited in either taxonomic works or faunal inventories. Spellings of floral records were corrected to conform
with the International Plant Name Index (www.ipni.org). Appendix 1 lists all plant species (including family)
upon which stingless bees are known to forage in the region. Plant synonymies were not updated, but rather
left as cited in the original publication.
Stingless bee type specimens are encountered in museums worldwide, as indicated in the catalog. 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 are
also listed following each institution. Of particular importance are the collections in the Natural History
Museum (London), with 52 primary types, Museum fr Naturkunde der Humboldt-Universitt (Berlin), with
15, and National Museum of Natural History (Washington, D.C.), with 10. As for the most prolific describers
of new taxa, Theodore D.A. Cockerell described 38 new taxa, Heinrich Friese 17, Frederick Smith 14, Herbert
F. Schwarz and Peter Cameron 11 each, and Shichi 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 been
found in the collections indicated by the late Sakagami nor in his private collection (Tadauchi et al. 1998). It
may be that the types are in those collections by Tadauchi et al. (1998), but that they are not labeled as types.
The type of Trigona kusutkana Dover is missing from the Raffles Museum of Biodiversity Research where it
was originally deposited. All of the types of Smith have been accounted for, although as stressed by Baker
(1993), several of the specimens labeled as types in the Natural History Museum (London) may not be true
types and putative type specimens from London needs to be examined and compared with specimens from
University Museum of Natural History (Oxford) when in doubt. A number of species described before the
standardization of taxonomy with the International Code of Zoological Nomenclature (latest edition is 1999)
are represented by syntypes (multiple primary types). In several cases a syntype series consists of multiple
biological and morphological species. Lectotypes are here designated for several species proposed by Friese
as the identity of his true types is often confusing (see Rasmussen & Ascher 2008).
Literature has been updated until mid-2007. References cited include those found in Zoological Record
(Zoological Society of London 1864present) and all references cited within these records, totaling 520
papers. A number of popular travel accounts are listed in the nearly complete stingless bees bibliography by
Schwarz (1948). These were all consulted, but only included if they provided either the scientific name or
some description of the natural history of the bees, which was infrequently the case.
Taxonomy and classification
I have followed the classification proposed by Moure (1961) and Rasmussen & Cameron (2007) in not treat-
ing some of the Old World stingless bees as subgenera of Trigona Jurine (Table 1). The phylogenetic groups
of Rasmussen & Cameron (2007) are fully congruent with the systematic arrangement proposed by Moure
(1961). Therefore, all supraspecific groups are treated at the generic level as in the New World stingless bee
catalog (Camargo & Pedro 2007). Other studies propose a simpler classification to avoid a profusion of for-
mal names, which is equally valid when based on monophyletic lineages. This dual approach to classifying
genera (chiefly Moure 1961; Michener 1990, 2007a), however, complicates literature searches with specific
taxa appearing under several different generic names. The genus-group name or synonymy is listed for each
taxon in Appendix 2.
Most synonymies here are based on the published examination of types by Schwarz (1937; 1939a) and
Moure (1961). However, a number of species were only tentatively synonymized by them, and re-examination
of primary types was necessary for confirming and establishing their synonymies. In those cases, I indicated
the types I examined. Just like the inconsistent usage of stingless bee generic combinations, some taxa have
been treated as subspecies (e.g., Schwarz 1939a; Sakagami et al. 1990), species (e.g., Klakasikorn et al.
INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 5

TERMS OF USE
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 the
fact 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 specialized
keys, although the fauna has not yet undergone the taxonomic scrutiny comparable to that of many New
World taxa (e.g., Camargo & Moure 1994; Camargo & Moure 1996; Pedro & Camargo 2003b). While
Michener (2007a) included genus-group keys, the synonymy of a larger part of the regional fauna under the
polyphyletic subgenus Heterotrigona, sensu Michener, makes that key less useful. The only other 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 need
to be consulted for poorly characterized genera like Tetragonula. For these genera, regional keys can be useful
(Sakagami 1978; Dollin et al. 1997). No workable key exists for Austroplebeia (although see Rayment 1935:
735), Platytrigona (although see Michener 1990: 127), and some of the varieties here considered as valid spe-
cies.
Distribution and biology
The distribution of stingless bees in the Indo-Malay/Australasian region stretches from India to the Solomon
Islands and from China (Yunnan, Hainan, Taiwan: Wu 2000) to Australia (New South Wales). Schwarz (1937)
suggested that the greater abundance of stingless bees in Thailand and Malaysia, including all of Borneo, is
due to the abundance of resin-secreting trees (Dipterocarpaceae) and humid tropical climate, although most
likely an array of factors are responsible for this concentration of species.
Natural history observations summarized from the present catalog are as follow:
Colony pests: Achroea grisella (Lepidoptera: Tineidae), Ceria cf. ornatifrons (Diptera: Syrphidae) (larvae
feeding from provision), and Protaetia mandarinea (Coleoptera: Scarabaeidae).
Insects and other organisms inhabiting nests or inquilines: Brachypeplus auritus, B. basalis, B. meyricki,
B. planus (Coleoptera: Nitidulidae), Cerophagus trigona (Acari: Acaridae), Homalota trigonae (Coleoptera:
Staphylinidae), Hypoaspis hoffmannae (Acari: Laelapidae), Neocypholaelaps phooni (Acari: Ameroseiidae),
and Tribolium myrmecophilum (Coleoptera: Tenebrionidae).
Predators recorded from flowers and around nests: Amblyseius (Acarina: Phytoseiidae) (of inquilines?),
Ardeola grayii (Ciconiiformes: Ardeidae), Argiope keyserlingi, A. sp. (Araneae: Araneidae), Bembix flavipes,
B. musca (Hymenoptera, Crabronidae), Cypsiurus balasiensis (Apodidae, Apodiformes), Dicrurus macrocer-
cus, D. paradiseus (Dicruridae: Passeriformes), Dorniphora trigonae (Diptera: Phoridae) (of brood?), Helarc-
tos malayanus (Carnivora: Ursidae), Hermetia illucens (Diptera: Stratiomyidae) (of brood), Merops
leschenaulti (Coraciiformes: Meropidae), Nephila maculata (Araneae: Nephilidae), Oecophylla smaragdina,
O. sp. (Hymenoptera: Formicidae), Pahabengkakia piliceps (Hemiptera: Reduviidae), Phoridae (Diptera),
small birds, Solenopsis geminata (Hymenoptera: Formicidae), Syrphidae (Diptera), and Thomisus spectabilis
(Araneae: Thomisidae). A bee was also reported clinging to the leg of a large green cicada, Dundubia spic-
ulata (Hemiptera: Cicadidae), although the details of that interaction was not provided.
Nest sites: aerial Crematogaster nests (Hymenoptera: Formicidae), bamboo stems and interspaces, beetle
burrows, cavities among roots, coco nuts, cracks, crevices, holes in limestone, hollow pillars, hollow trunks
(including Durio zibethinus, Bombacaceae; Eugenia aromatica, Myrtaceae; Ficus, Moraceae; Nephelium lap-
paceum, Sapindaceae; Shorea laevis, Dipterocarpaceae), house posts, a human skull, and old tree stumps.

TERMS OF USE
TABLE 1. Different classificatory schemes used in the literature.
Genus: This work and Moure (1961) Type species Genus: Michener (2007)
Austroplebeia Moure, 1961 Trigona cassiae Austroplebeia
Geniotrigona Moure, 1961 Trigona thoracica Trigona (Heterotrigona)
Heterotrigona Schwarz, 1939 Trigona itama T. (Heterotrigona)
Homotrigona Moure, 1961 Trigona fimbriata T. (Homotrigona)
Lepidotrigona Schwarz, 1939 Trigona nitidiventris T. (Lepidotrigona)
Lisotrigona Moure, 1961 Melipona cacciae Lisotrigona
Lophotrigona Moure, 1961 Trigona canifrons T. (Heterotrigona)
Odontotrigona Moure, 1961 Trigona haematoptera T. (Heterotrigona)
Papuatrigona Michener & Sakagami, 1990 Trigona genalis T. (Papuatrigona)
Pariotrigona Moure, 1961 Trigona pendleburyi Pariotrigona
Platytrigona Moure, 1961 Trigona planifrons T. (Heterotrigona)
Sundatrigona Inoue & Sakagami, 1993 Trigona moorei T. (Heterotrigona)
Tetragonilla Moure, 1961 Trigona atripes T. (Heterotrigona)
Tetragonula Moure, 1961 Trigona iridipennis T. (Heterotrigona)
Tetrigona Moure, 1961 Trigona apicalis T. (Heterotrigona)
Lastly, floral records are frequently reported and show that the stingless bees are truly polylectic with 69
families and 179 genera of plants recorded as hosts in the Indo-Malayan/Australasian region. This includes
plants visited for nectar without performing pollination. The diversity of plants includes both cycads (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 to
Frank Koch for hosting me during a visit to the Museum fr Naturkunde, David Furth and Brian Harris for
hosting me multiple times at National Museum of Natural History, and Holger H. Dathe and Stephan Blank
for hosting a brief visit to the DEI. David Notton kindly provided much information and loan of types from
the Natural History Museum. J.M.F. Camargo gave me unlimited access to his comprehensive stingless bee
library and collection which was helpful in forming the foundation for the present catalog, Camargo and 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 manuscript
and Ying Wang helped with translation of Chinese. Visits to institutions were supported by a Francis M. and
Harlie M. Clark Research Support Grant and a National Science Foundation grant (DEB 0446325) to S. A.
Cameron who also encouraged this study.

TERMS OF USE
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.
Mller.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. Mller 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); Michener
1990: 134, 137 (illustration, morphology, systematic position); Cardale 1993: 318 (distribution, floral
record, list); AUSTRALIA, NE coastal; Queensland; New South Wales; SE coastal; Murray-Darling
basin; Floral record: Alphitonia; Angophora; Callistemon; Eucalyptus; Melaleuca; Rapistrum; 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); Dollin
1996b: (nest); Nogueira-Neto 1997: 85, 204, 334; Dollin & Heard 1997: (domestication); Dollin &
Heard 1997: 4 (bee-keeping); Dollin 1997: (characteristics); Drumond et al. 1999: (oviposition); Crane
1999: 89, 90; Dollin & Heard 1999: (bee-keeping); Michener 2000: 782-783; Drumond et al. 2000:
(reproduction); Heard & Dollin 2000: (bee-keeping); Green et al. 2001: 91 (microsatellite); Serro
2001: (morphology (proventricular structure)); Palmer et al. 2002: 2112 (citation); Koedam et al. 2002:
231, 232 (comparative note, wax); Green & Oldroyd 2002: 197 (citation); Dollin 2002a: (bee-keeping);
Dollin 2002b: (bee-keeping); Costa et al. 2003: (systematics); Heard 2003: (citation); Tth et al. 2004: 6
(citation); Heiling & Herberstein 2004: (predator (Thomisus spectabilis: Thomisidae: Araneae)); Slaa et
al. 2006: 301 (pollination); Michener 2007a: 806-807; Temaru et al. 2007: 281 (honey properties);
Klumpp 2007: 5, 6, 10, 11, 15-18, 29, 47, 51, 56, 57, 76, 77, 82.
Austroplebeia cassiae (Cockerell, 1910)
Trigona cassi 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;

TERMS OF USE
Rayment 1930: 59 (comparative note); Rayment 1932a: 185 (common name (koochee)); Rayment
1932d: (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); Michener
1997: 10 (type species); Michener 2000: 791; Nagamitsu & Inoue 2005: 76 (distribution, foraging);
Michener 2007a: 815.
Austroplebeia cincta (Mocsry in Friese, 1898)
Trigona cincta Friese 1898: 431: Holotype (HNHM, worker): examined, "Friedrich- / Wilh.-hafen", "N.
Guinea / Bir 96" (taxonomy); Type locality: PAPUA NEW GUINEA "Neu-Guinea (Friedrich Wil-
helmshafen)" (1 worker); Friese 1909("1908"): 354, 356 (distribution); INDONESIA, Irian Jaya/PAPUA
NEW GUINEA, Cyclopen Gebirge; Moaif; Moso; Sentani; Tawarin; Timena; Friese 1909: 272, 273 (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 (probably
misidentification); 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);
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. /

TERMS OF USE
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; Dollin
1997: (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); Rayment
1935: 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; Klumpp
2007: 5.
Austroplebeia ornata (Rayment, 1932)
Trigona cockerelli ornata subsp. nov. Rayment 1932f: 107: Lectotype (MVMA, worker): here designated,
"4", "TYPE", "Trigona / cockerelli Raym. / ornata / C. York Q.", "SYNTYPE / T-11575 / Trigona / cock-
erelli / ornata", "MUS. VIC. / ENTO 2008-8 L"; Paralectotypes (MVMA (3)) (taxonomy (as manuscript
name)); Type locality: AUSTRALIA "Cape York, Queensland (Donald Thomson)" (workers); Rayment
1935: 735 (key to species, taxonomy (described as new species again)); Moure 1961: 197 (possible syn-
onymy); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution, list); AUSTRALIA, NE
coastal; Queensland; Dollin 1997: (characteristics); Klumpp 2007: 5.
Austroplebeia percinta (Cockerell, 1929)
Trigona cincta subsp. percinta nov. Cockerell 1929e: 242: Syntypes (ZMHB, 4 workers): examined, labeled
by Friese as Trigona cincta? (comparative note (corrected identification of Cockerell 1910)); Type 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); Moure
1961: 197 (systematic position); Michener 1965: 230 (citation); Cardale 1993: 320 (distribution, list);
AUSTRALIA, Lake Eyre basin, Northern Territory; Dollin 1996b: (nest); Dollin 1997: (characteristics);
Green et al. 2001: 91 (microsatellite); Klumpp 2007: 5.
Austroplebeia symei (Rayment, 1932)
Trigona symei Rayment 1932f: 106: Holotype (ANIC, worker): examined, "TYPE", "Trigona / symesi Raym

TERMS OF USE
/ 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: Dendrobium
speciosum; Moure 1961: 197 (possible synonymy); Michener 1965: 230 (citation); Cardale 1993: 320
(distribution, floral record, list); AUSTRALIA, Queensland; Floral record: Dendrobium; Dollin 1997:
(characteristics); Drumond et al. 1999: (oviposition); Camargo et al. 2000: 9 (comparative notes, 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); Tth et al. 2004: 6 (citation); Rasmussen & Cameron 2007: 29, 31, 33, 34 (systematics);
Klumpp 2007: 5.
Austroplebeia websteri (Rayment, 1932)
Trigona websteri Rayment 1932f: 105: Holotype (ANIC, worker): examined, "TYPE", "Trigona / websteri
Raym. / WYNDHAM, W.A. / [below] 25/1/1931", "AUST. NAT. / INS. COLL."; paratype (ANIC (1))
(taxonomy); Type locality: AUSTRALIA "Wyndham, North-western Australia (U.N. Webster, M.D.,
25th January, 1931)" (worker); Rayment 1932f: 107 (taxonomy); Rayment 1935: 735 (key to species);
Moure 1961: 197 (possible synonymy); Michener 1965: 230 (citation); Cardale 1993: 320 (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 & Dollin
2000: (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, native
collector, ded. F. Dupont" (3 workers, holotype, paratypes); "Todyamboe, 900m, C. Celebes" (3 workers);
"Wuasa, Kab. Poso, Sulteng, Sulawesi" (9 workers); "Lorei Lindu Nat. Park, C. Sulawesi" (3 workers);
Tadauchi et al. 1998: 242-243 (type depositories); Rasmussen & Cameron 2007: (systematics).
Geniotrigona lacteifasciata (Cameron, 1902)
Trigona bornensis 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-

TERMS OF USE
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 to
species); Schwarz 1937: 283, 291, 317-319*, 326, plate 3, 7 (key to species, taxonomy); Schwarz 1939a:
105, 106 (synonymy); Schwarz 1948: 91; Moure 1961: 213 (systematic position); Sakagami 1975: 56
(distribution, variation); SINGAPORE; THAILAND; Camargo 1988: 370 (distribution, taxonomy);
MALAYSIA, Sandakan; Sakagami et al. 1990: 127 (synonymy).
Geniotrigona thoracica (Smith, 1857)
Trigona ambusta Cockerell 1918: 387: Holotype (BMNH 17b.1131) (distribution, key to species); Type
locality: MALAYSIA/SINGAPORE "Sandakan and Singapore" (unknown); Cockerell 1919c: 78 (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); Cockerell
1919c: 79 (distribution); Cockerell 1923: 240 (key to species); Dover 1929: 62 (citation, distribution);
MALAYSIA, Kuala Tahan; Pahang; SINGAPORE; Friese 1933a: 46 (comparative note); Schwarz 1937:
317, 318; Schwarz 1939a: 85, 90, 92, 104-106* (distribution, key to species, taxonomy); Floral record:
Citrus limon; Tephrosia candida; Schwarz 1948: 83, 104, 114, 118; Moure 1961: 212, 213 (systematic
position); Yoshikawa et al. 1969: 172, 173 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang;
Tapah; Touaran; Sakagami 1975: 56* (distribution); SINGAPORE; THAILAND; Wille 1979b: 247
(morphology); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral
record: Xerospermum intermedium; Sakagami et al. 1983b: 2, 4, 13-15, 16, 20 (honey, nest (in Shorea
laevis: Dipterocarpaceae)); Sakagami et al. 1985: (distribution, key to species); Baker & 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: Averrhoa
carambola; Liow 1999: 106, 127 (distribution); Nagamitsu et al. 1999: (floral records (not by species),
foraging); Heard 1999a: 191, 192, 195 (citation); Michener 2000: 801-802; Chinajariyawong & 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

TERMS OF USE
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" (comparative
notes, key to species); Type locality: MALAYSIA "Penang Island (Baker 9068)" (worker); Cockerell
1930a: 157 (distribution, floral record, variation); CAMBODIA, Tung Nui; Satul; Floral record: 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); Cameron
1908: 193, 194 (key to species); Cockerell 1920a: 115 (comparative note); Cockerell 1923: 242 (key to
species); Schwarz 1937: 291, 319-320*, 323, 326, 327, plate 3, 6 (key to species, taxonomy); Floral
record: Melastoma; Schwarz 1939a: 84, 85, 86, 90, 96, 100-101* (distribution, key to species, 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); Michener
1990: 120, 126, 127 (systematic position); Sakagami et al. 1990: 131, 135 (distribution, key to species);
Michener & Roubik 1993: 260 (comparative note, morphology); Roubik 1996: 61 (distribution); BRU-
NEI, Kuala Belalong; Bruijn & Sommeijer 1997: 44 (citation (as erythrogaster!)); Nagamitsu & Inoue
1998: (foraging, morphology); Nagamitsu et al. 1999: (floral records (not by species), foraging); Heard
1999a: 187 (citation (as erythrogaster!)); Samejima et al. 2004: (population); MALAYSIA, Sarawak,
Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76, 78, 79, 81, 83, plate 6 (distribution, foraging);
Rasmussen & Cameron 2007: (systematics).
Trigona luteiventris Friese 1909("1908"): 354, 358, fig. 15-5: Lectotype (ZMHB, worker): here designated,
"Malakka / Perak / 1912", "Trigona / luteiventris / 1904 Friese det. Fr / n Fr.", "Type" (red label), "Coll. /
Friese"; paralectotype (ZMHB (1)); possible additional types (AMNH) (taxonomy); Type locality:
MALAYSIA/PHILIPPINES "Perak (Malaka)"; "Palawan" (workers); Cockerell 1919c: 77 (key to 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);

TERMS OF USE
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); Cockerell
1919a: 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, Hill
stream; Schwarz 1937: 283, 292, 319, 322-325*, 327, plate 3, 6 (key to species, taxonomy); Floral record:
Melastoma; Schwarz 1939a: 84, 85, 86, 90, 92, 95, 96-100*, 127, 133 (distribution, key to species, taxon-
omy); Floral record: Mimosa pudica; Tephrosia candida; Sandhouse 1943: 558 (citation); Schwarz
1948: 118, 119; Sakagami 1959: 119 (distribution); SINGAPORE; Moure 1961: 201, 203 (morphology,
systematic position); Wille 1961: 117; Yoshikawa et al. 1969: 168, 172, 173, 174 (distribution, nest);
MALAYSIA, Cameron Highland, Pahang; Fraser's Hill, Selagor-Pahang; George Town, Penang; Sabah,
Jesselton; Sabah, Touaran; Sarawak, Kuching; Wille & Michener 1973: 14, 48 (nest); Sakagami 1975:
55* (distribution, variation); MALAYSIA; THAILAND; Wille 1979b: 265 (comparative note); Appanah
1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum
intermedium; Sakagami et al. 1983b: 2-4, 13, 19, 20 (nest); Sakagami et al. 1985: (distribution, key to
species); Inoue et al. 1985a: (aggressiveness, foraging, nest); Kiew 1985: (luring (unwashed shorts));
Inoue et al. 1985b: 45, 47, 48 (foraging); Baker & Delfinado-Baker 1985: 231 (inquiline (Neocypholae-
laps phooni: Ameroseiidae: Acari)); Atmowidjojo & Adisoemarto 1986: 227, 229 (pollination); Floral
record: 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, Sumatera
Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest (tree cavities and others)); Michener &
Roubik 1993: 260 (comparative note, morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra;
Cunha 1994: (systematics); Tezuka & Maeta 1995: (foraging, pollen robbing); Roubik 1995: 151
(antropogenic nest); Camargo & Moure 1996: 109 (taxonomy (type series)); Salmah et al. 1996: (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 et
al. 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);

TERMS OF USE
Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu &
Inoue 2005: 76, 83, 84 (distribution, foraging); Chinh et al. 2005: 368 (citation); Klakasikorn et al.
2005: 6 (citation); Roubik 2006: 131 (nest); Rasmussen & Cameron 2007: (systematics); Leonhardt et
al. 2007: (foraging); Floral record: Averrhoa carambola; Cassia fistula; Michener 2007a: 826-827; Wat-
tanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note).
Heterotrigona indet. Roubik 2006: 127, 132 (nest).
Homotrigona
Homotrigona aliceae (Cockerell, 1929)
Trigona alice Cockerell 1929b: 139, 140: Holotype (BMNH 17b.1116) (comparative note, taxonomy);
Type locality: THAILAND "Siam: On trail in jungle between Pahtoop mountain and Nan, Jan. 9, 1928
(Alice Mackie)" (worker); Schwarz 1937: 305, 306; Schwarz 1939a: 85, 92, 104* (distribution, key to
species, taxonomy); Schwarz 1948: 118; Schwarz 1951: 63 (distribution); CAMBODIA, Angkor;
Moure 1961: 201 (systematic position); Sakagami & Yoshikawa 1961: 434-435 (distribution, 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, paratype
worker); Schwarz 1939a: 104* (distribution, key to species, taxonomy); Moure 1961: 201 (systematic
position); Sakagami 1975: 54 (synonymy); Sakagami et al. 1990: 127 (synonymy).
Homotrigona anamitica (Friese, 1909)
Trigona anamitica Friese 1909("1908"): 358-359, fig 15-4: Lectotype (ZMHB, worker): here designated,
"Sd-Annam / Xom-Gom / Februar / H. Fruhstorfer", "Trigona / anamitica / 1904 Friese det. / Fr."; para-
lectotype (ZMHB (1)) (comparative note, taxonomy); Type locality: VIETNAM "Xom-Gom, Sd-
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); Sakagami
1975: 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, comparative
notes, key to species, taxonomy); Type locality: MALAYSIA "Sandakan, Borneo (Baker 9222)"
(worker); Cockerell 1919b: 243 (comparative notes); Cockerell 1920b: 228 (distribution); MALAYSIA,
Sandakan; Schwarz 1937: 289, 305, 306, 320, 327 (key to species); Schwarz 1939a: 103, 104 (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-

TERMS OF USE
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); Schwarz
1939a: 84, 85, 90, 92, 102-103*, 104, 106, 132 (distribution, key to species, taxonomy); Schwarz 1948:
73, 94; Moure 1961: 200, 201, 215 (systematic position); Sakagami & Yoshikawa 1961: 435 (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 (systematic
position); Sakagami et al. 1990: (distribution, key to species); CAMBODIA; INDONESIA, Sumatra;
MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest); INDONESIA, Sumatera
Barat; Inoue et al. 1990b: (population); Roubik 1993: 541 (illustration, nest); Michener & Roubik 1993:
260 (comparative note, morphology); Cunha 1994: (systematics); Roubik 1996: 61 (distribution); 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 Forest
Reserve; Eltz et al. 2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz
2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram
area; Wattanachaiyingcharoen et al. 2004: (diversity, nest); Nagamitsu & Inoue 2005: 76, 80, 82, 83,
84, 85, plate 9 (distribution, foraging); Klakasikorn et al. 2005: 3, 6 (floral record); Floral record: 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; Mimosa
pudica; Ruellia tuberose; Senna siamea; Thunbergia laurifolia; Tridax procumbens; Zingiber officinale;
Rasmussen & Cameron 2007: (systematics); Raffudin & Crozier 2007: 544, 545, 546 (behavior);
Michener 2007a: 826-827; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note).
Trigona flavistigma Cameron 1902: 130: Holotype (BMNH 17b.1097) (taxonomy); Type locality:
MALAYSIA "Kuching, Sarawak" (male); Cameron 1908: 193 (key to species); Cockerell 1923: 241
(key to species); Cockerell 1929b: 139 (comparative note); Schwarz 1937: 304, 305, 306, 307, 326;
Schwarz 1939a: 102, 103 (synonymy); Moure 1961: 201 (systematic position); Sakagami 1975: 52 (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)" (2
workers); Schwarz 1937: 304, 306, 327, plate 4; Schwarz 1939a: 102, 103, 104 (synonymy); Schwarz
1948: 118; Moure 1961: 201 (synonymy); Sakagami 1975: 52 (synonymy); Sakagami et al. 1990: 127
(synonymy).

