Unveiling cryptic species of the bumblebee subgenus Bombus s. str. worldwide with COI barcodes (Hymenoptera: Apidae)

Systematics and Biodiversity

Volumn 10, Issue 1, 2012, Pages 21-56

  Williams, Paul H ^a   Brown, Mark J F ^b   Carolan, James C ^c   An, Jiandong ^d   Goulson, Dave ^e   Aytekin, A Murat ^f   Best, Lincoln R ^g   Byvaltsev, Alexandr M ^h   Cederberg, Björn ⁱ   Dawson, Robert ^e   Huang, Jiaxing ^d   Ito, Masao ^j   Monfared, Alireza ^k   Raina, Rifat H ^l   Schmid Hempel, Paul ^m   Sheffield, Cory S ^g   Šima, Peter ⁿ   Xie, Zenghua ^o  

^a DEPARTMENT OF LIFE SCIENCES   (United Kingdom)

^b UNIVERSITY OF LONDON   (United Kingdom)

^c NATIONAL UNIVERSITY OF IRELAND   (Ireland)

^d INSTITUTE OF PLANT PROTECTION   (China)

^e UNIVERSITY OF STIRLING   (United Kingdom)

^f HACETTEPE UNIVERSITY   (Turkey)

^g YORK UNIVERSITY   (Canada)

^h NOVOSIBIRSK STATE UNIVERSITY   (Russian Federation)

ⁱ SWEDISH UNIVERSITY OF AGRICULTURAL SCIENCES   (Sweden)

^j Yasouj University ^*  (Japan)

^k YASOUJ UNIVERSITY   (Iran)

^l SHER E KASHMIR UNIVERSITY OF AGRICULTURAL SCIENCES AND TECHNOLOGY OF KASHMIR   (India)

^m ETH ZURICH   (Switzerland)

ⁿ Koppert Sro   (Slovakia)

^o RESEARCH INSTITUTE OF FORESTRY   (China)

more..

Author keywords

Barcoding; Bombus; Bumblebees; Lucorum complex; Pollinators; Systematics; Taxonomy

Indexed keywords

APIDAE; BOMBUS CRYPTARUM; BOMBUS FRANKLINI; BOMBUS HYPOCRITA; BOMBUS IGNITUS; BOMBUS LUCORUM; BOMBUS MAGNUS; BOMBUS OCCIDENTALIS; BOMBUS PATAGIATUS; BOMBUS SPORADICUS; BOMBUS TERRESTRIS; BOMBUS TERRICOLA; HYMENOPTERA;

EID: 84863247111     PISSN: 14772000     EISSN: 14780933     Source Type: Journal    
DOI: 10.1080/14772000.2012.664574     Document Type: Review

References (172)