TERMS OF USE
Homotrigona lutea (Bingham, 1897)
Trigona ferrea Cockerell 1929b: 139-140: Holotype (BMNH 17b.1117) (comparative note, taxonomy); Type
locality: THAILAND "Siam: Mekami River, Feb. 3, 1928 (Cockerell)" (worker); Schwarz 1937: 305,
306; Schwarz 1939a: 85, 92, 103 (distribution, key to species, taxonomy); Moure 1961: 201 (synonymy);
Sakagami 1975: 52, 54, 55 (distribution, variation); MALAYSIA; THAILAND; Sakagami et al. 1990:
127 (synonymy); Klakasikorn et al. 2005: 6 (citation).
Melipona lutea Bingham 1897: 559, 560, 564: Type (BMNH 17b.1122) (distribution, illustration, key to spe-
cies, taxonomy; notice this is no longer a senior homonym of Trigona mocsaryi lutea Friese 1903, see
Camargo & Pedro 2007: 472); Type locality: MYANMAR "Northern Tenasserim; Karennee" (worker);
Cockerell 1929b: 140 (comparative note); Moure 1961: 201 (systematic position); Wu 2000: 387, 390,
plate (key to species, morphology).
Homotrigona indet. Roubik 2006: 130 (nest).
Lepidotrigona
Lepidotrigona arcifera (Cockerell, 1929)
Trigona arcifera Cockerell 1929c: 591-592: Holotype (BMNH 17b.1081). The ventralis species group (tax-
onomy); Type locality: INDIA "Testa bridge, Himalayas, India, 10.1.97 (Sladen)" (worker); Schwarz
1937: 295; Schwarz 1939a: 85, 88, 134, 135 (distribution, key to species, taxonomy); Nogueira-Neto
1949: 121 (bee-keeping); Moure 1961: 199 (possible synonymy); Sakagami 1975: 67, 71 (synonymy);
Sakagami et al. 1990: 126 (synonymy); Wu 2000: 387 (synonymy).
Lepidotrigona doipaensis (Schwarz, 1939)
Trigona (Lepidotrigona) ventralis variety doipaensis Schwarz 1939a: 85, 94, 136*: Holotype (USNM 53563,
worker): examined, "Doi Pa / Mai Deng / Siam 750m / 12-29-32", "HughSmith / Coll", "Type No./ 53563
/ U.S.N.M.", "Holotype", "Trigona ventralis / var. doipaensis / H.F. Schwarz". The ventralis species group
(distribution, key to species, taxonomy); Type locality: THAILAND "SIAM.-Doi Pa, Mai Deng, 750
meters, Dec. 29, 1932 (H. M. Smith)" (worker); Moure 1961: 199 (systematic position); Sakagami 1975:
67, 69, 71 (distribution, variation); THAILAND, N; Sakagami & Yamane 1984: 35; Sakagami et al.
1990: 126 (synonymy); Wu 2000: 387 (synonymy); Klakasikorn et al. 2005: 3, 6 (floral record); Floral
record: Ficus religiosa; Homalium grandiflorum.
Lepidotrigona flavibasis (Cockerell, 1929)
Trigona flavibasis Cockerell 1929c: 592: Holotype (AMNH). The ventralis species group (taxonomy); Type
locality: THAILAND "Doi Sutep, Siam, Feb. 9 (Alice Mackie)" (worker); Schwarz 1939a: 85, 94, 134-
135*, 136 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199 (systematic
position); Yoshikawa et al. 1969: 174 (distribution); MALAYSIA, Fraser's Hill, Selagor-Pahang; 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 et
al. 2001: 91 (microsatellite); Chinh et al. 2005: 366, 368-371 (nest (as ventralis in part)); VIETNAM,
Cucphuong National Forest, Red river delta; Chinh & Sommeijer 2005: (reproduction (as ventralis in

TERMS OF USE
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), but
as it was proposed as a geograhical restricted (=subspecific) form before 1961, it is considered valid
(45.6.1). The ventralis species group (taxonomy); Type locality: TAIWAN "Hoozan, Februar" ('a row' of
workers); Sakagami et al. 1985: (key to species); Sakagami & Yamane 1987: (oviposition); Salmah et
al. 1990: 155 (altitude); Bravo 1992: 876 (citation, altitude); Yamane et al. 1995: 294 (comparative note);
Sung et al. 2004: (morphology); Nagamitsu & Inoue 2005: 76 (distribution, foraging); Chinh & 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); Friese
1914a: 13 (distribution); INDONESIA, Nusa Kambangan; Friese 1914b: 58 (nest); Cockerell 1929c: 591
(comparative note); Schwarz 1937: 295, 296; Schwarz 1939a: 85, 90, 94, 140* (distribution, key to 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 (as
ventralis, 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).

TERMS OF USE
Lepidotrigona nitidiventris (Smith, 1857)
Trigona fulvopilosella Cameron 1908: 193, 194: Syntypes (BMNH 17b.1101, 2 workers) (key to species,
taxonomy); Type locality: MALAYSIA "Kuching, May and October (John Hewitt)" (worker); Schwarz
1937: 285, 287, 293-294*, 295, 296, 326 (key to species, taxonomy); Schwarz 1939a: 133 (synonymy);
Moure 1961: 199 (possible synonymy); Sakagami 1975: 65 (citation).
Trigona nitidiventris Smith 1857: 50-51: Type (OUMNH (=Wilson Saunders collection)). The nitidiventris
species group (taxonomy); Type locality: MALAYSIA "Malacca (Mount Ophir)" (worker); Smith 1871:
395 (distribution); MALAYSIA, Malacca; Dalla Torre 1896: 582 (citation); Bingham 1905: 58 (citation,
distribution); Cockerell 1920a: 115 (comparative note); Cockerell 1923: 241, 242 (key to species);
MALAYSIA, Mt. Ophir; Schwarz 1937: 292, 293, 296, 327 (key to species); Schwarz 1939a: 85, 94,
132, 133*, 134 (distribution, key to species, taxonomy); Sandhouse 1943: 543 (citation); Schwarz 1948:
118; Moure 1961: 197, 199 (systematic position); Baltazar 1966: 407 (distribution); PHILIPPINES, 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 (comparative
note); Uji 1987: (pollination); INDONESIA, E Kalimantan; Floral record: Nephelium lappaceum;
Michener 1990: 121, 122 (systematic position); Sakagami et al. 1990: (distribution, key to species);
INDONESIA, Sumatra; MALAYSIA, Borneo; THAILAND; Salmah et al. 1990: (distribution, nest);
INDONESIA, Sumatera Barat; Inoue et al. 1990b: (population); Yong 1993: (biochemistry); Cunha
1994: (systematics); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue
1997: 434, 436 (pollination); Floral record: Santiria laevigata; Michener 1997: 32 (type species); 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, Pasoh
Forest Reserve; Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Klakasikorn et al. 2005: 6
(citation); Michener 2007a: 827; Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative
note).
Lepidotrigona palavanica (Cockerell, 1915)
Trigona palavanica Cockerell 1915: 2-3: Holotype (BMNH 17b.1120). The nitidiventris species group (com-
parative notes, taxonomy); Type locality: PHILIPPINES "P. Princesa, Palawan (Baker coll 3839)"
(worker); Cockerell 1919c: 77 (key to species); PHILIPPINES, Palawan, Puerto Princesa; Cockerell
1920b: 229 (distribution); PHILIPPINES, Palawan; Schwarz 1939a: 85, 86, 133 (distribution, key to 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; near
Nam Geeung; Nan; Pah Meesung Mt.; Pahtoop Mt.; Cockerell 1934: 62, fig. 9 (illustration (mandible));
Schwarz 1937: 295, 296; Schwarz 1939a: 136, 137 (synonymy); Moure 1961: 199 (synonymy).
Trigona terminata Smith 1878: 169: Type (BMNH 17b.1100). The terminata species group (taxonomy);
Type locality: MYANMAR no precise locality, presumably "Maulmain, Tenasserim Provinces"
(worker); Dalla Torre 1896: 584 (citation); Bingham 1897: 563 (comparative note, taxonomy); 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;

TERMS OF USE
Pagden 1957: 17 (comparative note); Sakagami 1960: 146 (distribution, variation); CAMBODIA, Snoul;
Stung Treng; Moure 1961: 199 (systematic position); Wille 1961: 117, 121; Sakagami & Yoshikawa
1961: 440 (distribution); LAOS, Thakhek; THAILAND, Mae Fack; Yoshikawa et al. 1969: 168, 169, 174
(distribution, nest); MALAYSIA, George Town, Penang; Wille & Michener 1973: 14, 25, 49, 60 (nest);
Sakagami 1975: 65-66*, 72, 73, 74 (distribution, variation); LAOS; THAILAND; VIETNAM; Sakag-
ami 1978: 239 (comparative note); Sakagami et al. 1983b: 2, 15-19, 20 (nest); Sakagami & Yamane
1984: 40, 47; Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts));
Sakagami & Yamane 1987: 24, 25 (comparative note); Camargo 1988: 371 (distribution, taxonomy);
INDONESIA, Java; MALAYSIA, Penang; Roubik 1989: 181; Michener 1990: 95, 105, 120, 122 (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 & Roubik
1993: 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); Liow
1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species), foraging); Michener
2000: 800; Rajitparinya 2000: (distribution, nest); Rajitparinya 2000: (distribution, nest); Wu 2000:
386, 388-390 (key to species, morphology); Eltz et al. 2001b: 343, 345 (foraging); MALAYSIA, Borneo,
Sabah; Green et al. 2001: 91 (microsatellite); Eltz et al. 2002a: 30, 31, 32 (distribution, foraging, nest,
population); MALAYSIA, Danum Valley; Sepilok; Deramakot; Floral record: Rhizophora apiculata;
Klein et al. 2002: 1010 (floral record); Floral record: Coffea canephora; San & Osawa 2002: (popula-
tion); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest, tree preference); Klein
et al. 2003b: 957 (floral record); Floral record: Coffea canephora; Klein et al. 2003a: 841 (floral record);
Floral record: Coffea canephora; Eltz 2004: 322 (abundance); Samejima et al. 2004: (population);
MALAYSIA, Sarawak, Miri, Upper Baram area; Wattanachaiyingcharoen et al. 2004: (diversity, nest);
Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Chinh et al. 2005: 368, 371 (citation);
Klakasikorn et al. 2005: 3, 6 (floral record); Floral record: Ficus religiosa; Irvingia malayana; Slaa et al.
2006: 297, 298 (citation); Tangmitcharoen et al. 2006: (pollination); Floral record: Tectona grandis;
Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, Kanchanaburi, Thong Pha Phum;
Rasmussen & Cameron 2007: (systematics); Michener 2007a: 824; Wattanachaiyingcharoen &
Jongjitvimol 2007: 73 (comparative note).
Lepidotrigona trochanterica (Cockerell, 1920)
Trigona trochanterica Cockerell 1920a: 115: Holotype (BMNH 17b.1102) (taxonomy); Type locality:
MALAYSIA "Sandakan, Borneo (Baker)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA,
Sandakan; Schwarz 1937: 285, 287, 292-293*, 295, 296, 327 (key to species, taxonomy); Schwarz
1939a: 85, 94, 133-134* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 199
(systematic position); Sakagami 1975: 65 (citation); Michener 1990: 122 (systematic position); 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-

TERMS OF USE
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); Schwarz
1939a: 85, 94, 134*, 135, 136, 139 (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure
1961: 199 (systematic position); Sakagami 1975: 66, 67-75* (distribution, morphology, variation); 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 (as
ventra! in part)); Nagamitsu et al. 1999: (floral records (not by species), foraging); Rajitparinya 2000:
(distribution, nest); Rajitparinya 2000: (distribution, nest); Wu 2000: 386, 387-388 (key to species, 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 & Cameron
2007: (systematics); Wattanachaiyingcharoen & Jongjitvimol 2007: 73 (comparative note).
Lepidotrigona indet. Drumond et al. 1996a: 399 (behavioral comparison); Roubik 2006: 127 (nest).
"Trigona (Lepidotrigona) ventralis" nigribasis forma nov. Sakagami 1975: 71: Nomen nudum. The name is
unavailable for taxonomic purposes, as it must be deemed to be infrasubspecific (ICZN, art. 15.2 and
45.6.3.) (variation); VIETNAM.
Melipona tunneli Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as no
description or definition of the taxon was provided (ICZN, art. 13.1.1). The statement that it "is different
from the other two in appearance as it possesses somewhat yellow spots in the abdomen", however, could
make it a nomen dubium, although most likely then a junior synonymon of arcifera (common name (ngap
siwor), honey, nest).
Lisotrigona
Lisotrigona cacciae (Nurse, 1907)
Melipona cacci Nurse 1907: 619: Lectotype (BMNH 17b.1103, worker); Type locality: INDIA "Hoshang-

TERMS OF USE
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);
Trigona scintillans Cockerell 1920a: 116: Holotype (BMNH 17b.1115) (taxonomy); Type locality:
MALAYSIA "Sandakan, Borneo (Baker)" (worker); Cockerell 1920b: 228 (distribution); MALAYSIA,
Sandakan; Cockerell 1929c: 590 (distribution, ecology ("great nuisance in camp setting on one's hands
and face")); CAMBODIA, Angkor; Pah Meeung Mt.; Pahtoop Mt.; Schwarz 1937: 289, 290, 307*, 327,
plate 7 (key to species, taxonomy); Schwarz 1939a: 85, 86, 94, 130*, 132 (distribution, key to species,
taxonomy); Schwarz 1948: 118; Moure 1961: 193, 195 (morphology, systematic position); Sakagami &
Yoshikawa 1961: 440 (distribution); THAILAND, Bangkok; Sakagami 1975: 51-52* (distribution, 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; Floral
record: 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: Ngh An Province: Khe
Bo, 19 03N 104 43E, 25-28 April 1998, 123 m, James M. Carpenter" (worker); VIETNAM, Ha Thing, SE
Huong Son; Jobiraj & Narendran 2004: 39 (key to species); Chinh et al. 2005: 363, 365-367, 370
(nest); VIETNAM, Cucphuong National Forest, Red river delta; Michener 2007b: 130 (comparative
note).
Lisotrigona furva Engel, 2000
Lisotrigona furva Engel 2000: 235-236: Holotype (SEMC); 5 paratypes (AMNH, SEMC) (floral record, key
to species, taxonomy); Type locality: THAILAND "THAILAND: Nakhon Ratchasima Prov.: Tha
Chang, 10 February 1993, S. Boongird and C. Michener" (worker); THAILAND, Chiang Mai, W Fang;
Floral record: Buddleja asiatica; Callistemon; Jobiraj & Narendran 2004: 40 (key to species);
Michener 2007b: (morphology).
Lisotrigona mohandasi Jobiraj & Narendran, 2004

TERMS OF USE
Lisotrigona mohandasi Jobiraj & Narendran 2004: 39, 40-43, fig. 1-4: Holotype ("Calicut", worker);
paratypes ("Calicut" (2)) (floral record, key to species, taxonomy); Type locality: INDIA "India, Kerala
Forest Research Institute (Peechi), K. Mohandas, 3-IV-2000" (3 workers); Floral record: Tectona grandis.
Lisotrigona indet. Engel 2000; Jobiraj & Narendran 2004: 39; Momose & Hamid 2005; Roubik 2006:
127, 128 (nest).
Lophotrigona
Lophotrigona canifrons (Smith, 1857)
Trigona busara Cockerell 1918: 387: Holotype (USNM 29496, worker): examined, "Singapore / Coll.
Baker", "9072", Type No. / 29496 / U.S.N.M.", "Trigona / busara / Ckll TYPE" (distribution, key to 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 (BMNH
17b.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 (comparative
notes, taxonomy, misidentification, uncertain identity); Ihering 1912: 3 (citation); Friese 1914b: 53, 54,
57 (identification, previous misidentification); Cockerell 1918: 385 (comparative notes); Cockerell
1919b: 244 (comparative note); Prell 1919: (nest (in human skull)); Cockerell 1922a: 279 (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; Floral
record: Nephelium lappaceum; Camargo 1988: 370 (synonymy); Roubik 1989: 105, 193; Michener
1990: 88, 97, 105, 120, 126, 127 (systematic position); Sakagami et al. 1990: (distribution, key to 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,

TERMS OF USE
morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cunha 1994: (systematics); Yamane et
al. 1995: 294 (synonymy); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Dollin et al. 1997:
861 (comparative note); Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: Santiria laevi-
gata; Michener 1997: 34 (type species); Nagamitsu & Inoue 1998: (foraging, morphology); Momose et
al. 1998: (pollination); Floral record: Dryobalanops aromatica; Nagamitsu et al. 1999: (floral records
(not by species), foraging); Michener 2000: 780, 800-801; San & Osawa 2002: (population); 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 Baram
area; Nagamitsu & Inoue 2005: 76, 83, 84 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation);
Roubik 2006: 127, 131 (nest); Inson & Malaipan 2006: (distribution, foraging, nest); THAILAND, 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 to
species, taxonomy); Schwarz 1948: 118; Moure 1961: 214 (synonymy).
Trigona hmatoptera 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); Camargo
1988: 371 (distribution); MALAYSIA, Borneo, Sandakan; Michener 1990: 127 (systematic position);
Sakagami et al. 1990: 131 (distribution); Cunha 1994: (systematics); Roubik 1996: 61 (distribution);
BRUNEI, Kuala Belalong; Michener 1997: 42 (type species); Nagamitsu et al. 1999: (floral records (not
by species), foraging); Michener 2000: 801; Eltz et al. 2002a: 30 (distribution, foraging, nest, popula-
tion); MALAYSIA, Danum Valley; Sepilok; Deramakot; Eltz et al. 2003: (nest, tree preference); Eltz
2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram
area; Nagamitsu & Inoue 2005: 76 (distribution, foraging); Rasmussen & Cameron 2007: (systemat-
ics); Leonhardt et al. 2007: (foraging); Floral record: Ardisia elliptica; Dillenia excelsa; Mallotus;
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" (red
label), "Coll. Friese", "Zool. Mus. / Berlin"; paralectotype (ZMHB (3)) (distribution, taxonomy); Type
locality: INDONESIA (IRIAN JAYA) "Manikion, 14.-28. Februar 1903 (Neu-Guinea)" (several work-

TERMS OF USE
ers); Friese 1909: 272, 273, 276-277 (citation, key to species); Schwarz 1932: 245; Schwarz 1948: 118;
Michener 1961: 23, 42 (aggressiveness, nest); PAPUA NEW GUINEA, Gulf District, Murua River near
Kerema; Michener 1965: 231 (citation); Wille & Michener 1973: 14, 24, 48, 59, 70 (nest); Michener
1990: 95, 97, 105, 122, 127, 153-157 (distribution, illustration, lectotype designation, morphology, sys-
tematic position); INDONESIA, Irian Jaya, Bodem; Bodem Sarmi area; Klamono oil fields; Waris S of
Hollandia; PAPUA NEW GUINEA, Dreikikir; Marua river near Kerema; Northeast New Guinea; One;
Sepik; Michener & Roubik 1993: 260 (comparative note, morphology); Snelling 1996: 131 (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: Kuala
Teku, 500 feet, Dec. 8, 1921 (H. M. Pendlebury)" (workers); "N. BORNEO.-Bettotan, nr. Sandakan, (C.
B. Kloss and H. M. Pendlebury)" (workers, July 28, 1927, and Aug. 15, 1927); Schwarz 1948: 118;
Moure 1961: 193-194, 195 (morphology, systematic position); Yoshikawa et al. 1969: 174 (distribution);
MALAYSIA, Fraser's Hill, Selagor-Pahang; Kiew 1985: (luring (unwashed shorts)); Michener 1990: 131
(systematic position); Michener 2001: 235 (synonymy); San & Osawa 2002: (population); MALAYSIA,
Negri Sembiran, Pasoh Forest Reserve; Inson & Malaipan 2006: (distribution, foraging, nest); 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 "Cameron
Highlands, Rhododendron Hill, 5000 feet (H. M. Pendlebury)" (workers, holotype, June 19,1923, and
June 22, 1923); MALAYA.-State of Pahang: Cameron Highlands, G. Tarbakar, 4500 feet, June 9, 1923 (H.
M. Pendlebury); Cameron Highlands, Bukit Lendong, 5000 feet, May 21, 1931 (H. M. Pendlebury);
Lubok Tamang, 3500 feet, March 5, 1924, and 4000 feet, June 10, 1923 (H. M. Pendlebury); Fraser's Hill,
4600 feet, Pine Tree Hill, Oct. 25, 1932 (H. M. Pendlebury); Schwarz 1948: 118; Moure 1961: 192, 193-
194, 195 (morphology, systematic position); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran,
Pasoh Forest Reserve; Floral record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key
to species); Michener 1990: 96, 100, 105, 129, 131 (illustration, morphology, systematic position); 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.

TERMS OF USE
Platytrigona
Platytrigona atricornis (Smith, 1865)
Trigona atricornis Smith 1865: 94: Holotype (OUMNH, worker): According to Baker (1993) the specimen is
labeled " 'N' [New Guinea (Allen); white disc] and 'Trigona atricornis' 'Smith' ". The atricornis species
group (taxonomy); Type locality: INDONESIA (IRIAN JAYA) "New Guinea" (worker); Smith 1871:
396 (distribution); "NEW GUINEA"; Gribodo 1893: 263 (comparative note); Dalla Torre 1896: 576
(citation); Friese 1898: 428, 429 (citation, key to species); "NEW GUINEA"; Friese 1898: 429 (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): here
designated, "Cyclops / geb. / 13.IV.03", "Trigona / flaviventris / 1904 Friese det. / Fr.", "Type" (red label),
"Coll. / Friese"; paralectotype (ZMHB (2)). The planifrons species group (taxonomy); Type locality:
INDONESIA (IRIAN JAYA) "Cyclopen-Gebirge, 3.-14. April 1903 (New Guinea)" (4 workers); Friese
1909: 272, 275 (citation, key to species); INDONESIA, Irian Jaya, Cyclopen Gebirge; Cockerell 1919b:
243 (comparative notes); Cockerell 1929e: 243 (comparative note); Schwarz 1932: 245; Schwarz 1937:
289, 320 (key to species); Schwarz 1948: 118; Moure 1961: 204 (systematic position); Michener 1965:
231 (citation); Michener 1990: 120, 127, 153 (systematic position); Michener 2000: 800; Hohmann
2003: (communication, nest); Michener 2007a: 824.
Platytrigona hobbyi (Schwarz, 1937)
Trigona hobbyi Schwarz 1937: 283, 288, 298-300*, 328: Holotype (BMNH 17b.1118). The hobbyi species
group (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, 4000 feet, moss 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 (comparative
note, morphology); Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Samejima et al. 2004:
(population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76 (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 / Khn", "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); Type
locality: INDONESIA "Key-Eilanden (Amboina, Nederland India) durch H. Khn" (6 workers); Friese
1909("1908"): 354, 356, fig. 15-7 (distribution); INDONESIA, Irian Jaya, Moaif; Timena; Friese 1909:
272, 273, 277 (key to species, taxonomy); INDONESIA, Irian Jaya, Moaif; Timena; Strand 1911: 87 (dis-
tribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Holl. N.G.; Pauwi; Samberi; Teba; Friese
1912: 94 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, D. Neu Guinea; Pauwi; Sam-

TERMS OF USE
beri; Teba; Strand 1912: (distribution); Friese 1915: 4 (distribution); INDONESIA, Irian Jaya/PAPUA
NEW GUINEA, Moaif; Pauwi; Samberi; Teba; Timena; Zoutbron; Schwarz 1948: 118; Michener 1965:
231 (citation); Michener 1990: 120, 127, 153 (systematic position); Michener & Roubik 1993: 260
(comparative note, morphology); Snelling 1996: 131 (distribution); Michener 2000: 800; Hohmann
2003: (communication, nest); Michener 2007a: 824.
Platytrigona lamingtonia (Cockerell, 1929)
Trigona lamingtonia Cockerell 1929e: 243: Holotype (AMS, worker): examined, "Mt. Lamington / Northern
Division / Papua v.1927 / C.T. McNamara", "Trigona / lamingtonia / Ckll TYPE", "HOLOTYPE", "Aus-
tralian Museum K 238299". The planifrons species group (taxonomy); Type locality: PAPUA NEW
GUINEA "Mt. Lamington, May, 1927 (C.T.McNamara)" (workers); Michener 1965: 231 (citation).
Platytrigona planifrons (Smith, 1865)
Trigona planifrons Smith 1865: 93-94: Holotype (OUMNH, worker): According to Baker (1993) the 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 species
group (taxonomy); Type locality: INDONESIA (IRIAN JAYA) "New Guinea" (worker); Smith 1871:
396 (distribution); "NEW GUINEA"; Dalla Torre 1896: 582 (citation); Friese 1898: 427, 428 (citation,
key to species); MALAYSIA, Malacca (misidentification); "NEW GUINEA"; Friese 1909("1908"): 354,
356, fig. 15-8 (distribution); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai;
Jamur Gebiet; Manikion; Manokwari; Merauke; Moaif; Tawarin; Friese 1909: 272, 273, 274 (distribu-
tion, key to species); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Digul Fluss; Etna Bai; Janour
Gebiet; Manokwari; Merauke, Moaif; Tawarin; Friese 1915: 4 (distribution); INDONESIA, Irian Jaya/
PAPUA NEW GUINEA, Digul Fluss; Etna Bai; Manikion; Manokwari; Merauke; Moaif; Sattelberg;
Sekofro; Simbang; Tawarin; Tjahe Fluss; Zoutbron; Cockerell 1923: 241 (key to species); Cockerell
1929e: 242 (comparative note, distribution); PAPUA NEW GUINEA, Fly River; Mt. Lamington;
Schwarz 1948: 83, 109, 118; Moure 1961: 203, 204 (systematic position); Michener 1965: 231 (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, not
located, nor in SEHU); Paratypes (RMNH, 3 workers, 2 males) (morphology, taxonomy); Type locality:
INDONESIA "N.E. Sumatra, Tongkoh, probably Mt. Talamau (Pasaman), VI 1941, v.d. M. Mohr leg." (5
workers, 4 males, holotype worker, paratypes); "Koto Atas, Solok, Sumatera Barat" (6 workers); 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

TERMS OF USE
et al. 1998: 244-245 (type depositories); Nagamitsu & Inoue 2005: 76 (distribution, foraging).
Sundatrigona moorei (Schwarz, 1937)
Trigona (Tetragona) matsumurai Sakagami 1959: 120-121: Holotype (SEHU, worker); 1 paratype (SEHU);
Type locality: SINGAPORE "Singapore, Sept., 22, 1932, S. Matsumura leg." (2 workers); Sakagami
1975: 56, 57 (synonymy); Sakagami et al. 1990: 127 (synonymy); Tadauchi et al. 1998: 243 (type
depositories).
Trigona moorei Schwarz 1937: 283, 292, 321-322*, 328, plate 3, 4: Holotype (BMNH 17b.1124); 1 paratype
(AMNH) (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt. Dulit, R. Lejok, near
sweat and water, Oct. 5, 1932" (2 workers); Schwarz 1939a: 85, 90, 106* (distribution, key to species,
taxonomy); Schwarz 1948: 83, 118; Sakagami 1959: 120-121 (morphology); Sakagami 1975: 56-58*
(distribution, morphology, variation); MALAYSIA; Sakagami 1978: 239 (comparative note); Appanah
1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral record: Xerospermum
intermedium; Sakagami et al. 1983c: (reproduction); Sakagami et al. 1985: (distribution, key to species);
Inoue et al. 1985a: (aggressiveness, foraging, nest (in aerial Crematogaster: Formicidae: Hymenoptera
nests)); Inoue et al. 1985b: 45, 47, 48 (foraging); Sakagami & Inoue 1987: 610-617, 618, 619, 620, 621,
622, 623, 624 (morphology); Salmah et al. 1987: (development); Sakagami & Inoue 1989: 615, 617,
618 (morphology); Roubik 1989: 133, 196, 197, 218; Sakagami et al. 1989: (nest); Sakagami 1989:
(nest); Michener 1990: 98, 126, 127, 153 (morphology, systematic position); Sakagami et al. 1990: (dis-
tribution, key to species); INDONESIA, Sumatra; MALAYSIA; Salmah et al. 1990: (distribution, nest (in
aerial Crematogaster: Formicidae: Hymenoptera nests)); INDONESIA, Sumatera Barat; Inoue et al.
1990b: (population); Inoue 1990: 582 (nest (Crematogaster: Formicidae: Hymenoptera nests), popula-
tion); Roubik 1990c: 663 (citation); Salmah et al. 1991: 199 (comparative note); Inoue & Sakagami
1993: 769 (citation); Salmah et al. 1996: 402, 404, 407, 409, 410 (comparative note); Drumond et al.
1996b: 40 (citation); Nogueira-Neto 1997: 381; Michener 1997: 56, 59 (type species, Trigonella as 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); Michener
2000: 801-802; Amano et al. 2000: (citation); Rasmussen 2004: 599 (citation); Samejima et al. 2004:
(population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 76, 83 (distri-
bution, foraging); Roubik 2006: 135 (nest); Rasmussen & Cameron 2007: (systematics); Michener
2007a: 826.
Sundatrigona indet. Inoue & Sakagami 1993: 769 (citation); Inoue & Sakagami 1993: 769 (homonym (for
Trigonella)); Roubik 2006: 127 (nest).
Tetragonilla
Tetragonilla atripes (Smith, 1857)
Trigona atripes Smith 1857: 50: Type (OUMNH (=Wilson Saunders collection)) (taxonomy); Type locality:
MALAYSIA "Malacca (Mt. Ophir)" (worker); Smith 1871: 395 (distribution); MALAYSIA, Malacca;
Dalla Torre 1896: 576 (citation); Bingham 1897: 560, 561 (distribution, key to species, taxonomy);
MALAYSIA, Malacca; MYANMAR, Mergui, S Tenasserim; Friese 1901: 271 (comparative note); 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:

TERMS OF USE
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; Sakagami
1978: 230 (comparative note); Wille 1979b: 273 (citation, illustration); Wille 1979a: (morphology (pollen
press)); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Floral
record: Xerospermum intermedium; Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (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); Michener
2007a: 826.
Tetragonilla collina (Smith, 1857)
Trigona cambodiensis Cockerell 1926a: 224: Holotype (USNM 29470, worker): examined (nest, taxonomy);
Type locality: CAMBODIA "Angkov (=Angkor) Wat, Cambodia, Jan. 11, 1926 (H.M. Smith)" (4 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.

TERMS OF USE
1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Sakagami & Khoo 1987:
207, 209, 210 (comparative note); Sakagami & Inoue 1987: 626 (comparative note); Roubik 1989: 133,
196, 243, 387; Sakagami et al. 1989: 269 (comparative note); Sakagami et al. 1990: (distribution, key to
species); CAMBODIA; INDONESIA, Sumatra; MALAYSIA; MYANMAR; THAILAND; Sakagami &
Inoue 1990: 212; Salmah et al. 1990: (distribution, nest (subterranean among roots)); INDONESIA,
Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest (cavities among roots), 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 et
al. 2001a: (pollen foraging); Eltz et al. 2001b: (foraging); MALAYSIA, Borneo, Sabah; Floral record:
Caesalpinia; Citrus; Clausena; Croton; Durio; Jasminium; Manihot esculenta; Mimosa pudica; 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: Diospyros
rhodocalyx; Dipterocarpus alatus; Ficus religiosa; Irvingia malayana; Jongjitvimol et al. 2005: (nest);
THAILAND, Phitsanulok; Roubik 2006: 131, 137 (nest); Tangmitcharoen et al. 2006: (pollination);
Floral record: Tectona grandis; Jongjitvimol et al. 2006: (foraging); Jongjitvimol & Wattanachaiying-
charoen 2006: (foraging); THAILAND, Phitsanulok; Floral record: Agave angustifolia; Alangium salvii-
folium; Butea monosperma; Caryota bacsonensis; Cassia bakeriana; Coccinia grandis; Costus
speciosus; 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; Torenia
fournieri; 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; Dicrurus
paradiseus: 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

TERMS OF USE
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); Sakagami
1975: 59, 60 (comparative note); Sakagami et al. 1985: (distribution, key to species); Camargo 1988:
371 (distribution); MALAYSIA, Borneo, Sandakan; Sakagami et al. 1990: 131, 136 (distribution); Yong
1991: (biochemistry); Michener & Roubik 1993: 260 (comparative note, morphology (as rufibasis!));
Roubik 1996: 61 (distribution); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433, 436 (pollina-
tion); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu
et al. 1999: (floral records (not by species), foraging); Nagamitsu & Inoue 2002: 305, 307, 308 (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); Sakagami
et al. 1983a: 106 (comparative note); Sakagami & Inoue 1985: 177 (comparative note); Sakagami et al.
1985: (distribution); Sakagami & Inoue 1987: 626 (comparative note); Sakagami et al. 1990: 130, 131
(comparative note, distribution); Wu 2000: 392 (synonymy); Nagamitsu & Inoue 2005: 77 (distribution,
foraging).
Tetragonula biroi (Friese, 1898)
Trigona biri Friese 1898: 428, 429: Syntypes (ZMHB, 2 workers): examined, "N. Guinea / Bir 96",
"Friedrich- / Wilh.-hafen", "Trigona / biroi / det. Friese 1897" (1 worker); "Philippinen / Schadenberg",
"Trigona / biroi / det. Friese 1897 / n.sp.", "Coll. / Friese" (1 worker) (key to species, taxonomy); Type
locality: PHILIPPINES "Philippinen (Schadenburg)" (few workers); "N-Guinea (Berlinhafen und

TERMS OF USE
Friedrich Wilhelmshafen, 1896 Bir" (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 (former
Paranas); Floral record: Citrus; Melicope triphylla; Mimosa pudica; Hedicke 1926: 423 (distribution);
PHILIPPINES, Luzon; Mindanao; Siargao; Williams 1928: 91-92 (nest); Cockerell 1929d: 301 (compar-
ative note (as birsi!)); Cockerell 1930a: 157 (distribution, floral record, uncertain identity); CAMBODIA,
Tung Nui (misidentification); Satul (misidentification); Floral record: Melastoma malabaricum; Schwarz
1939a: 85, 86, 90, 101, 119-120* (distribution, key to species, taxonomy); Floral record: Citrus; Melicope
triphylla; Mimosa pudica; Nipa fruticans; Moure 1961: 209 (systematic position); Michener 1961: 28
(comparative note); Michener 1965: 231 (citation); Baltazar 1966: 406-407 (distribution); 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 (comparative
note, distribution); PHILIPPINES; New Guinea; Oshy & Arikumar 2000: 1651 (pollination); Floral
record: Bambusa vulgaris; Koshy et al. 2001: (see iridipennis); Cortopassi-Laurino et al. 2006: 303
(citation (as biro!)); Slaa et al. 2006: 303 (citation (as biro!)); Temaru et al. 2007: 282, 284 (honey 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 angophor Cockerell 1912: 225: Holotype (BMNH 17b.1139) (comparative notes, floral record,
taxonomy); Type locality: AUSTRALIA "Sydney, New South Wales, Dec. 1, 1910 (Froggatt, 118)"
(worker); Floral record: Angophora; Cockerell 1929d: 301 (comparative note, new status (as subspecies
of carbonaria)); Cockerell 1930b: 150 (distribution, floral record, taxonomy); AUSTRALIA, Sydney;
Floral record: Angophora; Rayment 1932f: 106 (taxonomy); Rayment 1935: 735, 740 (key to species,
taxonomy); Rayment 1939: (distribution); Schwarz 1948: 91; Rayment 1953: (distribution); Moure
1961: 209 (morphology, systematic position); Michener 1965: 231 (citation); Camargo 1988: 372 (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

TERMS OF USE
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); Lea
1910: 119, 207 (inquiline (Brachypeplus auritus; Brachypeplus basalis: Nitidulidae: Coleoptera)); Strand
1911: 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: Helianthus
annuus; Friese 1917: 9 (distribution); Cockerell 1922a: 279 (comparative note); Salt 1929: 452, 453
(citation); Cockerell 1929e: 243 (distribution); AUSTRALIA, Ourimbah, New South Wales; Cockerell
1929d: 301 (common name (karbi), distribution, nest); AUSTRALIA, Moreton; Cockerell 1929a: 8
(comparative note (identification corrected in footnote)); Cockerell 1930b: 149 (common name (karbi),
floral record, taxonomy); Floral record: Helianthus annuus; Cockerell 1930c: 149-150 (distribution, 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 (or
Epuraea): Nitidulidae: Coleoptera; Tribolium myrmecophilum: Tenebrionidae: Coleoptera)); Rayment
1932f: 106 (taxonomy); Cockerell 1934: 62, fig. 8 (illustration (mandible)); Rayment 1935: 735, 736,
737, 738, 739 (comparative note, key to species, taxonomy); Cannon 1937; Schwarz 1939a: 103 (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 Lamington
National Park); Brisbane; Bundaberg; Caboolture; Caloundra; Capalaba; Mount Edwards; Gladstone;
Ipswich; Jimboomba; Mackay; Maroochydore; Moggill (near Brisbane); Noosa; Petrie; Rockhampton;
Sarina; Tamborine; Tibrogargen (near Beerwah); Yaamba; Yarriman; Wille 1961: 117; Rayment 1961:
(nest); Wakefield 1962: (swarm (probably not stingless bees)); Michener 1964: 236 (nest); Michener
1965: 231 (citation); Pooley & Michener 1969: 429 (comparative note); Nogueira-Neto 1970: 218, 266,
314; Wille & Michener 1973: 13, 24, 47, 58, 70, 80, 81, 89, 92, 178 (nest); Michener 1974: 62, 341, 343
(colony, nest); McKenzie 1975: (common name (kabbai), honey); Kistner 1982: 122 (citation, inquiline
(Brachypeplus auritus: Nitidulidae: Coleoptera)); Wagner & Dollin 1982a: (distribution); Evans & 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); Sakagami
et al. 1983a: 106 (comparative note); Sakagami et al. 1983d: 454 (comparative note); Wagner & Briscoe
1983: (genetics); Hawkeswood 1983: (pollination); Floral record: Cupaniopsis anacardioides; Graur
1985: 194 (comparative note); Yong 1986: 628 (comparative note); Goebel 1986: (bee-keeping); Starr &
Sakagami 1987: 97, 106 (citation); Yong et al. 1987: 467 (biochemistry); Fain & Heard 1987: 197, 199,
202 (inquiline (Cerophagus trigona: Acaridae: Acari)); Milborrow et al. 1987: 15, 22, 23, 25 (wax);
Heard 1987: (nest); Camargo 1988: 372 (distribution, taxonomy); AUSTRALIA, Cape York, Gosford;
Maitland, New South Wales; Heard 1988: (bee-keeping); Adams & Lawson 1988: (pollination);
Michener 1990: 95, 101, 119, 120, 127, 128 (illustration, morphology, systematic position); Sakagami &
Inoue 1990: 211, 212, 215 (comparative note); Wittmann et al. 1991: 802, 806 (citation); Harriss &
Beatie 1991: (pollen transportation); Ornduff 1991: 207 (pollination); Floral record: Cycas media;
Adams et al. 1992: (orchid pollination); Michener & Roubik 1993: 259, 260 (comparative note, 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;

TERMS OF USE
Melastoma; Nymphaea; Oxylobium; Persoonia; Podolepis; Pultenaea; Rubus; Telopea; Xanthorrhoea;

Williams & Adam 1993: 480 (floral record); Gross 1993: 471, 472, 473 (pollination); Floral record:
Melastoma affine; Adams & Lawson 1993: (pollination); Floral record: Dendrobium teretifolium; Heard
1994: (pollination); Floral record: Macadamia integrifolia; Heard & Exley 1994: (pollination); Floral
record: Macadamia integrifolia; Forster et al. 1994: 219 (pollination); Floral record: Cycas; Bartareau
1994: (pollination); Floral record: Dendrobium toressae; Yamane et al. 1995: (oviposition); Wallace &
Trueman 1995: (seed dispersal (mellitochory)); Floral record: Eucalyptus torelliana; Disney & 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-Neto
1997: 85, 204, 205, 334; Dollin et al. 1997: (nest, revision, taxonomy); AUSTRALIA, New South Wales;
Queensland; Williams & Adam 1997: 75, 86, 88, 90-92 (competition, floral record); Floral record: 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; Hilrio 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); Hilrio et al. 2001: 195
(citation); Serro 2001: (morphology (proventricular structure)); White et al. 2001: (flower constancy);
Dollin et al. 2001: (bee-keeping); Green & Oldroyd 2002: (maternity of male); Wallace et al. 2002:
(pollination); Floral record: Persoonia virgata; Pick & Blochtein 2002: 827, 831, 833, 834 (citation);
Dollin 2002a: (bee-keeping); Dollin 2002b: (bee-keeping); Schmidt et al. 2003: 245 (citation); Gardener
et al. 2003: 120 (citation); Williams & Walker 2003: 143, 144, 145 (floral record); Floral record: Pollia
crispata; Heard 2003: (citation); Nieh 2004: 162, 164, 168 (citation, communication, comparative table);
Franck et al. 2004: (genetics, microsatellite, nest); Cameron et al. 2004: 2358 (genetics); Tth 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); Arias
et al. 2006: 267 (citation); Cortopassi-Laurino et al. 2006: (bee-keeping); Slaa et al. 2006: 297, 301,
306, 308 (citation); Bacelar-Lima et al. 2006: 344 (citation); Hoese et al. 2006: 769, 774 (predator (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); Klumpp
2007; Wallace et al. 2008: (dispersal, pollination); Floral record: Corymbia torelliana; Irish et al. 2008:
(honey properties).
Tetragonula clypearis (Friese, 1909)
Trigona laeviceps variety clypearis Friese 1909("1908"): 356, 358, fig. 15-3: Unknown; paralectotype
(ZMHB (1)) (distribution, taxonomy); Type locality: INDONESIA (IRIAN JAYA) "Manikion, 14.-28.
Februar 1903"; "Wendesi 29. Juni 1903, New Guinea" (worker); Friese 1909: 275; Moure 1961: 209
(systematic position); Michener 1961: 34 (synonymy); Michener 1965: 231 (citation); Wille &

TERMS OF USE
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); Tth 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 larger
form of laeviceps from the Philippines (taxonomy); Type locality: PHILIPPINES "Dapitan, Mindanao
(Baker, 23119)" (worker); Schwarz 1939a: 86, 112, 113 (synonymy); Moure 1961: 208 (morphology,
systematic position).
Tetragonula davenporti (Franck, 2004)
Trigona (Heterotrigona) davenporti Franck et al. 2004: 2325, 2326, 2327, 2328, 2330-2331, figs. 5a, 6a:
Holotype (QM, worker); paratypes (ANIC (2), BMNH (2), MNHN (2), QM (2), "INRA collection, 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 from
Mr. 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 & Ross
2000: 6, 13-14 (synonymy).