1. ALFORD, D.V. 1975. Bumblebees. Davis-Poynter, London.
2. AN, J.-D., HUANG, J.-X., WILLIAMS, P.H., WU, J. & ZHOU, B.-F. 2010. Species diversity and colony characteristics of bumblebees in the Hebei region of North China. Chinese Journal of Applied Ecology 21, 1542-1550.
3. AN, J.-D., YAO, J., HUANG, J.-X., SHAO, Y.-Q., WU, J., LI, J.-L. & SHI, H.-Y. 2008. Bombus fauna (Hymenoptera, Apidae) in Shanxi, China. Acta Zootaxonomica Sinica 33, 80-88.
4. ARRETZ, P.V.&MACFARLANE, R.P. 1982. The introduction of Bombus ruderatus to Chile for red clover pollination. Bee World 67, 15-22.
5. ASADA, S.&ONO,M. 1996. Crop pollination by Japanese bumblebees, Bombus spp. (Hymenoptera: Apidae): tomato foraging behaviour and pollination efficiency. Applied Entomology and Zoology 31, 581-586.
6. ASHMEAD, W.H. 1902. Papers from the Harriman Alaska Expedition. XXVIII. Hymenoptera. Proceedings of the Washington Academy of Sciences 4, 117-274.
7. BAKER, A.J., TAVARES, E.S. & ELBOURNE, R.F. 2009. Countering criticisms of single mitochondrial DNA gene barcoding in birds. Molecular Ecology Resources 9 (Supplement 1), 257-268.
8. BAKER, D.B. 1996. On a collection of Bombus and Psithyrus principally from Sutherland, with notes on the nomenclature or status of three species (Hymenoptera, Apoidea). British Journal of Entomology and Natural History 9, 7-19.
9. BALLARD, J.W.O. & WHITLOCK, M.C. 2004. The incomplete natural history of mitochondria. Molecular Ecology 13, 729-744. (Pubitemid 38390584)
10. BERGSTRöM, G., KULLENBERG, B. & STÄLLERG-STENHAGEN, S. 1973. Studies on natural odoriferous compounds. VII. Recognition of two forms of Bombus lucorum L. (Hymenoptera, Apidae) by analysis of the volatile marking secretion from individual males. Chemica Scripta 4, 174-182.
11. BERTSCH,A. 2009. Barcoding cryptic bumblebee taxa: B. lucorum, B. cryptarum and B. magnus, a case study (Hymenoptera: Apidae: Bombus). Beiträge zur Entomologie 59, 287-310.
12. BERTSCH, A. 2010. A phylogenetic framework for the bumblebee species of the subgenus Bombus sensu stricto based on mitochondrial DNA markers, with a short description of the neglected taxon B. minshanicola Bischoff, 1936 n. status. Beiträge zur Entomologie 60, 471-487.
13. BERTSCH, A. & SCHWEER, H. 2011. Labial gland marking secretions of male Bombus lucorum bumblebees from Europe and China reveal two separate species: B. lucorum (Linnaeus 1761) and Bombus minshanicola (Bischoff 1936). Biochemical Systematics and Ecology 2011, 1-7.
14. BERTSCH, A.,HABRÉ DE ANGELIS, M. & PRZEMECK, G.K.H. 2010a. Phylogenetic relationships of the bumblebees Bombus moderatus, B. albocinctus, B. burjaeticus, B. florilegus and B. cryptarum based on mitochondrial DNA markers: a complex of closely related taxa with circumpolar distribution. Beiträge zur Entomologie 60, 13-32.
15. BERTSCH, A.,HRABÉ DE ANGELIS,M. & PRZEMECK, G.K.H. 2010b. A phylogenetic framework for the North American bumblebee species of the subgenus Bombus sensu stricto (Bombus affinis, B. franklini, B. moderatus, B. occidentalis & B. terricola) based on mitochondrial DNA markers. Beiträge zur Entomologie 60, 229-242.
16. BERTSCH, A., SCHWEER, H. & TITZE, A. 2004. Discrimination of the bumblebee species Bombus lucorum, B. cryptarum and B. magnus by morphological characters and male labial gland secretions. Beiträge zur Entomologie 54, 365-386. (Pubitemid 40102136)
17. BERTSCH, A., SCHWEER, H., TITZE, A. & TANAKA, H. 2005. Male labial gland secretions and mitochondrial DNA markers support species status of Bombus cryptarum and B. magnus (Hymenoptera, Apidae). Insectes Sociaux 52, 45-54. (Pubitemid 40856338)
18. BISCHOFF, H. 1930. Entomologische Ergebnisse der schwedischen Kamtschatka Expedition 1920-1922. 29. Bombinae (Hymen.). Arkiv för Zoologi 21, 1-6.
19. BISCHOFF, H. 1936. Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sü Ping-chang. Insekten gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927-1930. 56. Hymenoptera. 10. Bombinae. Arkiv för Zoologi 27A, 1-27.
20. BROWER, A.V.Z. 2006. Problems with DNA barcodes for species delimitation: 'ten species' of Astraptes fulgerator reassessed (Lepidoptera: Hesperiidae). Systematics and Biodiversity 4, 127-132. (Pubitemid 43825010)
21. BROWN, M.J.F. 2011. The trouble with bumblebees. Nature 469, 169-170.
22. BUCHMANN, S., KEVAN, P.G., ADAMS, L. & DALSIMER, A. 2008. Birds, bees, and bases: how military natural resource managers can help local pollinators and why they should care. Natural Selections 4, 5-6.
23. CAMERON, S.A., HINES, H.M. & WILLIAMS, P.H. 2007. A comprehensive phylogeny of the bumble bees (Bombus). Biological Journal of the Linnean Society 91, 161-188. (Pubitemid 46642958)
24. CAMERON, S.A., LOZIER, J.D., STRANGE, J.P., KOCH, J.B., CORDES, N., SOLTER, L.F. & GRISWOLD, T.L. 2011. Patterns of widespread decline in North American bumble bees. Proceedings of the National Academy of Sciences USA 108, 662-667.