TERMS OF USE
Tetragonula drescheri (Schwarz, 1939)
Trigona (Tetragona) sarawakensis variety drescheri Schwarz 1939a: 85, 93, 106-107*: Holotype (AMNH);
paratype (AMNH (1) (distribution, key to species, taxonomy); Type locality: INDONESIA "M. JAVA.-
South Banjoemas, Koebangkangkoeng, 25 meters, July 1935 (F. C. Drescher)" (1 worker, holotype); "E.
JAVA.-Mt. Goemitir, Nov. 3 (R. van der Veen)" (1 worker, paratype); Schwarz 1948: 118; Sakagami
1959: 120 (citation); Moure 1961: 210 (morphology, systematic position); Yoshikawa et al. 1969: 174
(distribution); MALAYSIA, Tapah; Sakagami 1978: 210 (synonymy); Sakagami et al. 1983b: 10-12, 20
(nest); Sakagami & Inoue 1985: 174, 176, 177, 178, 179, 180, 181, 182-184*, 186, 187-188 (key to 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 clinging
to the leg of a large green cicada (Platylomia spiculata (=Dundubia spiculata): Cicadidae: Hemiptera)"));
CAMBODIA, Pahtoop Mt.; Cockerell 1930a: 157 (distribution, ecology (attracted to sweat)); CAMBO-
DIA, Chun Het; Trang; Yasumatsu 1935: 94 (common name (mitsubachi), distribution); PALAU
ISLANDS; Esaki 1936: 44, 46-37 (common name (galadngikl; chaladngikl; mitsubachi), nest); PALAU
ISLANDS; Schwarz 1937: 290, 310, 311*, 316, 328 (taxonomy); Schwarz 1939a: 107, 108 (synonymy);
Schwarz 1939b: 152 (synonymy); Cockerell 1939a: 61, 62, 64 (distribution ("presumably introduced by
man")); PALAU ISLANDS, Melekeiok on Babelthuap; Ngeremlengui, Ogiwal; Schwarz 1948: 69, 117;
Krombein 1950: 133 (synonymy); Moure 1961: 209 (morphology, systematic position); Sakagami
1978: 223, 224 (synonymy); Camargo 1988: 371 (synonymy, taxonomy); Sakagami et al. 1990: 128
(synonymy).
Trigona brevis Friese 1925: 41: Nomen nudum. The name is unavailable for taxonomic purposes, as no
description or definition of the taxon was provided (ICZN, art. 12.1) (comparative note).
Trigona erythrostoma Cameron 1908: 193-194: Syntypes (BMNH 17b.1113, 2 workers) (key to species, tax-
onomy); Type locality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker or queen); Cockerell
1918: 386 (comparative notes); Cockerell 1920a: 116 (comparative note); Schwarz 1937: 290, 310*, 326
(key to species, taxonomy); Schwarz 1939a: 112 (synonymy); Moure 1961: 195, 209 (synonymy).
Trigona fusco-balteata Cameron 1908: 193, 194: Lectotype (BMNH 17b.1112): Designated Moure 1961
(key to species, taxonomy); Type locality: MALAYSIA "Medang, Sarawak (Hewitt)" (worker); Schwarz
1937: 290, 310-311*, 316, 326 (key to species, taxonomy); Schwarz 1939a: 85, 86, 88, 90, 93, 96, 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; Schwarz
1948: 40, 56, 69, 83, 90, 118; Krombein 1950: 102, 133, 135 (distribution, morphology); MICRONESIA,
Caroline Islands; Nogueira-Neto 1951: (comparative note); Pagden 1957: 15, 17 (flight); Sakagami

TERMS OF USE
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); Sakagami
1982: 389 (citation); Appanah 1982: (pollination); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve;
Floral record: Xerospermum intermedium; Wille 1983: 50 (nest); Sakagami et al. 1983b: 2, 9-10, 20
(nest); Sakagami et al. 1983a: 106 (comparative note); Sakagami & Inoue 1985: 177 (comparative
note); Sakagami et al. 1985: (distribution, key to species); Kiew 1985: (luring (unwashed shorts)); Yong
1986: (allozyme); Sakagami & Khoo 1987: 213 (comparative note); Sakagami & Inoue 1987: 626
(comparative note); Starr & Sakagami 1987: (common name (kiyot), nest); Yong et al. 1987: (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 & Saiboon
2000: (distribution, nest); THAILAND, Songkhla; Amano et al. 2000: (citation); Michener 2001: 232
(morphology); Green et al. 2001: 91 (microsatellite); Eltz et al. 2003: 309 (comparative note); Franck et
al. 2004: 2318 (citation); Cameron et al. 2004: 2358 (nest); Eltz 2004: 322 (abundance); Samejima et al.
2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Amano 2004: 3 (introduction, 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);
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 / (erroneous
locality) / det. J.S.Ascher"; paralectotypes (DEI (3)) (taxonomy); Type locality: UNCERTAIN "Campi-
nas (So 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

TERMS OF USE
1979a: (morphology (pollen press)); Michener & Roubik 1993: 260 (comparative note, morphology);
Camargo & Pedro 2003a: (synonymy); Camargo & Pedro 2007: 524 (synonymy).
Trigona pygmaea Friese 1933b: 147: Lectotype (ZMHB, worker): here designated "O-Sumatra / Mandau /
7.1933", "Trigona / pygmaea / Fr. / 1933 Friese det. / n."; paralectotype (DEI (1), ZMHB (1)) (common
name (kloeloet itam ketjid)); Type locality: INDONESIA "Beringin, in der Wldern 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); Schwarz
1937: 282, 290, 311-313* (key to species, taxonomy); Schwarz 1939a: 101, 117 (synonymy); Moure
1961: 209 (synonymy); Sakagami 1978: 195, 196 (synonymy).
Trigona geissleri Cockerell 1918: 385-386, 387: Unknown: Cockerell used a Friese manuscript name for his
description (key to species, distribution, manuscript name of Friese, taxonomy, variation); Type locality:
MALAYSIA "Sintang, North Borneo" (male); "Singapore by Baker" (couple of workers); Cockerell
1919b: 242 (taxonomy, variation); MALAYSIA, Sandakan; Cockerell 1920b: 228 (distribution);
MALAYSIA, Sandakan; Schwarz 1937: 311, 312, 313, 327; Schwarz 1939a: 85, 90, 95, 109, 114, 115,
117-119*, 120, 136 (distribution, key to species, taxonomy (probably several distinct species were
included here)); Schwarz 1948: 83, 118, 233; Moure 1961: 209 (morphology, systematic position); 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, Pasoh
Forest 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 & Khoo
1987: 208, 209, 210, 211, 212 (morphology); Sakagami & Inoue 1987: 625, 626 (morphology); Starr &
Sakagami 1987: 97 (citation); Camargo 1988: 371 (distribution); MALAYSIA, Borneo, Sandakan; 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 (as
geisslerrii!)); Liow 1999: 106, 127 (distribution (as geisslerei! in part)); Nagamitsu et al. 1999: (floral
records (not by species), foraging); Liow et al. 2001: 184; San & Osawa 2002: (population); 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"

TERMS OF USE
(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 (comparative
note); 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 (1
worker); "Mae Klang" (2 workers); "Mae Fack" (1 worker); "Lamphun" (1 worker); "Uthaithani" (1
worker); Sakagami & Inoue 1985: 177, 182 (comparative note); Sakagami et al. 1985: (distribution);
Sakagami & Inoue 1987: 625, 626 (morphology); Sakagami et al. 1990: 131 (distribution); Dollin et al.
1997: 873 (comparative note); Tadauchi et al. 1998: 243 (type depositories); Nagamitsu & Inoue 2005:
77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (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 (comparative
note, morphology, systematic position); Michener 1961: 10, 18, 23-27, 29, 40, 42, 44 (aggressiveness,
nest); Wille 1961: 117, 125, 129; Michener 1965: 231 (citation); Kerr et al. 1967: 283 (citation);
Darchen 1969: 462 (illustration, physiology); Wille & Michener 1973: 14, 24, 38, 48, 59, 71, 94, 100
(nest); Michener 1974: 334, 343, 344 (nest); Wagner & Dollin 1982b: (swarming); Wagner & Dollin
1982c; Sakagami et al. 1983b: 20 (comparative note); Sakagami et al. 1983a: 106 (comparative note);
Sakagami et al. 1983d: 454 (comparative note); Dollin & Dollin 1983: (nest); Starr & Sakagami 1987:
97 (citation); Michener 1990: 120 (systematic position); Sakagami & Inoue 1990: 211, 212, 215 (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); Dollin
2002a: (bee-keeping); Dollin 2002b: (bee-keeping); Gardener et al. 2003: (foraging, nectar choice);
Franck et al. 2004: (genetics, microsatellite, nest); Tth 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

TERMS OF USE
& Cameron 2007: (systematics); Michener 2007a: 824-825; Klumpp 2007: 4, 5, 6, 8, 11, 15, 46, 77, 85;
Wallace et al. 2008: (dispersal, pollination); Floral record: Corymbia torelliana.
Tetragonula iridipennis (Smith, 1854)
Trigona iridipennis Smith 1854: 413-414: Holotype (BMNH 17b.1114, worker): examined, "Type" (orange
border), "iridipennis / Type Sm.", "B.M.TYPE / HYM. / 17B.1114", "Trigona / iridipennis / TYPE.
Smith.", "Ceylon" (reverse side "53 / 23"). Citations for iridipennis are included in a broad sense, as 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); Type
locality: SRI LANKA "Ceylon" (worker); Tennent 1859: 282 (list); Tennent 1861: 454 (list); Castets
1893: 468, 481-488 (nest); Dalla Torre 1896: 579 (citation); Bingham 1897: 560, 563-564 (distribution,
key to species, taxonomy); MYANMAR, Tenasserim; SRI LANKA; Friese 1904: 140 (see laeviceps);
Friese 1909: 273 (comparative note); Strand 1913a: 149-150 (distribution (as laeviceps)); SRI LANKA;
Friese 1914a: 12-13 (see laeviceps); Friese 1914a: 12 (synonymy (as canifrons)); Friese 1914a: 12 (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 (see
laeviceps); George 1933: (physiology (as irridipennis!)); George 1934: (brood, distribution, morphology,
nest (crevice of wall), physiology (as irridipennis!)); INDIA, Coimbatore; Schwarz 1937: 283, 290, 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 the
specimens 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-Arajo & Kerr 1959: 226 (comparative
note); Sakagami 1960: 146 (see laeviceps); Lindauer & Kerr 1960: 31, 32, 34 (communication); SRI
LANKA; Lindauer & Kerr 1960: 31, 32, 34; Kerr 1960: 386 (citation); Moure 1961: 206, 207, 208
(morphology, systematic position); Michener 1961: 3, 10, 28-33, 35, 38, 39, 42 (see laeviceps); 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: (comparative
note); Schmid 1964: 588 (citation); Michener 1965: 231 (citation); Kerr & Esch 1965: (comparative
note); Baltazar 1966: 407 (see laeviceps); Phadke 1968: (honey, uncertain identity (unidentified));
INDIA, Mysore, Madhya Pradesh; Kerr 1969: 166 (systematics); Darchen 1969: 482 (flight);
Yoshikawa et al. 1969: 168, 169, 173 (see laeviceps); Jander & Jander 1970: 361 (citation); Sakagami
1971: 345 (citation); Wille & Michener 1973: 14, 24, 30, 48, 59, 71, 94, 96, 98, 100, 102-103, 257 (nest);
Michener 1974: 57, 153, 154, 343 (colony, nest); Sakagami 1975: 59 (comparative note); Kerr &
Cunha 1976: (systematics); Zeuner & Manning 1976: 222-224, plate 2 (morphology); Wille 1977: (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,

TERMS OF USE
216-219*, 220, 221, 222, 225, 226, 227, 228, 230, 234, 235, 236, 237, 238, 239, 247, plate 5 (taxonomy);
INDIA; SRI LANKA; Wille 1979b: 243 (comparative note); Koeniger & Vorvohl 1979: (foraging); SRI
LANKA; Floral record: Boehmeria; Cassia; Cleome; Cocos; Erigeron; Homonoia; Lannea; Mimosa;
Ocimum; Phyllanthus; Sapium; Smithia; Solanum; Syzygium; Taxillus; Toona; Tridax; Vitex; Divan
1981("1980"): (behavior); Sakagami & Ito 1981: 368, 373 (morphology); Wille 1983: 53 (population);
Sakagami et al. 1983b: 6 (comparative note); Sakagami et al. 1983a: 101, 103, 106 (comparative note);
Sakagami & Inoue 1985: 177, 182 (comparative note); Sakagami et al. 1985: (distribution); Utami
1986: (see laeviceps); Sakagami & Khoo 1987: 208, 209 (comparative note); Sakagami & Inoue 1987:
625, 626 (morphology); Uji 1987: (see laeviceps); Bichee & Sharma 1988: (pollination); Floral record:
Helianthus annuus; Mohana Rao 1988("1987"): (foraging); Floral record: Trigonella corniculata;
Camargo & Wittmann 1989: 223 (citation); Kumar & Kumar 1989: (foraging (as irridipensis!)); 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; Floral
record: Areca vestiaria; Percy 1989: (bee-keeping); Michener 1990: 89, 119, 120, 126, 127 (systematic
position); Sakagami et al. 1990: (misidentification, synonymy); Sakagami & Inoue 1990: 212 (taxon-
omy); Goel & Kumar 1990: 44 (pollination); INDIA, W Uttar Pradesh; Floral record: Helianthus
annuus; 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; Delonix
regia; Eucalyptus globulus; Lawsonia alba; Leucaena leucocephala; Loranthus longiflorus; Peltophorum
pterocarpum; Prosopis juliflora; Rotala densiflora; Tamarindus indica; Michener & Roubik 1993: 260
(comparative note, morphology); Cunha 1994: (systematics); Roubik 1995: 24, 41, 42 (pollination); 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 & Suryanarayana
1999("1997"): (foraging (as inridipennis!)); Floral record: Ceiba pentandra; Joshi et al. 1998: (honey,
pollen); INDIA, Pune, Maharashtra; Floral record: Ageratum conyzoides; Amaranthus; Bombax ceiba;
Cocos nucifera; Moringa oleifera; Peltophorum pterocarpum; Sorghum vulgare; Zea mays; Heard
1999a: 193, 194, 197, 198 (citation); Crane 1999: 301; Michener 2000: 800-802; Jarau et al. 2000: 86
(citation); Wu 2000: 387, 392-393 (key to species, morphology); Atluri et al. 2000: (pollination, 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 et
al. 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); Floral
record: Wrightia tinctoria; Raju 2005: 106 (floral record); Floral record: Woodfordia floribunda; Raju et

TERMS OF USE
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 lviceps 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 be
resolved in a separate paper with C.D. Michener. Citations below for laeviceps are in a broad sense, as
species limits are uncertain and may include both valdezi and testaceitarsis. These taxa have previously
been proposed as junior synonymons of laeviceps, but are slightly larger (taxonomy); Type locality: SIN-
GAPORE "Singapore" (worker); Smith 1858: (propolis, wax); Smith 1859: 135 (distribution); INDO-
NESIA, Aru; INDIA (misidentification); SINGAPORE; Smith 1865: 93 (distribution); "NEW
GUINEA"; 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 (comparative
notes, synonymy, uncertain identity (as canifrons)); AUSTRALIA, Adelaide River (misidentification);
SRI LANKA (misidentification, *iridipennis group); Bingham 1905: 59 (citation, distribution, nest
(house-post in village)); Brown 1906: 686 (distribution); PHILIPPINES; Schulz 1907: (nest, predator
(Oecophylla smaragdina: Formicidae: Hymenoptera)); Friese 1909("1908"): 354, 355 (distribution,
uncertain identity (as praeterita)); INDONESIA, Irian Jaya, Etna Bai; Merauke; Friese 1909("1908"):
354, 356 (distribution); INDONESIA, Irian Jaya, Manikion; Wendesi; Friese 1909: 272, 275 (key to spe-
cies, nest (in structural wood)); INDONESIA, Ost-Java; Sumatra; Schulz 1909: (nest (as canifrons in
part)); Friese 1912: 94 (distribution); PAPUA NEW GUINEA, Sepik-Bivak; Ihering 1912: 3 (citation (as
laevicops!)); Strand 1913a: 149-150 (see iridipennis); Friese 1914a: 12-13 (distribution, floral record,
uncertain identity (as iridipennis)); INDONESIA, Bandung; Batavia; Buitenzorg; Nongkodjadjar; Nusa
Kambangan; Salatiga; Samarang; Floral record: Canna flavescens; Friese 1915: 4 (distribution, uncertain
identity (as praeterita)); INDONESIA, Irian Jaya/PAPUA NEW GUINEA, Etna Bai; Jasa; Merauke;
Njao; Zoutbron; Cockerell 1918: 385 (comparative notes); Cockerell 1919c: 77, 79 (distribution, 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); Cockerell
1925b: (misidentification (was a Diptera)); Cockerell 1926b; Salt 1929: 460 (citation); Cockerell 1929c:
590, 592 (comparative note); Cockerell 1929e: 242, 243 (comparative note); Cockerell 1929d: 300 (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 specimens
examined by Schwarz and identified as iridipennis are actually laeviceps, see iridipennis)); Floral record:
Typha angustifolia; Schwarz 1945: 141 (citation); Schwarz 1948: 23, 84, 141; Nogueira-Neto 1949: 121
(synonymy); Sakagami 1959: 120 (comparative note); Sakagami 1960: 146 (comparative note, distribu-

TERMS OF USE
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: (see
minangkabau); Sakagami & Yamane 1984: 37, 48; Inoue et al. 1984a: (see minangkabau); Salmah et al.
1984: (see minangkabau); Sakagami & Inoue 1985: 175, 176, 177, 180, 181, 182, 184, 186 (comparative
note); Sakagami et al. 1985: (distribution, key to species); Utami 1986: (pollination, uncertain identity
(as irridipenis!)); INDONESIA, West Java, Pandeglang; Floral record: Metroxylon sagu; Sakagami &
Khoo 1987: 208, 209, 210, 211, 212 (morphology); Sakagami & Inoue 1987: 611, 624 (comparative
note); Starr & Sakagami 1987: 97 (citation); Salmah et al. 1987: 384 (synonymy); Uji 1987: (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-Fonseca
1990: (see minangkabau); Salmah et al. 1991: 196 (synonymy); Prasad & Hemanth 1992: (predator
(Ardeola grayii: Ardeidae: Ciconiiformes) (as leviseps!)); Michener & Roubik 1993: 260 (comparative
note, morphology); Inoue et al. 1993: (nest); INDONESIA, Sumatra; Cardale 1993: 323-324 (see 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-Neto
1997: 94 (citation); Dollin et al. 1997: (revision, synonymy, taxonomy); Nagamitsu & Inoue 1997: 435,
436 (pollination); Floral record: Santiria laevigata; Dollin 1997: (compared to sapiens); Nagamitsu &
Inoue 1998: (foraging, morphology); Liow 1999: 106, 116, 127 (nest); Nagamitsu et al. 1999: (floral
records (not by species), foraging); Peters et al. 1999: 382 (citation); Crane 1999: 89; van Veen & 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);

TERMS OF USE
MALAYSIA, Danum Valley; Sepilok; Deramakot; Nagamitsu & Inoue 2002: 309 (citation); San &
Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Eltz et al. 2003: (nest,
tree preference); Camargo & Pedro 2003a: (comparative notes, systematics, taxonomy); Hohmann
2003: (communication, nest); Biesmeijer & Slaa 2004: 148 (citation, foraging); Franck et al. 2004:
(genetics, microsatellite); Eltz 2004: 322 (abundance); Samejima et al. 2004: (population); MALAYSIA,
Sarawak, Miri, Upper Baram area; Wattanachaiyingcharoen et al. 2004: (diversity, nest); Nagamitsu &
Inoue 2005: 77, 78, 82, 83, 84, 85 (distribution, foraging); Chinh et al. 2005: 363, 364, 367-368, 370
(nest); VIETNAM, Cucphuong National Forest, Red river delta; Klakasikorn et al. 2005: 2, 3, 5, 6 (floral
record); Floral record: Hevea brasiliensis; Roubik 2006: 134 (nest); Tangmitcharoen et al. 2006: (polli-
nation); Floral record: Tectona grandis; Inson & Malaipan 2006: (distribution, foraging, nest, uncertain
identity (as iridipennis)); THAILAND, Kanchanaburi, Thong Pha Phum; Rasmussen & Cameron 2007:
(systematics); Leonhardt et al. 2007: (foraging); Floral record: Ardisia pyramidalis; Diospyros 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 taxon
is labeled "Bandjarmas" (?=Bandjarmasin, near Liangtelan) (F. Penati, pers. com.) (taxonomy); Type
locality: MALAYSIA "Liangtelan (Borneo)" (1 worker); Dalla Torre 1896: 580 (citation); Cockerell
1919b: 242 (distribution); MALAYSIA, Sandakan; Cockerell 1920b: 228 (distribution); MALAYSIA,
Sandakan; Schwarz 1937: 282, 283, 290, 315*, 316, 326 (key to species, taxonomy); Schwarz 1939a: 85,
93, 120* (distribution, key to species, taxonomy); Schwarz 1948: 118; Moure 1961: 210 (systematic
position); Yoshikawa et al. 1969: 172, 174 (distribution); MALAYSIA, Sabah, Touaran; Sakagami &
Inoue 1985: 181 (comparative note); Sakagami et al. 1985: (distribution); Sakagami et al. 1990: 131
(distribution); Inoue et al. 1995: 97, 98 (pollination); Floral record: Dendrobium setifolium; Neuwiedia
borneensis; Roubik 1996: 61, 62 (distribution, predator (Helarctos malayanus: Ursidae: Carnivora));
BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: 433, 435, 436, 437 (pollination); Floral record:
Santiria laevigata; Nagamitsu & Inoue 1998: (foraging, morphology); Nagamitsu et al. 1999: (floral
records (not by species), foraging); Eltz et al. 2001a: (pollen foraging); Eltz et al. 2001b: 343, 345 (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; Floral
record: Rhizophora apiculata; Nagamitsu & Inoue 2002: 305, 307, 308, 310, 312 (foraging); Eltz et al.
2003: (nest, tree preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz 2004: 322 (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); Type
locality: MALAYSIA "Kuching, Borneo (John Hewitt)" (worker); Schwarz 1937: 315, 326; Schwarz
1939a: 120 (synonymy); Moure 1961: 210 (synonymy); Michener & Boongird 2004: 146 (comparative
note).
Tetragonula melina (Gribodo, 1893)
Trigona melina Gribodo 1893: 262-263, 264: Syntypes (MSNG, 3 workers; unknown depository, 2 workers):
Two syntypes are labeled "Bandjarmas" (?=Bandjarmasin, near Liangtelan), while the third syntype carry
no locality label (F. Penati, pers. com.) (taxonomy); Type locality: MALAYSIA "Liangtelan (Borneo)" (2

TERMS OF USE
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); Kiew
1985: (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 & Inoue
1997: (foraging, pollination); Floral record: Santiria laevigata; Nagamitsu & Inoue 1998: (foraging,
morphology); Liow 1999: 106, 110, 127 (nest); Nagamitsu et al. 1999: (floral records (not by species),
foraging); Liow et al. 2001: 184; Eltz et al. 2001a: (pollen foraging); Eltz et al. 2001b: 343, 345 (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: Rhizophora
apiculata; Nagamitsu & Inoue 2002: 305, 307, 308, 310, 312 (foraging); Eltz et al. 2003: (nest, tree
preference); Biesmeijer & Slaa 2004: 148 (citation, foraging); Eltz 2004: 322 (abundance); Samejima et
al. 2004: (population); MALAYSIA, Sarawak, Miri, Upper Baram area; Nagamitsu & Inoue 2005: 77,
80, 82, 83, 84, 85, 86, 88 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation); Rasmussen &
Cameron 2007: (systematics).
Tetragonula mellipes (Friese, 1898)
Trigona mellipes Friese 1898: 428, 429: Syntypes (ZMHB, 1 worker, 1 male): examined, "C. Austr. / v.
Mller.93", "Trigona / mellipes / det. Friese 1897", "Zool. Mus. / Berlin", "Coll. / Friese" (only worker).
Friese stated the type locality was "S. Australia", probably an error for C. Australia, as ZMHB specimens
match the original description and the taxon is not known from southern Australia (key to species, 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; Western
Australia; Dollin 1997: (characteristics); Crane 1999: 89; Heard & Dollin 2000: (bee-keeping); Green et
al. 2001: 91 (microsatellite); Serro 2001: (morphology (proventricular structure)); Palmer et al. 2002:
(maternity); Franck et al. 2004: (genetics, microsatellite, nest); Tth 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:

TERMS OF USE
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, ix
1981 in mass flight, S.F. Skagami" (1 worker, holotype); "Lubuk Mintrun" (many workers, 25 males,
paratypes); Sakagami et al. 1983a: (nest, taxonomy (misidentification; as laeviceps)); INDONESIA,
Sumatera Barat at Lubuk; Sakagami et al. 1983d: (oviposition behavior (misidentification; as laeviceps));
Inoue et al. 1984b: (brood, nest, reproduction (as laeviceps)); Inoue et al. 1984a: (physiology, 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); Salmah
et al. 1987: 383, 388, 389, 390 (development); Roubik 1989: 86, 133, 137; Sakagami et al. 1989: 271,
272 (comparative note); Bego et al. 1989: 48, 53 (foraging, greenhouse, introduction); Sakagami 1989:
164; Sakagami et al. 1990: (distribution, key to species); INDONESIA, Sumatra; Sakagami & Inoue
1990: 201, 202, 203, 205, 210, 211, 212, 215 (comparative note); Salmah et al. 1990: (distribution, nest
(hollow rambutan branches (Nephelium lappaceum: Sapindaceae); bamboo; interspaces)); INDONESIA,
Sumatera Barat; Inoue et al. 1990b: (population); Inoue 1990: 582 (nest, population); Suka et al. 1990:
(nestmate recognition); Roubik 1990a: 115 (citation); Engels & Imperatriz-Fonseca 1990: 186, 207
(citation (as laeviceps)); Inoue et al. 1990a: 229; Salmah et al. 1991: (reproduction); Kakutani et al.
1993: (pollination); Floral record: Fragaria; Suka & Inoue 1993: (behavior); Inoue et al. 1993: (nest
(hollow rambutan branches (Nephelium lappaceum: Sapindaceae))); INDONESIA, Sumatra; Hoshiba &
Imai 1993: 469, 477, 481, 487 (chromosome); Yamane et al. 1995: 276, 282, 293 (comparative note);
Roubik 1995: 151 (antropogenic nest); Inoue et al. 1995: 98 (pollination); Floral record: Neuwiedia
borneensis; Inoue et al. 1996b: (development, nest, variation); Salmah et al. 1996: 402, 407, 408, 409,
410 (comparative note); Inoue et al. 1996a: (population); Tadauchi et al. 1998: 243-244 (type 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); Tth et al. 2003: 366 (citation); Costa et al. 2004: 187
(citation); Tth 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); Sakagami
et al. 1990: 131 (distribution); Tadauchi et al. 1998: 244 (type depositories); Nagamitsu & Inoue 2005:
77 (distribution, foraging); Klakasikorn et al. 2005: 2-3, 4, 5, 6 (floral record, morphology); Floral
record: Homalium grandiflorum; Tectona grandis; Rasmussen & Cameron 2007: (systematics).

TERMS OF USE
Tetragonula pagdeni (Schwarz, 1939)
Trigona (Tetragona) fusco-balteata variety pagdeni Schwarz 1939a: 85, 93, 96, 110-111*, 113: Holotype
(USNM 53564, male): examined, "Nakon / SriTamarat / Siam 7/5/28", "HughSmith / coll", Type No. /
53564 / U.S.N.M.", "Holotype", "Trigona / fusco-balteata / var. pagdeni / H.F. Schwarz"; paratype
(AMNH (1)) (distribution, key to species, taxonomy); Type locality: THAILAND "SIAM.-Nakon Sri
Tamarat, July 5, 1928 (Hugh M. Smith)" (workers, males); "Singora, June 1929 (Hugh M. Smith)" (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 (comparative
note, nest, taxonomy); CAMBODIA, Udong; THAILAND, Cheng Mai; Doi Dao; Kamphangpet; Korat;
Yoshikawa et al. 1969: 168, 169 (nest); Wille & Michener 1973: 14, 24, 59 (nest); Sakagami 1978: 166-
194, 200, 211, 212, 213, 218, 219-223*, 224, 225, 226, 227, 228, 234, 236, 237, 238, 239, 240, 247, plate
5 (taxonomy); Sakagami et al. 1983b: 2, 6-7, 8, 20 (nest); Sakagami et al. 1983a: 103, 105, 106 (compar-
ative note); Sakagami et al. 1983d: (oviposition behavior); Sakagami & Inoue 1985: 177 (comparative
note); Sakagami et al. 1985: (distribution); Kiew 1985: (luring (unwashed shorts)); Yong 1986: 627
(comparative note); Sakagami & Khoo 1987: 213-214 (morphology); Sakagami & Inoue 1987: 626
(comparative note); Starr & Sakagami 1987: 97 (citation); Sakagami et al. 1990: 130, 131 (comparative
note, distribution); Sakagami & Inoue 1990: 201, 202, 203, 204, 205, 206, 210, 211, 212, 215 (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" (1
worker); "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 (comparative
note); Sakagami et al. 1990: 130, 131 (comparative note, distribution); Michener & Roubik 1993: 260
(comparative note, morphology); Dollin et al. 1997: 873 (comparative note); Tadauchi et al. 1998: 244
(type depositories); San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest
Reserve; Nagamitsu & Inoue 2005: 77 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation);
Roubik 2006: 132 (nest).
Tetragonula penangensis (Cockerell, 1919)
Trigona penangensis Cockerell 1919c: 78, 79: Holotype (AMNH) (comparative notes, key to species); Type
locality: MALAYSIA "Penang Island (Baker 9075)" (worker); Schwarz 1937: 309; Schwarz 1939a: 112
(synonymy); Moure 1961: 209 (systematic position); Sakagami 1978: 200 (synonymy); Sakagami et al.
1990: 129 (synonymy); Wu 2000: 391 (synonymy (as penengensis!)).

TERMS OF USE
Tetragonula praeterita (Walker, 1860)
Trigona prterita Walker 1860: 305-306: Holotype (BMNH 17b.1185, worker): examined, "63 / 52", "Type"
(green border), "prterita / 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); "Cameron
Highland" (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 (comparative
note); 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 (bhnga), honey, nest, taxonomy); Type locality: INDIA "Mainpur, 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-Neto
1951: (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, nec
Lamarck. Lamarck 1817 (and repeated in 1835) only listed Melipona favosa, M. amalthea, M. ruficrus, M.
postica, and M. pallida. The replacement name therefore remains enigmatic, unless mistaken for M. 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-

TERMS OF USE
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); Cockerell
1929e: 242-243 (comparative note, distribution, taxonomy); SOLOMON ISLANDS, Lavoro Plantations,
Gaudalcanal Is.; New Georgia, W. Solomon; Cockerell 1936: 225 (distribution); SOLOMON ISLANDS,
Halaita, Nggela; Schwarz 1939a: 111, 113 (synonymy); Krombein 1949: 14 (distribution); Krombein
1951: 295 (synonymy); Moure 1961: 209 (systematic position); Michener 1965: 231 (citation); 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; Adelaide
River; 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); Floral
record: Corymbia torelliana.
Trigona wallacei Tadauchi et al. 1998: 245: Nomen nudum. The name is unavailable for taxonomic purposes,
as no description or definition of the taxon was provided (ICZN, art. 15.1) (type depositories).
Tetragonula sarawakensis (Schwarz, 1937)
Trigona sarawakensis Schwarz 1937: 283, 290, 313-315*, 316, 328, plate 2, 4: Holotype (BMNH 17b.1110);
paratypes (AMNH, BMNH?) (key to species, taxonomy); Type locality: MALAYSIA "Sarawak: Mt.
Dulit, 4000 ft." (2 workers Oct. 18, 1932; 2 workers Oct. 19, 1932; 1 worker Oct. 22, 1932); Schwarz
1939a: 85, 92, 106* (distribution, key to species, taxonomy); Schwarz 1948: 118; Sakagami 1959: 119-
120 (distribution, morphology, variation); SINGAPORE; Moure 1961: 210 (morphology, systematic
position); Yoshikawa et al. 1969: 168, 169 (nest); Wille & Michener 1973: 14, 25, 59 (nest); Sakagami
1978: 166-194, 210-211*, 228, 229, 230, 236, 237, 238 (taxonomy); Sakagami et al. 1983b: 2, 10 (nest);
Sakagami et al. 1983a: 106 (comparative note); Sakagami & Inoue 1985: 174, 176, 177, 178, 179, 180,
181, 182*, 184, 186 (key to species, taxonomy); Sakagami et al. 1985: (distribution); Starr & Sakagami
1987: 97 (citation); Sakagami et al. 1990: 131; Nagamitsu & Inoue 2005: 77 (distribution, foraging);
Klakasikorn et al. 2005: 6 (citation); Rasmussen & Cameron 2007: (systematics).
Tetragonula sirindhornae (Michener & Boongird, 2004)
Trigona (Tetragonula) sirindhornae Michener & Boongird 2004: 143-146: Holotype (SEMC, worker);
paratypes (AMNH (3), Insect Museum, Chakthong Building, Department of Agriculture, Bangkok, Thai-
land (3), SEMK (11) (taxonomy); Type locality: THAILAND "Thailand, on the Isthmus of Kra: Ranong
Province: Muang district, 913 November 2002" (11 workers); "Ngow Waterfall National Park" (7 work-

TERMS OF USE
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; Schwarz
1948: 118, 233; Krombein 1950: 102, 133, 135 (distribution, morphology); MICRONESIA, Caroline
Islands; Schwarz 1951: 63 (distribution); CAMBODIA, Angkor; Mt. Suept; Sakagami 1960: 146 (com-
parative note); Moure 1961: 208 (morphology, systematic position); Michener 1961: 28 (comparative
note); Sakagami & Yoshikawa 1961: 435 (comparative note, distribution); CAMBODIA, Phnom Penh;
LAOS, Thakhek; THAILAND, Mae Fack; Baltazar 1966: 407 (distribution); PHILIPPINES, Luzon,
Laguna; Mindanao, Davao; Sakagami 1978: 166, 197, 200, 201, 205, 206, 208 (synonymy); Sakagami et
al. 1983a: 101 (comparative note); Camargo 1988: 372 (synonymy); Sakagami et al. 1990: 129, 130
(synonymy, variation); Ikudome & Kusigemati 1996: 20 (taxonomy); Wu 2000: 391 (synonymy);
Klakasikorn et al. 2005: 6 (citation).
Tetragonula zucchii (Sakagami, 1978)
Trigona (Tetragonula) zucchii Sakagami 1978: 166-194, 196, 197, 198, 208-210*, 228, 229, 236, 237, 238,
247, plate 5: Holotype (SEHU) (nest, taxonomy); Type locality: MALAYSIA "Fraser's Hill-b, Malaya"
(9 workers, holotype, 8 paratypes); "Cameron Highlands" (8 worker); "Gombak" (1 worker); "Penang" (1
worker); "Templer Park" (17 workers); Yoshikawa et al. 1969: 173 (citation (as iridipennis)); Sakagami
et al. 1985: (distribution); Sakagami et al. 1990: 131 (distribution); Tadauchi et al. 1998: 244 (type
depositories); Nagamitsu & Inoue 2005: 77 (distribution, foraging); Rasmussen & Cameron 2007:
(systematics).
Tetragonula indet. Anderson et al. 1982: (pollination); Floral record: Mangifera indica; Camargo 1988:
372 (taxonomy); Drumond et al. 1996a: 396, 398, 399 (behavioral comparison); Dollin et al. 1997: (tax-
onomy); Roubik 2006: 126, 127, 130, 132 (nest).
Tetrigona
Tetrigona apicalis (Smith, 1857)

TERMS OF USE
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)); Beccari
1904: 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; Schwarz
1937: 282, 283, 288, 297, 298, 300-302*, 303, 304, 326 (key to species, taxonomy); Schwarz 1939a: 85,
94, 95, 100, 121, 122, 126-128*, 129, 130, 133 (distribution, key to species, taxonomy); Schwarz 1948:
19, 23, 91, 117, 141, 165, 290; Pagden 1957: 17-18 (flight); Sakagami 1960: 146 (distribution); 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 trunks
of Ficus: Moraceae; Durio zibethinus: Bombacaceae; crevices)); INDONESIA, Sumatera Barat; Inoue et
al. 1990b: (population); Johnson & Appanah 1990: (predator (Oecophylla: Formicidae: Hymenoptera));
Roubik 1993: 541 (illustration, nest); Michener & Roubik 1993: 260 (comparative note, morphology);
Cunha 1994: (systematics); Yamane et al. 1995: 294 (synonymy); Roubik 1996: 61, 62 (distribution,
nest); BRUNEI, Kuala Belalong; Nagamitsu & Inoue 1997: (foraging, pollination); Floral record: 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); Michener
2000: 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

TERMS OF USE
note).
Trigona hemileuca Cockerell 1929b: 140: Holotype (BMNH 17b.1098) (comparative note, taxonomy); Type
locality: THAILAND "Siam: Nan, Dec. 31 and Jan. 31 (Cockerell)" (worker); CAMBODIA, Ban Maa
Hia; Mekami river; Cockerell 1929c: 591 (distribution); CAMBODIA, W Weing Sa; Schwarz 1937: 300,
302; Schwarz 1939a: 126 (synonymy); Moure 1961: 216, 217 (morphology, systematic position); 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); Type
locality: MALAYSIA/INDONESIA "Meliau (Borneo) 4. Juni 1932"; "Sanggau (Borneo) 24. Juli 1932"
(several workers); Schwarz 1937: 282, 300, 301, 302, 327; Schwarz 1939a: 126 (synonymy); Moure
1961: 216 (synonymy).
Tetrigona binghami (Schwarz, 1937)
Trigona apicalis variety binghami Schwarz 1937: 288, 300, 301, 303-304*, 328, plate 2, 5, 7: Holotype
(BMNH 17b.1142); paratypes (AMNH, BMNH, USNM) (key to species, taxonomy); Type locality:
MYANMAR "Tenasserim: Dawnat Range, Jan., 1891 (Col. C.T. Bingham)" (1 worker); Bingham 1897:
560 (as apicalis, but most belong to this taxon); Cockerell 1918: 387 (distribution, key to species (as 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, tree
preference); Eltz 2004: 322 (abundance); Nagamitsu & Inoue 2005: 76 (distribution, foraging);
Klakasikorn et al. 2005: 2, 3, 5, 6 (floral record, morphology); Floral record: Tectona grandis; 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 synonymon
of vidua. See below (comparative note, taxonomy); Type locality: THAILAND "Siam: Nan, Jan. 13,
1928 (Alice Mackie)" (worker); Schwarz 1937: 300, 302, 304; Schwarz 1939a: 85, 94, 95, 98, 129*, 130
(distribution, key to species, taxonomy); Schwarz 1948: 117; Moure 1961: 216-217 (morphology, 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); Rajitparinya
2000: (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-

TERMS OF USE
tion); Type locality: MALAYSIA "Perak, F.M.S., Batang Padang, Jor Camp, 1800 ft., June 4, 1923
(Pendlebury)"; CAMBODIA, Patalung; MALAYSIA, Kualu Lumpur, Gombak valley; Schwarz 1937:
302; Schwarz 1939a: 85, 94, 102, 127, 129-130* (distribution, key to species, taxonomy); Schwarz 1948:
117; Moure 1961: 216 (morphology, systematic position); Sakagami 1975: 61, 62, 63, 64* (distribution,
morphology, variation); MALAYSIA; THAILAND; Sakagami et al. 1985: (distribution, key to species);
Michener 1990: 127 (systematic position); Sakagami et al. 1990: 131, 136 (distribution, key to species);
San & Osawa 2002: (population); MALAYSIA, Negri Sembiran, Pasoh Forest Reserve; Nagamitsu &
Inoue 2005: 76 (distribution, foraging); Klakasikorn et al. 2005: 6 (citation).
Tetrigona vidua (Lepeletier de Saint Fargeau, 1836)
Melipona vidua Lepeletier de Saint Fargeau 1836: 429: Syntype (MNHN, 1 worker): putative syntype
examined, "Museum Paris / Bengale / Diard & Duvaucel 1815", "vidua", "Diard et Duvaucel", "type",
"TYPE", "M. Vidua / Lep. S. Farg / Bengale". I consider this a true type: Alfred Duvaucel (1793-1825)
and Pierre-Mdard Diard (1794-1863) arrived on their first expedition to Kolkata, India, in January 1818
and spent years collecting in the Indo-Malayan region. The label date is wrong as they were still in France
in 1815, but did reside for long periods in India later (Claeys: 1954) and I consider the label provenance
merely the port of embarkation of the specimen, rather than the collecting locality. The type is similar to
melanoleuca and could be considered a senior synonymon of that. However, melanoleuca is a variable
taxon, and is here retained until further studies may document if in fact it is only a single taxon (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; Castets
1893: 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 bees
settled 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; Moure
1961: 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 (key
to species, taxonomy (as Melipona)); Waterhouse 1900: (nest); Cockerell 1920a: 116 (distribution);
Cockerell 1930b: 149; Rayment 1932a: (morphology, nest); Bibby 1947: 78 (distribution); Michener
1961: (common name (native bee; sweat bee), nest); Michener 1965: 231 (citation); Michener 1965:
230; Smythe 1970: (pollination (as Trigona)); Floral record: Cymbidium madidum; Dendrobium; Frith
1978: (nest (in Rhizophora apiculata: Rhizophoraceae); predator (Oecophylla smaragdina: Formicidae:
Hymenoptera)); Schneider 1982: (pollination); Appanah 1982: (pollination); MALAYSIA, Negri 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);

TERMS OF USE
Raju & Reddi 1989: (pollination); Floral record: Anisomeles indica; Anisomeles malabarica; Heard et
al. 1990: 1006, 1007, 1012 (pollination); Floral record: Anacardium occidentale; Boonithee et al. 1991:
(pollination); Adams et al. 1992: (orchid pollination); Appanah 1993: 466 (pollination); Adams & Law-
son 1993: (pollination (as Trigona)); Floral record: Caladenia carnea; Cymbidium madidum; Cymbidium
suave; Dendrobium adae; Dendrobium kingianum; Dendrobium monophyllum; Dendrobium speciosum;
Dendrobium lichenastrum; Sarcochilus moorei; Pomatocalpa macphersonii; Heard 1993: (pollination
(as spp.)); Floral record: Macadamia integrifolia; Forster et al. 1994: (pollination); Rinderer et al. 1996:
(foraging); Aluri & Subba Reddi 1996: (pollination); Momose et al. 1998: (pollination); Heard 1998:
(pollination (as spp.)); Floral record: Macadamia integrifolia; Crane 1999: 86-90 (honey); Viraktamath
et al. 1999: (pollen); Heard 1999b: (pollination (as spp.)); Floral record: Mangifera indica; Yumoto
2000: (pollination); Heard 2000: (pollination (as spp.)); Jagadish et al. 2002: (common name (dammer
bee), pollination); Floral record: Hygrophila auriculata; Leucas aspera; Tangmitcharoen et al. 2002;
Chourykaew et al. 2002: 35, 36, 38, 41 (pollination (contrary to conclusion)); Floral record: Afgekia 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; Telm, Perak-Pahang boundary, 4,000 feet" (male); Melipona basimaculata
Bingham 1905: 60, plate A11 (citation).
Trigona pallidicincta Cockerell 1918: 386: Holotype (BMNH 17b.1134, male) (key to species, comparative
notes, taxonomy); Type locality: SINGAPORE "Singapore (Baker)" (male).
Nomina nuda
Melipona pulla Illiger 1806: 158: Nomen nudum. The name is unavailable for taxonomic purposes, as no
description or definition of the taxon was provided (ICZN, art. 12.1) (citation); INDONESIA, Sumatra;
Melipona pulla Dalla Torre 1896: 582 (citation).
Melipona Khasiana Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as no
description or definition of the taxon was provided (ICZN, art. 13.1.1) (common name (nagp hamang),
honey, nest).
Melipona terrestris Pugh 1947: 62: Nomen nudum. The name is unavailable for taxonomic purposes, as no
description or definition of the taxon was provided (ICZN, art. 13.1.1) (common name (nagp khyndew),
honey, nest).
Trigona (Pacificotrigona) okazawai Tadauchi et al. 1998: 245: Nomen nudum (as Trigona (Pacificotrigona)
okazawai irianensis). The name is unavailable for taxonomic purposes, as no description or definition of
the taxon was provided (ICZN, art. 15.1).
Trigona (Pacificotrigona) okazawai irianensis Tadauchi et al. 1998: 245: Nomen nudum. The name is

TERMS OF USE
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)

TERMS OF USE
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)

TERMS OF USE
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)

TERMS OF USE
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)

TERMS OF USE
Appendix 2. Stingless bee taxon name followed by current combination or synonymy
aliceae, Homotrigona aliceae (Cockerell, 1929)

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

TERMS OF USE
hemileuca, see apicalis

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

TERMS OF USE
sericea, see apicalis

sirindhornae, Tetragonula sirindhornae (Michener & Boongird, 2004)
smithii, see ruficornis
symei, Austroplebeia symei (Rayment, 1932)
terminata, Lepidotrigona terminata (Smith, 1878)
terrestris, nomen nudum
testaceinerva, see melanocephala
testaceitarsis, Tetragonula testaceitarsis (Cameron, 1901)
thoracica, Geniotrigona thoracica (Smith, 1857)
trochanterica, Lepidotrigona trochanterica (Cockerell, 1920)
tunneli, nomen nudum, see Lepidotrigona
valdezi, Tetragonula valdezi (Cockerell, 1918)
ventralis, Lepidotrigona ventralis (Smith, 1857)
versicolor, see fimbriata
vidua, Tetrigona vidua (Lepeletier de Saint Fargeau, 1836)
wallacei, nomen nudum, see sapiens
websteri, Austroplebeia websteri (Rayment, 1932)
wybenica, see clypearis
zucchii, Tetragonula zucchii (Sakagami, 1978)
Appendix 3. Museum acronyms. Curators and sources of information concerning the collection are
listed in parenthesis
AMNH USA, New York, New York, American Museum of Natural History (Jerome G.
Rozen, John S. Ascher)
AMS Australia, New South Wales, Sydney, Australian Museum (David Britton, Max
Beatson)
ANIC Australia, Australian Capital Territory, Canberra City, CSIRO, Australian
National Insect Collection (John La Salle, Nicole Fisher)
BMNH United Kingdom, London, The Natural History Museum (David Notton)
BPBM USA, Hawaii, Honolulu, Bernice P. Bishop Museum (Shepherd P. Myers)
Calicut India, Kerala, Calicut University, Zoology Department
DEI Germany, Mncheberg, 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 (Sndor Cssz)
IEBR Vietnam, Hanoi, Institute of Ecology and Biological Resources
MNHN France, Paris, Musum National d'Histoire Naturelle (Claire Villemant)
MSNG (Collezione Gribodo)Italy, Genova, Museo Civico di Storia Naturale "Giacomo Doria" (Fabio Penati)
MVMA Australia, Victoria, Abbotsford, Museum of Victoria (Ken Walker)
MZB Indonesia, Bogor, Museum Zoologicum Bogoriense (Yayuk R. Suhardjono)
OUMNH (=Wilson Saunders collection)United Kingdom, Oxford, University Museum of Natural History
(James Hogan)
QM Australia, Queensland, South Brisbane, Queensland Museum
RMNH Netherlands, Leiden, Nationaal Natuurhistorische Museum (Naturalis) (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

TERMS OF USE
SFS Shichi F. Sakagamis collection (see SEHU)

SMNS Germany, Stuttgart, Staatliches Museum fr Naturkunde
USNM USA, Washington D.C., National Museum of Natural History, (formerly, United
States National Museum) (David Furth, Brian Harris)
ZMHB Germany, Berlin, Museum fr Naturkunde der Humboldt-Universitt (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, 97101.
Adams, P.B. & Lawson, S.B. (1993) Pollination in Australian orchids: a critical assessment of the literature 18821992.
Australian Journal of Botany, 41, 553575.
Adams, P.B. & Lawson, S.D. (1988) Multiple bee pollinators of Dendrobium kingianum Bidw. in the natural habitat. The
Orchadian (Australasian Native Orchid Society), 9, 103, 105107.
Aluri, R.J.S. & Subba Reddi, C. (1996) Floral biology and pollination in Gliricidia sepium (Fabaceae). Journal of Nature
Conservation, 8, 6567.
Amano, K. (2004). Attempts to introduce stingless bees for the pollination of crops under greenhouse conditions in
Japan. 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, 183190.
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, 541548.
Anderson, J. (1890) The Selungs of the Mergui archipelago. Trbner & 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, 1134.
Appanah, S. (1993) Mass flowering of dipterocarp forests in the aseasonal tropics. Journal of Bioscience, 18, 457474.
Appanah, S., Willemstein, S.C., & Marshall, A.G. (1986) Pollen foraging by two Trigona colonies in a Malaysian rain
forest. Malayan Nature Journal, 39, 177191.
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, 259274.
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, 957971.
Atluri, J.B., Rao, S.P., & Reddi, C.S. (2000) Pollination ecology of Helicteres isora Linn. (Sterculiaceae). Current Sci-
ence (India), 78, 713718.
Atmowidjojo, A.H. & Adisoemarto, S. (1986) Potential pollentransferring insects of Indigofera spp. Treubia, 29, 225
235.
BacelarLima, C.G., Freire, D.d.C.B., ColettoSilva, A., Costa, K.B., Laray, J.P.B., VilasBoas, H.C., & CarvalhoZilse,
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 Amazona Cen-
tral, Brasil. Acta Amazonica, 36, 343348.
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, 1311, 14 pl.
Baker, E.W. & DelfinadoBaker, M. (1985) An unusual new species of Neocypholaelaps (Acari: Ameroseiidae) from the
nests of stingless bees (Apidae: Meliponinae). International Journal of Acarology, 11, 227232.
Baltazar, C.R. (1966) A catalogue of Philippine Hymenoptera (with a bibliography, 17581963). Pacific Insects Mono-
graph, 8, 1488.
Barile, G.E. & Cervancia, C.R. (1995) Foraging and nesting behavior of Trigona biroi Friese (Hymenoptera: Apidae).
Philippine Entomologist, 9, 555566.
Bartareau, T. (1994) The reproductive ecology of Dendrobium toressae (Bailey) Dockr., a geographically restricted spe-
cies in northeast Queensland. The Orchadian (Australasian Native Orchid Society), 11, 106111.
Bartareau, T. (1995) The pollination of Cymbidium madidum Lindley and C. suave R. Br. by native bees a case of floral
reward deception? The Orchadian (Australasian Native Orchid Society), 11, 487495.

TERMS OF USE
Bartareau, T. (1996) Foraging behaviour of Trigona carbonaria (Hymenoptera: Apidae) at multiple choice feeding sta-
tions. Australian Journal of Zoology, 44, 143153.
Beccari, O. (1904) Wanderings in the great forests of Borneo: Travels and researches of a naturalist in Sarawak [from
italian "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 stingless
bee, Nannotrigona testaceicornis, kept in a greenhouse (Hymenoptera, Apidae). Bulletin of the Faculty of Agricul-
ture (Shimane University), 23, 4654.
Bibby, F.F. (1947) Notes on the insect fauna of the Samar group, Philippines. Philippine Journal of Science, 77, 6181.
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, 6568.
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,
19011902. Zoology. Vol. I (appendix), University Press of Liverpool, London, pp. iiivii.
Bingham, C.T. (1905). Report on the 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,
19011902. Zoology. Vol. III, University Press of Liverpool, London, pp. 1660, plate A.
Blamires, S.J., Hochuli, D.F., & Thompson, M.B. (2008) Why cross the web: decoration spectral properties and prey
capture in an orb spider (Argiope keyserlingi) web. Biological Journal of the Linnean Society, 94, 221229.
Boonithee, A., Juntawong, N., Peckhacker, H., & Httinger, E. (1991) Floral visits to select crops by four Apis species
and Trigona sp. in Thailand. Acta Horticulturae (Wageningen), 288, 7480.
Borges, F.v.B. & Blochtein, B. (2005) Atividades externas de Melipona marginata obscurior Moure (Hymenoptera, Api-
dae), em distintas pocas do ano, em So Francisco de Paula, Rio Grande do Sul, Brasil. Revista Brasileira de Zoo-
logia, 22, 680686.
Bravo, F. (1992) Sistemtica e distribuo de Parapartamona Schwarz (Hymenoptera, Apidae, Meliponinae). Revista
Brasileira de Entomologia, 36, 863878.
Breed, M.D., McGlynn, T.P., Sanctuary, M.D., Stocker, E.M., & Cruz, R. (1999) Distribution and abundance of colonies
of selected meliponine species in a Costa Rican tropical wet forest. Journal of Tropical Ecology, 15, 765777.
Brown, R.E. (1906) A catalogue of Philippine Hymenoptera, with descriptions of new species. Philippine Journal of Sci-
ence, 1, 683695.
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, 38333845.
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, 3547.
Camargo, J.M.F. (1988) Meliponinae (Hymenoptera, Apidae) da coleo do "Instituto di entomologia agraria", Portici,
Itlia. Revista Brasileira de Entomologia, 32, 351374.
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 (Wille
and Chandler). American Museum Novitates, 3293, 124.
Camargo, J.M.F. & Moure, J.S. (1994) Meliponinae neotropicais: os gneros Paratrigona Schwarz, 1938 e Aparatrigona
Moure, 1951 (Hymenoptera, Apidae). Arquivos de Zoologia, 32, 33109.
Camargo, J.M.F. & Moure, J.S. (1996) Meliponini neotropicais: o gnero Geotrigona Moure, 1943 (Apinae, Apidae,
Hymenoptera), com especial referncia filogenia e biogeografia. Arquivos de Zoologia, 33, 95161.
Camargo, J.M.F. & Pedro, S.R.M. (2003a). Sobre as relaes filogenticas 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, Cricima, pp. 109122.
Camargo, J.M.F. & Pedro, S.R.M. (2003b) Meliponini neotropicais: o gnero Partamona Schwarz, 1939 (Hymenoptera,
Apidae, Apinae) bionomia e biogeografia. Revista Brasileira de Entomologia, 47, 311372.
Camargo, J.M.F. & Pedro, S.R.M. (2007). Meliponini. In: J.S. Moure, D. Urban & G.A.R. Melo (Eds.), Catalogue of
bees (Hymenoptera, Apoidea) in the Neotropical region, Sociedade Brasileira de Entomologia, Curitiba (Paran),
pp. 272578.
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, 213229.
Cameron, E.C., Franck, P., & Oldroyd, B.P. (2004) Genetic structure of nest aggregations and drone congregations of the
southeast Asian stingless bee Trigona collina. Molecular Ecology, 13, 23572364.

TERMS OF USE
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, 15144.
Cameron, P. (1901) On the Hymenoptera collected during the "Skeat Expedition" to the Malay Peninsula, 18991900.
Proceedings of the Zoological Society of London, 1901, 1644.
Cameron, P. (1902) On the Hymenoptera collected by Mr. Robert Shelford at Sarawak, and on the Hymenoptera of the
Sarawak museum. Journal of the Straits Branch of the Royal Asiatic Society, 37, 29131.
Cameron, P. (1908) On some bornean species of Trigona (Apidae). The Entomologist, 41, 192195.
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, 13.
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, 465488, plate.
Cepeda, O.I. (2006) Division of labor during brood production in stingless bees with special reference to individual par-
ticipation. Apidologie, 37, 175190.
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. 7880.
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, 3742.
Chinajariyawong, A. & Saiboon, S. (2000) Stingless bees in Wongkhla province. In Proceedings of the 38th Kasetsart
University Annual Conference: Plants, Agricultural Extension and Communication, pp. 445449, 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, 493503.
Chinh, T.X., Sommeijer, M.J., Boot, W.J., & Michener, C.D. (2005) Nest and colony characteristics of three stingless bee
species in Vietnam with the first description of the nest of Lisotrigona carpenteri (Hymenoptera: Apidae: Melipo-
nini). Journal of the Kansas Entomological Society, 78, 363372.
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, 3144.
Claeys, J.Y. (1954) Deux Franais la conqute de Singapour. Les naturalistes Alfred Duvaucel et PierreMdard Diard.
FranceAsie (Association pour une Meilleure Connaissance de l'Asie), 99, 10011007.
Cobara, B. & Dejan, A. (1998) A stingless bee nesting inside antgardens in French Guiana (Hymenoptera: Apidae).
Sociobiology, 32, 489492.
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, 199249.
Cockerell, T.D.A. (1911) The bees of the Solomon Islands. Proceedings of the Linnean Society of New South Wales, 36,
160178.
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, 409415.
Cockerell, T.D.A. (1915) Descriptions and records of bees. LXVIII. Annals and Magazine of Natural History, [8]16,
19.
Cockerell, T.D.A. (1918) Descriptions and records of bees. LXXX. Annals and Magazine of Natural History, [9]2,
384390.
Cockerell, T.D.A. (1919a) Descriptions and records of bees. LXXXIII. Annals and Magazine of Natural History, [9]3,
118125.
Cockerell, T.D.A. (1919b) Descriptions and records of bees. LXXXV. Annals and Magazine of Natural History, [9]3,
240250.
Cockerell, T.D.A. (1919c) The social bees of the Philippine Islands. Philippine Journal of Science, 14, 7781.
Cockerell, T.D.A. (1920a) Descriptions and records of bees. LXXXVIII. Annals and Magazine of Natural History,
[9]5, 113119.
Cockerell, T.D.A. (1920b) Some bees from Sandakan, Borneo. Philippine Journal of Science, 17, 221229.
Cockerell, T.D.A. (1921) Fossil arthropods in the British Museum. VII. Annals and Magazine of Natural History, [9]8,
541545.
Cockerell, T.D.A. (1922a) Australian bees in the Queensland museum. Memoirs of the Queensland Museum, 7, 257279.
Cockerell, T.D.A. (1922b) Fossils in burmese amber. Nature, 109, 713714.
Cockerell, T.D.A. (1923) Descriptions and records of bees. XCVIII. Annals and Magazine of Natural History, [9]12,
238247.
Cockerell, T.D.A. (1925a) Descriptions and records of bees. CIV. Annals and Magazine of Natural History, [9]15,

TERMS OF USE
489496.
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, 5355.
Cockerell, T.D.A. (1926a) Descriptions and records of bees. CXII. Annals and Magazine of Natural History, [9]18,
216227.
Cockerell, T.D.A. (1926b) Trigona laeviceps not a pest. Journal of Economic Entomology, 19, 188189.
Cockerell, T.D.A. (1927) Bees from the Malay Peninsula. Annals and Magazine of Natural History, [9]20, 530541.
Cockerell, T.D.A. (1929a) Bees from the Australian region. American Museum Novitates, 346, 118.
Cockerell, T.D.A. (1929b) Descriptions and records of bees. CXVII. Annals and Magazine of Natural History, [10]4,
132141.
Cockerell, T.D.A. (1929c) Descriptions and records of bees. CXX. Annals and Magazine of Natural History, [10]4,
584594.
Cockerell, T.D.A. (1929d) Bees in the Queensland Museum. Memoirs of the Queensland Museum, 9, 298323 [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, 199243.
Cockerell, T.D.A. (1930a) Descriptions and records of bees. CXXII. Annals and Magazine of Natural History, [10]5,
156163.
Cockerell, T.D.A. (1930b) The bees of Australia. Australian Zoologist, 6, 137156, 205238.
Cockerell, T.D.A. (1930c) Australian bees in the Museum of Comparative Zoology. Psyche, 37, 141154.
Cockerell, T.D.A. (1934) Some African meliponine bees. Revue de Zoologie et de Botanique Africaines, 26, 4662.
Cockerell, T.D.A. (1936) Bees from the Solomon Islands. Proceedings of the Royal Entomological Society of London,
Series B, Taxonomy, 5, 225226.
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, 6166.
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, 133140.
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.
CortopassiLaurino, M., Imperatriz-Fonseca, V.L., Roubik, D.W., Dollin, A.E., Heard, T.A., Aguilar, I., Venturieri, G.C.,
Eardley, C.D., & NogueiraNeto, P. (2006) Global meliponiculture: challenges and opportunities. Apidologie, 37,
275292.
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,
187190.
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, 7384.
Crane, E. (1999) The World History of Beekeeping and Honey Hunting. Routledge, New York, 682 pp.
Cunha, R.A.d. (1994) Estudo fentico supraespecfico de Trigonini do Velho Mundo (Hymenoptera, Apidae). Revista
Brasileira de Entomologia, 38, 613626.
Cunningham, S.A., Fitzgibbon, F., & Heard, T.A. (2002) The future of pollination in Australia. Australian Journal of
Agricultural Research, 53, 893900.
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 Mlipones (Hymnoptres Apid) a la lumire des travaux rcents.
L'Anne Biologique (Paris), 8, 455490.
Darchen, R. & Louis, J. (1961) Les mlipones et leur levage. Melipona Trigona Lestrimelitta. Annales de l'Institut
National de la Recherche Agronomique. Srie C bis, Annales de l'abeille, 4, 539.
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, 229239.
Divan, V.V. (1981("1980")) Honey bee behaviour during total solar eclipse. Indian Bee Journal, 42, 9799.
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,

TERMS OF USE
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, 219223.
Dollin, A.E. & Dollin, L.J. (1994) An elusive Australian native bee drone the key to renaming a species. The Australa-
sian beekeeper, 95, 291292.
Dollin, A.E., Dollin, L.J., & Sakagami, S.F. (1997) Australian stingless bees of the genus Trigona (Hymenoptera: Api-
dae). Invertebrate Taxonomy, 11, 861896.
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, 104107.
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. Srie Zoologia (Porto
Alegre), 79, 3945.
Drumond, P.M., Oldroyd, B.P., Dollin, A.E., & Dollin, L.J. (1999) Oviposition behaviour of two Australian stingless
bees, Austroplebeia symei Rayment and Austroplebeia australis Friese (Hymenoptera: Apidae: Meliponini). Austra-
lian Journal of Entomology, 38, 234241.
Drumond, P.M., Oldroyd, B.P., & Osborne, K. (2000) Worker reproduction in Austroplebeia australia Friese
(Hymenoptera, Apidae, Meliponini). Insectes Sociaux, 47, 333336.
Drumond, P.M., Zucchi, R., Mateus, S., Bego, L.R., Yamane, S., & Sakagami, S.F. (1996a) Oviposition behavior of the
stingless bees. XVII. Plebeia (Plebeia) droryana and an ethological comparison with other Meliponine taxa
(Hymenoptera, Apidae). Japanese Journal of Entomology, 64, 385400.
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, 3155.
Eardley, C.D. (2004) Taxonomic revision of the African stingless bees (Apoidea: Apidae: Apinae: Meliponini). African
Plant Protection, 10, 6396.
Eltz, T. (2004) Spatio-temporal variation of apine bee attraction to honeybaits in Bornean forests. Journal of Tropical
Ecology, 20, 317324.
Eltz, T., Brhl, C., van der Kaars, S., & Linsenmair, K.E. (2002a) Determinants of stingless bee nest density in lowland
dipterocarp forests of Sabah, Malaysia. Oecologia, 131, 2734.
Eltz, T., Brhl, C.A., & Grke, C. (2002b) Collection of mold (Rhizopus sp.) spores in lieu of pollen by the stingless bee
Trigona collina. Insectes Sociaux, 49, 2830.
Eltz, T., Brhl, 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 Ecology
and Management, 172, 301313.
Eltz, T., Brhl, 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,
273279.
Eltz, T., Brhl, C.A., van der Kaars, S., & Linsenmair, K.E. (2001b) Assessing stingless bee pollen diet by analysis of
garbage pellets: a new method. Apidologie, 32, 341353.
Engel, M.S. (2000) A review of the Indo-Malayan Meliponine genus Lisotrigona, with two new species (Hymenoptera:
Apidae). Oriental Insects, 34, 229237.
Engel, M.S. (2001) A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the
American Museum of Natural History, 259, 1192.
Engels, W. & Imperatriz-Fonseca, V.L. (1990). Caste development, reproductive strategies, and control of fertility in
honey bees and stingless bees. In: W. Engels (Ed.), Social insects, an evolutionary approach to castes and reproduc-

TERMS OF USE
tion, Springer, Berlin, pp. 167230.

Esaki, T. (1936) Einige biologische Beobachtungen ber die Bienen und Wespen Mikronesiens. Mushi, 9, 4447.
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 chowchow (Sechium edule). Journal of Ecobiol-
ogy (India), 17, 509516.
Eswarappa, G., Kuberappa, G.C., Sreenath, T.N., Sreeramulu, K.R., & Gowda, G. (2005b) Role of different species of
honeybees in cross pollination of chowchow (Sechium edule). Journal of Ecobiology (India), 17, 101107.
Eswarappa, G., Kuberappa, G.C., Sreenath, T.N., Sreeramulu, K.R., & Gowda, G. (2005c) Nectar secretion and its sugar
concentration on foraging activity of different species of honey bees in chowchow (Sechium edule). Journal of
Ecobiology (India), 17, 337343.
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, 7180.
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, 1387.
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), associated
with the stingless bee Trigona carbonaria Smith in Australia. Bulletin de l'Institut Royal des Sciences Naturelles de
Belgique, Entomologie, 57, 197202.
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, 133140.
Forster, P.I., Machin, P.J., Mound, L., & Wilson, G.W. (1994) Insects associated with reproductive structures of cycads in
Queensland and northeast New South Wales, Australia. Biotropica, 26, 217222.
Franck, P., Cameron, E.C., Good, G., Rasplus, J.Y., & Oldroyd, B.P. (2004) Nest architecture and genetic differentiation
in a species complex of Australian stingless bees. Molecular Ecology, 13, 23172331.
Friese, H. (1898) Die TrigonaArten Australiens. Termszetrajzi Fzetek, 21, 427431.
Friese, H. (1900) Neue Arten der Bienengattungen Melipona Ill., und Trigona Jur. Termszetrajzi Fzetek, 23, 381394.
Friese, H. (1901) Neue Arten der Bienengattung Trigona Jur. (Hym.). Zeitschrift fr systematische Hymenopterologie
und Dipterologie, 1, 265271.
Friese, H. (1904) Eine Bienenausbeute von Java (Hym.). Allgemeine Zeitschrift fr Entomologie, 9, 138140.
Friese, H. (1905) Ein Bienennest mit Vorratskammern (Lithurgus dentipes Sm.). Zeitschrift fr wissenschaftliche Insek-
tenbiologie, 1, 118119.
Friese, H. (1909) Die Bienenfauna von NeuGuinea. Annales HistoricoNaturales Musei Nationalis Hungarici (Ter-
mszettudomnyi Mzeum vknyve), 7, 179288.
Friese, H. (1909("1908")). Hymenoptera II. Apidae. In: A. Wichmann (Ed.), Nova Guinea. Rsultats de l'expdition sci-
entifique nerlandaise la NouvelleGuine en 1903. Vol. 5 (Zoologie), E.J. Brill, Leiden, pp. 353359.
Friese, H. (1912) Zur Bienenfauna von Neu-Guinea und Oceanien. bersicht des von den Herren Dahl, Finsch, Krmer,
Moszkowski, Neuhaus, Schoede und L. Schultze in den letzten Jahren dem Zoologischen Museum in Berlin ber-
wiesenen Forschungsmaterials. Mitteilungen aus dem Zoologischen Museum in Berlin, 6, 9196.
Friese, H. (1914a) Die Bienenfauna von Java. Tijdschrift voor Entomologie, 57, 113.
Friese, H. (1914b) Anhang zur: "Bienenfauna von Java". Tijdschrift voor Entomologie, 57, 1461.
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, 14.
Friese, H. (1917) Results of Dr. E. Mjberg's Swedish scientific expeditions to Australia 19101913. 13. Apidae. Arkiv
fr Zoologi (Stockholm), 11(2), 19.
Friese, H. (1918) Wissenschaftliche Ergebnisse einer Forschungsreise nach Ostindien, ausgefhrt im Auftrage der Kgl.
Preu. Akademie der Wissenschaften zu Berlin von H. v. Buttel-Reepen. VII. Bienen aus Sumatra, Java, Malakka
und Ceylon. Gesammelt von Herrn Prof. Dr. v. Buttel-Reepen in den Jahren 19111912. Zoologische Jahrbcher,
Abteilung fr Systematik, Geographie und Biologie der Tiere, 41, 489520.
Friese, H. (1925) Neue neotropischen Bienenarten, zugleich II. Nachtrag zur Bienenfauna von Costa Rica (Hym.). Stet-
tiner Entomologische Zeitung, 86, 141.
Friese, H. (1933a) Neue TrigonaArten von Borneo (Hym.). Natuurhistorisch Maandblad, 22, 4546.
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, 194195.
Gardener, M.C., Rowe, R.J., & Gillman, M.P. (2003) Tropical bees (Trigona hockingsi) show no preference for nectar
with amino acids. Biotropica, 35, 119125.
George, C.J. (1933) Excretion in Melipona during metamorphosis. Journal of the University of Bombay, 1, 5861.
George, C.J. (1934) The bionomics, morphology, and metamorphosis of Melipona irridipennis (sic). Journal of the Uni-

TERMS OF USE
versity of Bombay, 2, 116.

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, 412417.
Goebel, R.L. (1986) Australian native bees. Queensland Agricultural Journal, 112, 285286.
Goel, S.C. & Kumar, A. (1990) Insect pests and predators associated to sunflower in winters of northern India. Indian
Journal of Entomology, 52, 3945.
Graur, D. (1985) Gene diversity in Hymenoptera. Evolution, 39, 190199.
Greco, M., SpoonerHart, 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, 97100.
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, 8992.
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, 196202.
Gribodo, G. (1891) Contribuzioni imenotterologiche sopra alcune specie nuove o poco conosciute di imenotteri antofili
(Generi Ctenoplectra, Xylocopa, Centris, Trigona e Bombus). Bollettino della Societ Entomologica Italiana, Gen-
ova, 23, 102119.
Gribodo, G. (1893) Note imenotterologische. Nota II. Nuovi generi e nuove specie di imenotteri antofili ed osservazioni
sopra alcune specie gi conosciute. Bollettino della Societ Entomologica Italiana, Genova, 25, 248287.
Grimaldi, D.A., Shedrinsky, A., Ross, A.J., & Baer, N.S. (1995) Forgeries of fossils in amber: history, identification
and case studies. Curator (New York), 37, 251274.
Gross, C.L. (1993) The breeding system and pollinators of Melastoma affine (Melastomataceae); a pioneer shrub in trop-
ical Australia. Biotropica, 25, 468474.
Haddon, A.C. (1901) The omen animals of Sarawak. Popular Science Monthly, 60, 8088.
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 stingless
bees. Apidologie, 37, 144163.
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, 1220.
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, 303304.
Heard, T.A. (1993) Pollinator requirements and flowering patterns of Macademia integrifolia. Australian Journal of Bot-
any, 41, 491497.
Heard, T.A. (1994) Behaviour and pollinator efficiency of stingless bees and honey bees on macadamia flowers. Journal
of Apicultural Research, 33, 191198.
Heard, T.A. (1996) Stingless bees. Nature Australia, 1996(spring), 5055.
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, 183206.
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, 2021.
Heard, T.A. (2003) Native stingless bees: biology, conservation and economic utilisation. News Bulletin of the Entomo-
logical Society of Queensland, 31, 134141.
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, 91100.
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, 343353.
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, 11011114.
Hedicke, H. (1926) Beitrge zur Apidenfauna der Philippinen (Hym.). (2. Beitrag zur Kenntnis orientalischer Apiden).
Deutsche Entomologische Zeitschrift, 1926, 413423.

TERMS OF USE
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, S196S198.
Hilrio, S.D., Imperatriz-Fonseca, V.L., & Leinert, A.d.M.P. (2000) Flight activity and colony strength in the stingless
bee Melipona bicolor bicolor (Apidae, Meliponinae). Revista Brasileira de Biologia, 60, 299306.
Hilrio, S.D., Imperatriz-Fonseca, V.L., & Leinert, A.d.M.P. (2001) Responses to climatic factors by foragers of Plebeia
pugnax Moure (in litt.) (Apidae, Meliponinae). Revista Brasileira de Biologia, 61, 191196.
Hockings, H.J. (1884) Notes on two Australian species of Trigona. Transactions of the Entomological Society of London,
32, 149157.
Hoese, F.J., Law, E.A.J., Rao, D., & Heberstein, M.E. (2006) Distinctive yellow bands on a sit-and-wait predator: prey
attractant or camouflage? Behaviour, 143, 763781.
Hofstede, F.E. & Sommeijer, M.J. (2006) Effect of food availability on individual foraging specialisation in the stingless
bee Plebeia tobagoensis (Hymenoptera, Meliponini). Apidologie, 37, 387397.
Hohmann, H. (2003) Zur Kommunikation stachelloser Bienen in Neuguinea. TenDenZen (Jahrbuch berseemuseum
Bremen), 11, 926.
Horne, C. & Smith, F. (1870) Notes on the habits of some hymenopterous insects from the North-West provinces of
India. 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, 161196.
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, 465492.
Ihering, H.v. (1912) Zur Biologie der brasilianischen Meliponiden. Zeitschrift fr wissenschaftliche Insektenbiologie, 8,
15, 4346.
Ikudome, S. & Kusigemati, K. (1996) Notes on some bees from the Palau Islands (Hymenoptera, Apoidea). Kagoshima
University Research Center for the South Pacific Occasional Papers, 30, 1721.
Illiger, K. (1806) William Kirby's Familien der Bienenartigen Insekten mit Zustzen, Nachweisungen und Bemerkungen.
Magazin fr Insektenkunde, 5, 28175.
Imperatriz-Fonseca, V.L. & Zucchi, R. (1995) Virgin queens in stingless bee (Apidae, Meliponinae) colonies: A review.
Apidologie, 26, 231244.
Inoue, K., Kato, M., & Inoue, T. (1995) Pollination ecology of Dendrobium setifolium, Neuwiedia borneensis, and
Lecanorchis multiflora (Orchidaceae) in Sarawak. Tropics, 5, 95100.
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. 377400.
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 (International
union for the study of social insects), Oxford, New Delhi, pp. 581582.
Inoue, T., Adri, & Salmah, S. (1990a). Nest site selection and reproductive ecology of the Asian honey bee, Apis cerana
indica, 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. 219232.
Inoue, T., Nakamura, K., Salmah, S., & Abbas, I. (1993) Population dynamics of animals in unpredictably-changing
tropical environments. Journal of Bioscience, 18, 425455.
Inoue, T., Roubik, D.W., & Suka, T. (1999) Nestmate recognition in the stingless bee Melipona panamica (Apidae, Meli-
ponini). Insectes Sociaux, 46, 208218.
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 bee
Trigona (Tetragonula) laeviceps. Journal of Apicultural Research, 23, 136142.
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, 100111.
Inoue, T., Salmah, S., Abbas, I., & Yusuf, E. (1985a) Foraging behavior of individual workers and foraging dynamics of
colonies of three sumatran stingless bees. Researches on Population Ecology (Kyoto), 27, 373392.
Inoue, T., Salmah, S., & Sakagami, S.F. (1996b) Individual variations in worker polyethism of the Sumatran stingless
bee, Trigona (Tetragonula) minangkabau (Apidae, Meliponinae). Japanese Journal of Entomology, 64, 641668.
Inoue, T., Salmah, S., Sakagami, S.F., Yamane, S., & Kato, M. (1990b). An analysis of anthophilous insects in central
Sumatra. 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. 175200.
Inoue, T., Salmah, S., & Yusuf, E. (1985b). Foraging behavior of individual workers and foraging dynamics of colonies
of three Sumatran stingless bees [summary of Researches on Population Ecology (Kyoto), 27: 373392]. In:
Sumatra nature study (Entomology) (Ed.), Evolutionary ecology of insects in humid tropics, especially in central
Sumatra, Kanazawa University, Kanazawa (Japan), pp. 4449.

TERMS OF USE
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, Thong
Pha Phum district, Kanchanaburi Province]. BRT (The Thai Response to Biodiversity), 2549, 2031.
International Commission on Zoological Nomenclature (1999) International code of zoological nomenclature, fourth
edition. 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 bee
Trigona carbonaria. International Journal of Antimicrobial Agents, 32, 8998.
Jagadish, K.S., Venkatesh-Gowda, T.N., Lakshmikantha, B.P., & Gowda, A.N.S. (2002) Foraging behaviour of honey
bees 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, 6770.
Jain, P.C. & Kushwaha, K.S. (1972) Morphometrics of bees, Apis dorsata F., A. indica F., A. florea F., Megachile lanata
F., Trigona smithii B. (Hymenoptera: Apidae) and their foraging habits at Udaipur. University of Udaipur Research
Journal, [5], 3840 [reference not seen].
Jander, R. & Jander, U. (1970) ber die Phylogenie der Geotaxis innerhalb der Bienen (Apoidea). Zeitschrift fr vergle-
ichende Physiologie, 66, 355368.
Jarau, S., Hrncir, M., Zucchi, R., & Barth, F.G. (2000) Recruitment behavior in stingless bees, Melipona scutellaris and
M. quadrifasciata. I. Foraging at food sources differing in direction and distance. Apidologie, 31, 8191.
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, 3943.
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 of
IUSSI, 1990 (International union for the study of social insects), Oxford, New Delhi, pp. 585586.
Jongjitvimol, T., Boontawon, K., Wattanachaiyingcharoen, W., & Deowanish, S. (2005) Nest dispersion of a stingless
bee species, Trigona collina Smith, 1857 (Apidae, Meliponinae) in a mixed deciduous forest in Thailand. Natural
History Journal of Chulalongkorn University, 5, 6971.
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. Natural
History Journal of Chulalongkorn University, 6, 7582.
Jongjitvimol, T. & Wattanachaiyingcharoen, W. (2007) Distribution, nesting sites and nest structures of the stingless bee
species, Trigona collina Smith, 1857 (Apidae, Meliponinae) in Thailand. Natural History Journal of Chulalongkorn
University, 7, 2534.
Jongjitvimol, T., Wattanachaiyingcharoen, W., Deowanish, S., & Wattanachaiyingcharoen, D. (2006) [Original title in
Thai=Factors affecting foraging behaviors and food resource partitioning of three Trigona species (Trigona apicalis
Smith, T. collina Smith and T. fimbriata Smith) in the Phitsanulok area]. BRT (The Thai Response to Biodiversity),
2549, 246252.
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, 9098.
Kajobe, R. & Echazarreta, C.M. (2005) Temporal resource partitioning and climatological influences on colony flight
and foraging of stingless bees (Apidae; Meliponini) in Ugandan Tropical Forests. African Journal of Ecology, 43,
267275.
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 Population
Ecology (Kyoto), 35, 95111.
Kerr, W.E. (1960) Evolution of communication in bees and its role in speciation. Evolution, 14, 386387.
Kerr, W.E. (1969) Some aspects of the evolution of social bees (Apidae). Evolutionary Biology, 3, 119175.
Kerr, W.E., Carvalho, G.A., & Nascimento, V.A. (1996) Abelha Uruu biologia manejo e conservao. Fundao
Acanga, Belo horizonte, 144 pp.
Kerr, W.E. & Cunha, R.A.d. (1976) Taxonomic position of two social bees (Apidae). Revista de Biologa Tropical, 24,
3543.
Kerr, W.E. & Esch, H. (1965) Comunicao entre as abelhas sociais brasileiras e sua contribuio para o entendimento da
sua evoluo. Cincia e Cultura, 17, 529538.
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, 8090.
Kerr, W.E. & Maule, V. (1964) Geographic distribution of stingless bees and its implications (Hymenoptera: Apidae).
Journal of the New York Entomological Society, 72, 218.
Kerr, W.E., Sakagami, S.F., Zucchi, R., Portugal-Arajo, V.d., & Camargo, J.M.F. (1967) Observaes sbre a
arquitetura dos ninhos e comportamento de algumas espcies de abelhas sem ferro das vizinhaas de Manaus,
Amazonas (Hymenoptera, Apoidea). Atas do Simpsio Sbre a Biota Amaznica, Zoologia, 5, 255309.
Kiew, B.H. (1985) Effective and economical lure for catching sweat bees. Malayan Naturalist, 38, 30.

TERMS OF USE
Kistner, D.H. (1982). The social insects' bestiary. In: H.R. Hermann (Ed.), Social insects. Vol. 3, Academic Press, New
York, pp. 1244.
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, 17.
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, 10031014.
Klein, A.-M., Steffan-Dewenter, I., & Tscharntke, T. (2003a) Pollination of Coffea canephora in relation to local and
regional agroforestry management. Journal of Applied Ecology, 40, 837845.
Klein, A.-M., Steffan-Dewenter, I., & Tscharntke, T. (2003b) Fruit set of highland coffee increases with the diversity of
pollinating bees. Proceedings of the Royal Society of London, Series B, 270, 955961.
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) and
their relationships to other Orchidaceae. International Journal of Plant Sciences, 162, 847867.
Koedam, D., Jungnickel, H., Tentschert, J., Jones, G.R., & Morgan, E.D. (2002) Production of wax by virgin queens of
the stingless bee Melipona bicolor (Apidae, Meliponinae). Insectes Sociaux, 49, 229233.
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, 95109.
Koshy, K.C., Harikumar, D., & Narendran, T.C. (2001) Insect visits to some bamboos of the Western Ghats, India. Cur-
rent Science (India), 81, 833838.
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, 1014.
Krombein, K.V. (1950) The aculeata Hymenoptera of Micronesia, II. Colletidae, Halictidae, Megachilidae, and Apidae.
Proceedings of the Hawaiian Entomological Society, 14, 101142.
Krombein, K.V. (1951) Additional notes on the bees of the Solomon islands (Hymenoptera: Apoidea). Proceedings of the
Hawaiian Entomological Society, 14, 277295.
Kshirsagar, K.K. & Chauhan, R.M. (1977) Preliminary observations on oviposition behaviour in Trigona (Tetragona) iri-
dipennis Smith. Indian Bee Journal, 39, 2223.
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, 5963.
Kumar, S. & Kumar, P. (1989) A study of the pollen carrying capacity of honey bees found in Lucknow in relation to
their body weight. Bulletin of Entomology (Madras), 30, 172177.
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, 165169.
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, 116230.
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, 3178.
Leonhardt, S.D., Dworschak, K., Eltz, T., & Blthgen, N. (2007) Foraging loads of stingless bees and utilisation of stored
nectar for pollen harvesting. Apidologie, 38, 125135.
Lepeletier de Saint Fargeau, A.L.M. (1836). Histoire naturelle des insectes. Hymnoptres. In. Vol. 1, Librairie encyclo-
pdique de Roret, Paris, pp. 1547.
Lindauer, M. (1956) ber die Verstndigung bei indischen Bienen. Zeitschrift fr vergleichende Physiologie, 38, 521
557.
Lindauer, M. & Kerr, W.E. (1958) Die gegenseitige Verstndigung bei den stachellosen Bienen. Zeitschrift fr vergle-
ichende Physiologie, 41, 405434.
Lindauer, M. & Kerr, W.E. (1960) Communication between the workers of stingless bees. Bee World, 41, 2941, 6571.
Liow, L.H. (1999) Bee diversity along a gradient of disturbance in tropical lowland forests of Southeast Asia. CBS:s
Skriftserie (Sweden), 3, 101130.
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, 180192.
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, 2526.
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, 318328.
McKenzie, E. (1975) Growing up with Aborigines. The Queensland Naturalist, 21, 4651.
Michener, C.D. (1961) Observations on the nests and behavior of Trigona in Australia and New Guinea (Hymenoptera,
Apidae). American Museum Novitates, 2026, 146.
Michener, C.D. (1964) Evolution of the nests of bees. American Zoologist, 4, 227239.

TERMS OF USE
Michener, C.D. (1965) A classification of the bees of the Australian and south Pacific regions. Bulletin of the American
Museum of Natural History, 130, 1362.
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, 75163.
Michener, C.D. (1997) Genus-group names of bees and supplemental family-group names. Scientific Papers, Natural
History Museum, University of Kansas, 181.
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, 231236.
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, 130135.
Michener, C.D. & Boongird, S. (2004) A new species of Trigona from Peninsular Thailand (Hymenoptera: Apidae:
Meliponini). Journal of the Kansas Entomological Society, 77, 143146.
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. 251266.
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, 1525.
Mohan, R. & Devanesan, S. (2000) Studies on the behaviour of the stingless bee, Trigona iridipennis Smith (Apidae:
Meliponinae). Indian Bee Journal, 62, 5962.
Mohana Rao, G. (1988("1987")) Scented methi a promising bee forage crop. Indian Bee Journal, 49, 1415.
Mohana Rao, G. & Suryanarayana, M.C. (1989("1988")) Studies on pollination of watermelon (Citrullus lanatus
(Thunb.) Mansf.). Indian Bee Journal, 50, 58.
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, 3133.
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 the
plant-pollinator community in a lowland dipterocarp forest. American Journal of Botany, 85, 14771501.
Moure, J.S. (1961) A preliminary supra-specific classification of the old world meliponine bees (Hymenoptera,
Apoidea). Studia Entomologica, 4, 181242.
Moure, J.S. & Camargo, J.M.F. (1978) A fossil stingless bee from copal (Hymenoptera: Apidae). Journal of the Kansas
Entomological Society, 51, 560566.
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, 481493.
Nagamitsu, T. & Inoue, T. (1997) Aggressive foraging of social bees a mechanism of floral resource partitioning in an
Asian tropical rainforest. Oecologia, 110, 432439.
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 to
general flowering in Sarawak, Malaysia. Apidologie, 33, 303314.
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. 7388.
Nagamitsu, T., Momose, K., Inoue, T., & Roubik, D.W. (1999) Preference in flower visits and partitioning in pollen diets
of stingless bees in an Asian tropical rain forest. Researches on Population Ecology (Kyoto), 41, 195202.
Nieh, J.C. (2004) Recruitment communication in stingless bees (Hymenoptera, Apidae, Meliponini). Apidologie, 35,
159182.
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, 238246.
Nogueira-Neto, P. (1949) India and stingless bees. Indian Bee Journal, 11, 121122.
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, 419451.
Nogueira-Neto, P. (1970) A criao de abelhas indgenas sem ferro, 2. ed. Tecnapis, So Paulo, 365.
Nogueira-Neto, P. (1997) Vida e criao de abelhas indgenas sem ferro. Edio Nogueirapis, So Paulo, 446.

TERMS OF USE
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, 173181.
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, 37.
Oliveira, M.L., Morato, E.F., & Garcia, M.V.B. (1995) Diversidade de espcies e densidade de ninhos de abelhas sociais
sem ferro (Hymenoptera, Apidae, Meliponinae) em floresta de terra firme na Amazonia Central. Revista Brasileira
de Zoologia, 12, 1324.
Ornduff, R. (1991) Size classes, reproductive behavior, and insect associates of Cycas media (Cycadaceae) in Australia.
Botanical Gazette, 152, 203207.
Oshy, K.C.K. & Arikumar, D.H. (2000) Flowering incidences and breeding system in Bambusa vulgaris. Current Sci-
ence (India), 79, 16501652.
Packard, A.S. (1867) The home of the bees. American Naturalist, 1, 364378.
Pagden, H.T. (1957) Notes of communal flight activites of stingless bees. Malayan Nature Journal, 12, 1519.
Palmer, K.A., Oldroyd, B.P., Quezada-Eun, 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, 21072113.
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, 1315.
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, 4243.
Parish, C.S.P. (1866) Pwai-ngyet. Science-Gossip, 2, 198200.
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 gnero Partamona Schwarz, 1939 (Hymenoptera,
Apidae). Revista Brasileira de Entomologia, 47, 1117.
Peeters, C. & Ito, F. (2001) Colony dispersal and the evolution of queen morphology in social Hymenoptera. Annual
Review of Entomology, 46, 601630.
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,
379384.
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, 38.
Pick, R.A. & Blochtein, B. (2002) Atividades de vo de Plebeia saiqui (Holmberg) (Hymenoptera, Apidae, Meliponini)
durante o perodo de postura da rainha e em diapausa. Revista Brasileira de Zoologia, 19, 827839.
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, 423430.
Portugal-Arajo, V.d. & Kerr, W.E. (1959) A case of sibling species among social bees. Revista Brasileira de Biologia,
19, 224228.
Prasad, J.N. & Hemanth, J. (1992) Pond heron Ardeola grayii (Sykes) feeding on bees. Journal of the Bombay Natural
History Society, 89, 246.
Prell, H. (1919) Menschenschdel als Bienenwohnung (Hym.). Entomologische Mitteilungen, 8, 157162.
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 "Agriculture
in 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 Phylogenetics
and Evolution, 43, 543552.
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, 103108.
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, 23.
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, 960967.
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, 939943.
Raju, A.J.S. & Reddi, C.S. (1989) Pollination biology of Anisomeles indica and A. malabarica (Lamiaceae). Plant Spe-
cies Biology, 4, 157167.

TERMS OF USE
Raju, A.J.S., Zafar, R., & Rao, S.P. (2005) Floral device for obligate selfing by remote insect activity and anemochory in
Wrightia tinctoria (Roxb.) R.Br. (Apocynaceae). Current Science (India), 88, 13781380.
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, 2528.
Ranger, S. & O'Donnell (1999) Genotypic effects on forager behavior in the neotropical stingless bee Partamona bilin-
eata (Hymenoptera: Meliponidae). Naturwissenschaften, 86, 187190.
Rao, S.P., Atluri, J.B., & Subba Reddi, C. (1999("1997")) Crataeva magna (Lour.) D.C. as a pollen source for the honey
bees. Indian Bee Journal, 59, 8587.
Rasmussen, C. (2004) A stingless bee nesting with a paper wasp (Hymenoptera: Apidae, Vespidae). Journal of the Kan-
sas Entomological Society, 77, 593601.
Rasmussen, C. (2008). Molecular phylogeny of stingless bees: Insights into divergence times, biogeography, and nest
architecture 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 (18601948): Names proposed and notes on a pioneer melittologist
(Hymenoptera, Anthophila). Zootaxa, 1833, 1118.
Rasmussen, C. & Camargo, J.M.F. (2008) A molecular phylogeny and the evolution of nest architecture and behavior in
Trigona s.s. (Hymenoptera: Apidae: Meliponini). Apidologie, 39, 102118.
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, 2639.
Rayment, P.T. (1930) New and remarkable bees. Proceedings of the Royal Society of Victoria, 43, 4261.
Rayment, P.T. (1932a) The stingless bees of Australia. 1. Introduction. Victorian Naturalist, 48, 183189.
Rayment, P.T. (1932b) The stingless bees of Australia. 2. The architecture. Victorian Naturalist, 48, 203212.
Rayment, P.T. (1932c) The stingless bees of Australia. 3. Behaviour of the individual. Victorian Naturalist, 48, 246254.
Rayment, P.T. (1932d) The stingless bees of Australia. 4. Food of the larval bee. Victorian Naturalist, 49, 914.
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, 104107.
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, 263294.
Rayment, P.T. (1953) The distribution of the bee genus, Trigona. Western Australian Naturalist, 4, 2223.
Rayment, P.T. (1961) A tiny bee and a very large flower. Proceedings of the Royal Zoological Society of New South
Wales, 19581959, 9697.
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, 423425.
Rocha, M.P., Pompolo, S.d.G., & Campos, L.A.O. (2003). Citogentica 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, Cricima, pp. 311320.
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, 115120.
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 equatorial
Sumatra, Hokkaido University Press, Sapporo, pp. 245257.
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, Academic
Press, San Diego, pp. 327354.
Roubik, D.W. (1993) Direct costs of forest reproduction, bee-cycling and the efficiency of pollination modes. Journal of
Bioscience, 18, 537552.
Roubik, D.W., ed. (1995) Pollination of cultivated plants in the tropics. Vol. 118, pp x+1196. 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. 5966.
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. 89103. Springer.

TERMS OF USE
Roubik, D.W. (2006) Stingless bee nesting biology. Apidologie, 37, 124143.
Sakagami, S.F. (1959) Stingless bees collected by Prof. S. Matsumura from Singapore (Hymenoptera, Apidae). Insecta
Matsumurana, 22, 119121.
Sakagami, S.F. (1960) Some bees of Apinae and Xylocopinae collected in Cambodia. Konty, 28, 146147.
Sakagami, S.F. (1963) On the male of Trigona (Tetragona) fimbriata Smith (Hymenoptera: Apidae). Journal of the New
York Entomological Society, 71, 3944.
Sakagami, S.F. (1971) Ethosoziologischer Vergleich zwischen Honigbienen und stachellosen Bienen. Zeitschrift fr
Tierpsychologie, 28, 337350.
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, 4976.
Sakagami, S.F. (1978) Tetragonula stingless bees of the continental Asia and Sri Lanka (Hymenoptera, Apidae). Journal
of the Faculty of Science, Hokkaido University, Series VI, Zoology, 21, 165247.
Sakagami, S.F. (1982). Stingless bees. In: H.R. Hermann (Ed.), Social insects. Vol. 3, Academic Press, New York, pp.
361423.
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, 164175.
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, 163189.
Sakagami, S.F. & Inoue, T. (1985) Taxonomic notes on three bicolorous Tetragonula stingless bees in Southeast Asia.
Konty, 53, 174189.
Sakagami, S.F. & Inoue, T. (1987) Stinglee bees of the genus Trigona (subgenus Trigonella) with notes on the reduction
of spatha in male genitalia of the subgenus Tetragonula (Hymenoptera, Apidae). Konty, 55, 610627.
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, 605620.
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 social
wasps and bees in equatorial Sumatra, Hokkaido University Press, Sapporo, pp. 201217.
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 in
central Sumatra, Kanazawa University, Kanazawa (Japan), pp. 3743.
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. 125137.
Sakagami, S.F., Inoue, T., Yamane, S., & Salmah, S. (1983a) Nest architecture and colony composition of the Sumatran
stingless bee Trigona (Tetragonula) laeviceps. Konty, 51, 100111.
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, 265274.
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, 365376.
Sakagami, S.F. & Khoo, S.G. (1987) Taxonomic status of the Malesian stingless bee Trigona reepeni, with discovery of
Trigona pagdeni from northern Malaya. Konty, 55, 207214.
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,
3748.
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, 1727.
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, 121.
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 Central
Sumatra with special reference to wasps and bees, pp. 4651.
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]. Konty, 51, 441457.
Sakagami, S.F. & Yoshikawa, K. (1961) Bees of Xylocopinae and Apinae collected by the Osaka city University biolog-
ical expedition to Southeast Asia 195758, 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 postemergence pigmentation in the

TERMS OF USE
sumatran stingless bee, Trigona (Trigonella) moorei. Konty, 55, 383390.

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, 5558.
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. 139174.
Salmah, S., Inoue, T., & Sakagami, S.F. (1991) Incubation period in the sumatran stingless bee Trigona (Tetragonula)
minangkabau. Treubia, 30, 195201.
Salmah, S., Inoue, T., & Sakagami, S.F. (1996) Incubation period and post-emergence pigmentation in the Sumatran
stingless bee Trigona (Heterotrigona) itama (Apidae, Meliponinae). Japanese Journal of Entomology, 64, 401411.
Salt, G. (1929) A contribution to the ethology of the Meliponinae. Transactions of the Entomological Society of London,
77, 431470.
Samejima, H., Marzuki, M., Nagamitsu, T., & Nakasizuka, T. (2004) The effects of human disturbance on a stingless bee
community in a tropical rainforest. Biological Conservation, 120, 577587.
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, 519619.
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 Strung des Bienengedchtnisses durch Narkosemittel, zugleich ein Beitrag zur Strung
der sozialen Bindung durch Narkose. Zeitschrift fr vergleichende Physiologie, 47, 559595.
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, 237248.
Schneider, E.L. (1982) Observations on the pollination biology of Nymphaea gigantea W. J. Hooker (Nymphaeaceae).
Western Australian Naturalist, 15, 7172.
Schulz, W.A. (1907) Die indoaustralische Trigona laeviceps F. Sm. und ihr Nest. Zeitschrift fr wissenschaftliche Insek-
tenbiologie, 3, 6573.
Schulz, W.A. (1909) Ein javanisches Nest von Trigona canifrons F. Sm. in einem Bambusstabe. Zeitschrift fr wissen-
schaftliche Insektenbiologie, 5, 338341.
Schwarz, H.F. (1932) The genus Melipona. The type genus of the Meliponidae or stingless bees. Bulletin of the American
Museum of Natural History, 63, 231460.
Schwarz, H.F. (1937) Results of the Oxford University Sarawak (Borneo) expedition: Bornean stingless bees of the
genus Trigona. Bulletin of the American Museum of Natural History, 73, 281328.
Schwarz, H.F. (1939a) The Indo-Malayan species of Trigona. Bulletin of the American Museum of Natural History, 76,
83141.
Schwarz, H.F. (1939b) Two Trigona bees collected by Prof. Teiso Esaki on the Palau and East Caroline Islands. Mushi,
12, 151152.
Schwarz, H.F. (1945) The wax of stingless bees (Meliponidae) and the uses to which it has been put. Journal of the New
York Entomological Society, 53, 137144.
Schwarz, H.F. (1948) Stingless bees (Meliponidae) of the western hemisphere. Lestrimelitta and the following subgenera
of Trigona: Trigona, Paratrigona, Schwarziana, Parapartamona, Cephalotrigona, Oxytrigona, Scaura, and
Mourella. Bulletin of the American Museum of Natural History, 90, 1546.
Schwarz, H.F. (1951) Stingless bees collected by R. Takahashi in Indo-China and Siam (Hymenoptera). Mushi, 22, 63.
Serro, J.E. (2001) A comparative study of the proventricular structure in corbiculate Apinae (Hymenoptera, Apidae).
Micron, 32, 379385.
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: practice
and perspectives. Apidologie, 37, 293315.
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 Biologa,
Universidad Nacional Autnoma de Mxico. Serie Zoologa, 67, 197203.
Smith, F. (1854) Catalogue of the Hymenopterous Insects in the Collection of the British Museum. Part II, Apid. Brit-
ish Museum (Natural History), London, 197.
Smith, F. (1857) Catalogue of the hymenopterous insects collected at Sarawak, Borneo; Mount Ophir, Malacca; and at
Singapore, by A. R. Wallace. Journal of the Proceedings of the Linnean Society, Zoology, 2, 4288, pl. 12.
Smith, F. (1858) Mr. Smith exhibited some specimens of a small species of Trigona from Moulmein... Transactions of

TERMS OF USE
the Entomological Society of London, 9, 9899.

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, 132178.
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, 181182.
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, 6194, plate 4.
Smith, F. (1871) A catalogue of the Aculeate Hymenoptera and Ichneumonidae of India and the eastern archipelago. By
F. Smith, with introductory remarks by A. R. Wallace. Journal of the Proceedings of the Linnean Society, Zoology,
11, 285415.
Smith, F. (1878) List of Hymenoptera obtained by Mr. Ossian Limborg east of Maulmain, Tenasserim provinces, during
the months december 1876, january, march and april 1877, with descriptions of new species. Journal of the Asiatic
Society of Bengal (Physical Science), 47, 167169.
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; Natural
history 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. 3947, 131139, 140143.
Spinola, M. (1853) Compte rendu des Hymnoptres indits provenant du voyagae entomologique de M. Ghiliani dans
le Para en 1846. Memorie della Accademia delle Scienze di Torino, [2]13, 1994.
Starr, C.K. & Sakagami, S.F. (1987) An extraordinary concentration of stingless bee colonies in the Philippines, with
notes on nest structure (Hymenoptera: Apidae: Trigona spp.). Insectes Sociaux, 34, 96107.
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, 422424.
Strand, E. (1911) Zur Kenntnis papuanischer und australischer Hymenopteren, insbesondere Schlupfwespen. Internatio-
nale Entomologische Zeitschrift, 5, 8687.
Strand, E. (1912) Biologische Notiz ber papuanische Trigonen. Internationale Entomologische Zeitschrift, 6, 11.
Strand, E. (1913a) Apidae con Ceylon, gesammelt 1899 von Herrn Dr. W. Horn. Archiv fr Naturgeschichte. Abteilung
A, 79, 135150.
Strand, E. (1913b) [H. Sauter's Formosa-Ausbeute] Apidae I (Hym.). Supplementa Entomologica, 2, 2367.
Strand, E. (1915) Beitrge zur Systematik und insbesondere zur Verbreitung der Apidae. Archiv fr Naturgeschichte.
Abteilung A, 81(11), 124139.
Suka, T. & Inoue, T. (1993) Nestmate recognition of the stingless bee Trigona (Tetragonula) minangkabau (Apidae:
Meliponinae). Journal of Ethology, 11, 141147.
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, 129147.
Sung, I.-H., Lin, M.-Y., Chang, C.-H., Cheng, A.-S., & Chen, W.-S. (2006b) Pollinators and their behaviors on mango
flowers in southern Taiwan. Formosan Entomologist, 26, 161170.
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 honeybee
Apis cerana in Taiwan (Hymenoptera: Apidae)]. Japanese Journal of Entomology, N.S., 9, 3345.
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, 263269.
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, 229246.
Tangmitcharoen, S., Pianhanuruk, P., & Cunningham, R. (2002) [Original title in Thai=Pollination ecology in relation to
fruit setting of teak (Tectona grandis Linn.f.): natural and plantation forests]. In Proceedings of the Forestry Confer-
ence, pp. 202214.
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, 6774.

TERMS OF USE
Temaru, E., Shimura, S., Amano, K., & Karasawa, T. (2007) Antibacterial activity of honey from stingless honeybees
(Hymenoptera; Apidae; Meliponinae). Polish Journal of Microbiology, 56, 281285.
Tennent, J.E. (1859) Ceylon, an account of the island physical, historical, and topographical with notices of its natural
history, 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 and
instincts 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, 759762.
Thomson, V.P., Nicotra, A.B., & Cunningham, S.A. (2004) Herbivory differentially affects male and female reproductive
traits of Cucumis sativus. Plant Biology, 6, 621628.
Thummajitsakul, S., Klinbunga, S., Smith, D., & Sittipraneed, S. (2008) Genetic diversity and population structure of
Trigona pagdeni Schwarz in Thailand. Apidologie, 39, 446455.
Tth, E., Queller, D.C., Dollin, A.E., & Strassmann, J.E. (2004) Conflict over male parentage in stingless bees. Insectes
Sociaux, 51, 111.
Tth, E., Strassmann, J.E., Imperatriz-Fonseca, V.L., & Queller, D.C. (2003) Queens, not workers, produce the males in
the stingless bee Schwarziana quadripunctata quadripunctata. Animal Behaviour, 66, 359368.
Tsuji, K., Hasyim, A., Nakamura, H., & Nakamura, K. (2004) Asian weaver ants, Oecophylla smaragdina, and their
repelling of pollinators. Ecological Research, 19, 669673.
Uji, T. (1987) Penyerbukan pada rambutan (Nephelium lappaceum var. lappaceum) [=Pollination in Nephelium lappa-
ceum var. lappaceum]. Berita Biologi, 32, 3134.
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 on
the pinang yaki (Areca vestiaria Giseke) in the Bogor Botanical Gardens]. Berita Biologi, 3, 470472.
van Veen, J.W. & Sommeijer, M.J. (2000) Colony reproduction in Tetragonisca angustula (Apidae, Meliponini). Insectes
Sociaux, 47, 7075.
Velthuis, H.H.W., Koedam, D., & Imperatriz-Fonseca, V.L. (2005) The males of Melipona and other stingless bees, and
their mothers. Apidologie, 36, 169185.
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, 18771879, beschreven door de leden der expeditie, onder toezicht van Prof. P.J. Veth. Vol. 4 (Natuurlijke his-
torie, fauna), E.J. Brill, Leiden, pp. 123.
Viraktamath, S., Lingappa, S., Kallesha, G.R., Holi, K.S., & Hunur, S. (1999) Comparative studies on pollen sources to
Trigona sp. Apis mellifera L. and Apis cerana F. in Dharwad, Karnataka, India. Indian Bee Journal, 61, 7173.
Wagner, A.E. & Briscoe, D.A. (1983) An absence of enzyme variability within two species of Trigona (Hymenoptera).
Heredity, 50, 97103.
Wagner, A.E. & Dollin, L.J. (1982a) Probing the mysteries of the N.S.W. native bees. The Australasian Beekeeper, 83,
157161.
Wagner, A.E. & Dollin, L.J. (1982b) Swarming in Australian native bees help solve the mystery! The Australasian
Beekeeper, 84, 3438.
Wagner, A.E. & Dollin, L.J. (1982c) North Queensland's native bees the little Aussie battlers. The Australasian Bee-
keeper, 84, 7072.
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, 304311.
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, 8794.
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, 5559.
Wallace, H.M. & Trueman, S.J. (1995) Dispersal of Eucalyptus torelliana seeds by the resin-collecting stingless bee,
Trigona carbonaria. Oecologia, 104, 1216.
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, 133136, 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, 7174.

TERMS OF USE
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 conservation
area, Lopburee province]. NU (Naresuan University) Science Journal, 1, 6374.
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, 2328.
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.), The
naturalist's library. Vol. 26, W.H. Lizars, Edinburgh, pp. viii+17301, 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, 6164.
Wijesekara, A. (2001) An annotated list of bees (Hymenoptera: Apoidea: Apiformis) of Sri Lanka. Tijdschrift voor Ento-
mologie, 144, 145158.
Wille, A. (1961) Evolutionary trends in the ventral nerve cord of the stingless bees (Meliponini). Revista de Biologa
Tropical, 9, 117129.
Wille, A. (1962) A new species of stingless bee (Meliponini) from Costa Rica. Revista de Biologa Tropical, 10, 177
181.
Wille, A. (1977) A general review of the fossil stingless bees. Revista de Biologa Tropical, 25, 4346.
Wille, A. (1979a) A comparative study of the pollen press and nearby structures in the bees of the family Apidae. Revista
de Biologa Tropical, 27, 217222.
Wille, A. (1979b) Phylogeny and relationships among the genera and subgenera of the stingless bees (Meliponinae) of
the world. Revista de Biologa Tropical, 27, 241277.
Wille, A. (1983) Biology of the stingless bees. Annual Review of Entomology, 28, 4164.
Wille, A. & Michener, C.D. (1973) The nest architecture of stingless bees with special reference to those of Costa Rica.
Revista de Biologa Tropical, 21(supl.), 9278.
Williams, F.X. (1928) The natural history of a Philippine nipa house with descriptions of new wasps. Philippine Journal
of Science, 35, 53118.
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,
141146.
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, 6995.
Wittmann, D., Bego, L.R., Zucchi, R., & Sakagami, S.F. (1991) Oviposition behavior and related aspects of the stingless
bees. XIV. Plebeia (Mourella) caerulea, with comparative notes on the evolution of the oviposition patterns (Api-
dae, Meliponinae). Japanese Journal of Ecology, 59, 793809.
Wu, Y. (2000). [Original title in Chinese=Hymenoptera Melittidae Apidae]. In: Fauna Sinica, Insecta. Vol. 20, Science
Press, Beijing, pp. 1442, pls. 19.
Yamane, S. (1996) [Original title in Japanese=Social wasps in Australia]. Nature and Insects, 31, 1015.
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, 275296.
Yasumatsu, K. (1935) Notes on two Apoidea collected by Prof. Tohru Uchida on the Mariana and West Caroline islands.
Mushi, 8, 9496.
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 Molecular
Biology, 99, 625628.
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 of
the first International conference on the Asian honey bees and bee mites, Wicwas Press, Cheshire, Connecticut, pp.
5156.
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, 465467.
Yoshikawa, K., Ohgushi, R., & Sakagami, S.F. (1969) Preliminary report on entomology of the Osaka city University 5th

TERMS OF USE
scientific expedition to Southeast Asia 1966. With descriptions of two new genera of stenogastrine waps by J. van
der Vecht. Nature and Life in Southeast Asia, 6, 153182.
Yumoto, T. (2000) Bird-pollination of three Durio species (Bombacaceae) in a tropical rainforest in Sarawak, Malaysia.
American Journal of Botany, 87, 11811188.
Zeuner, F.E. & Manning, F.J. (1976) A monograph on fossil bees (Hymenoptera: Apoidea), edited with an appendix by
Samuel Francis Morris. Bulletin of the British Musem (Natural History), series Geology, 27, 149268.
Zherikhin, V.V. & Ross, A.J. (2000) A review of the history, geology and age of Burmese amber (Burmite). Bulletin of
the Natural History Museum (Geology), 56, 310.
Zoological Society of London, ed. (1864present) The Zoological Record [former Record of Zoological Literature]. J.V.
Vorst, London.

Rasmussen Catalog Kelulut Indo Malayan PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Rasmussen Catalog Kelulut Indo Malayan PDF

Uploaded by

Copyright:

Available Formats

TERMS OF USE

This pdf is provided by Magnolia Press for private/research use.

Catalog of the Indo-Malayan/Australasian stingless bees

FIRST PUBLISHED IN 2008 BY

2008 Magnolia Press

ISSN 1175-5326 (Print edition)

2 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Zootaxa 1935: 180 (2008) ISSN 1175-5326 (print edition)

Catalog of the Indo-Malayan/Australasian stingless bees

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

Key words: Austroplebeia, Geniotrigona, Heterotrigona, Homotrigona, Lepidotrigona, Lisotrigona, Lophotrigona,

4 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Taxonomy and classification

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

Distribution and biology

6 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

TABLE 1. Different classificatory schemes used in the literature.

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 7

Catalog of the indo-malayan/australasian stingless bees

Austroplebeia australis (Friese, 1898)

Austroplebeia cassiae (Cockerell, 1910)

8 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Austroplebeia cincta (Mocsry in Friese, 1898)

Austroplebeia cockerelli (Rayment, 1930)

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 9

Austroplebeia essingtoni (Cockerell, 1905)

Austroplebeia ornata (Rayment, 1932)

Austroplebeia percinta (Cockerell, 1929)

Austroplebeia symei (Rayment, 1932)

10 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Austroplebeia websteri (Rayment, 1932)

Geniotrigona incisa (Sakagami & Inoue, 1989)

Geniotrigona lacteifasciata (Cameron, 1902)

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 11

Geniotrigona thoracica (Smith, 1857)

12 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Geniotrigona indet. Roubik 2006: 130 (nest).

Heterotrigona bakeri (Cockerell, 1919)

Heterotrigona erythrogastra (Cameron, 1902)

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 13

Heterotrigona itama (Cockerell, 1918)

14 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Heterotrigona indet. Roubik 2006: 127, 132 (nest).

Homotrigona aliceae (Cockerell, 1929)

Homotrigona anamitica (Friese, 1909)

Homotrigona fimbriata (Smith, 1857)

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 15

16 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Homotrigona lutea (Bingham, 1897)

Homotrigona indet. Roubik 2006: 130 (nest).

Lepidotrigona arcifera (Cockerell, 1929)

Lepidotrigona doipaensis (Schwarz, 1939)

Lepidotrigona flavibasis (Cockerell, 1929)

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 17

Lepidotrigona hoozana (Strand, 1913), comb. nov.

Lepidotrigona javanica (Gribodo, 1891)

Lepidotrigona latebalteata (Cameron, 1902)

Lepidotrigona latipes (Friese, 1900)

18 Zootaxa 1935 2008 Magnolia Press RASMUSSEN

Lepidotrigona nitidiventris (Smith, 1857)

Lepidotrigona palavanica (Cockerell, 1915)

Lepidotrigona terminata (Smith, 1878)

INDO-MALAYAN/AUSTRALASIAN STINGLESS BEES Zootaxa 1935 2008 Magnolia Press 19