25. CAROLAN, J.C., MURRAY, T.E., FITZPATRICK, U., CROSSLEY, J., SCHMIDT, H., CEDERBERG, B., MCNALLY, L., PAXTON, R.J., WILLIAMS, P.H. & BROWN, M.J.F. 2012. Colour patterns do not diagnose species: quantitative evaluation of a DNA barcoded cryptic bumblebee complex. PLoS ONE 7, 662-667.
26. CHEN, W.-F., AN, J.-D., DONG, J., DING, K.-F. & GAO, S. 2011. Flower-visiting behavior and pollination ecology of different bee species on greenhouse strawberry. Chinese Journal of Ecology 30, 290-296.
27. COCKERELL, T.D.A. 1905. Descriptions and records of bees.-I. Annals and Magazine of Natural History 16 (7), 216-225.
28. COCKERELL, T.D.A. 1911. Bees in the collection of the U. S. National Museum. 1. Proceedings of the United States National Museum 39, 635-658.
29. COLLA, S.R. & PACKER, L. 2008. Evidence for decline in eastern North American bumblebees (Hymenoptera: Apidae), with special focus on Bombus affinis Cresson. Biodiversity and Conservation 17, 1379-1391.
30. CRESSON, E.T. 1863. List of the North American species of Bombus and Apathus. Proceedings of the Entomological Society of Philadelphia 2, 83-116.
31. CRESSON, E.T. 1916. The Cresson types ofHymenoptera. Memoirs of the American Entomological Society 1, 1-141.
32. DALLA TORRE, K.W.V. 1882. Bermerkungen zur Gattung Bombus Latr. II. 3. Zur Synonymie und geographischen Verbreitung der Gattung Bombus Latr. Bericht des Naturwissenschaftlichmedizinischen Vereins in Innsbruck 12, 14-31.
33. DAVYDOVA, N.G. 2001. On the taxonomic status of the bumble bee Bombus albocinctus (Hymenoptera, Apidae). Entomological Review 81, 1197-1201. (Pubitemid 36420897)
34. DAY, M.C. 1979. The species of Hymenoptera described by Linnaeus in the genera Sphex, Chrysis, Vespa, Apis and Mutilla. Biological Journal of the Linnean Society 12, 45-84.
35. DE MEULEMEESTER, T., GERBAUX, P., BOULVIN, M., COPPÉE, A. & RASMONT, P. 2011. A simplified protocol for bumble bee species identification by cephalic secretion analysis. Insectes Sociaux 58, 227-236.
36. DELMAS, R. 1976. Contribution à l'étude de la faune française des Bombinae (Hymenoptera, Apoidea, Bombidae). Annales de la Société entomologique de France (Nouvelle serie) 12, 247-290.
37. DELMAS, R. 1981. Systematics and geographical variation in the Bombinae. In: HOWSE, P.E. & CLEMENT, J.-L., Eds., Biosystematics of Social Insects, vol. 19. Systematics Association Special Volume. Academic Press, London and New York, pp. 223-229.
38. DIAS, B.S.F., RAW, A. & IMPERATRI-FONSECA, V.L. 1999. International Pollinators Initiative: the São Paulo declaration on pollinators. Report on the recommendations of the workshop on the conservation and sustainable use of pollinators in agriculture with emphasis on bees. Brazilian Ministry of the Environment, Brasília, pp. 79.
39. DRUMMOND, A.J. & RAMBAUT, A. 2007. BEAST: Bayesian evolutionary analysis by sampling trees. BioMed Central Evolutionary Biology 7, 214.
40. EBACH, M.C. & CARVALHO, M.R.D. 2010. Anti-intellectualism in the DNA barcoding enterprise. Zoologia 27, 165-178.
41. EDWARDS, S.V. & BEERLI, P. 2000. Gene divergence, population divergence, and the variance in coalescence time in phylogeographic studies. Evolution 54, 1839-1854. (Pubitemid 32113816)
42. ERLANDSSON, A. 1979. Bombus canariensis Pérez, 1895 n. stat and Bombus maderensis n. sp. from the Macaronesian Islands. Entomologica Scandinavica 10, 187-192.
43. ESTOUP, A., SOLIGNAC, M.,CORNUET, J.-M., GOUDET, J. & SCHOLL, A. 1996. Genetic differentiation of continental and island populations of Bombus terrestris (Hymenoptera: Apidae) in Europe. Molecular Ecology 5, 19-31.
44. EVANS, E.,THORP, R., JEPSON, S.&HOFFMAN BLACK, S. 2008. Status reviewof three formerly common species of bumble bee inthe subgenus Bombus ed. The Xerces Society for Invertebrate Conservation, Portland (Oregon), pp. 63.
45. EVERSMANN, E. 1852. Fauna hymenopterologica Volgo- Uralensis. (Continuatio). Familia anthophilarium seu apidarum. Izvêstiya Moskovskago éntomologicheskago obshchestva 25, 3-137.
46. FABRICIUS, J.C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Flensburgi and Lipsiae.
47. FRANKLIN, H.J. 1913. The Bombidae of the New World. Transactions of the American Entomological Society 38, 177-486.
48. FRIESE,H. 1916.Über einige neueHummelformen. Bombus 1916, 107-110.
49. FRIESE, H. 1918. Über Hummelformen aus dem Himalaja. Deutsche entomologische Zeitschrift 1918, 81-86.
50. FRISON, T.H. 1921. New distribution records for North American Bremidae, with description of a new species (Hym.). Entomological News 32, 144-148.
51. GERTH, M.,GEISSLER, A. & BLEIDORN, C. 2011. Wolbachia infection in bees (Anthophila) and possible implications for DNA barcoding. Systematics and Biodiversity 9, 319-327.
52. GOULSON, D. 2003. Effects of introduced bees on native ecosystems. Annual Review of Ecology, Evolution, and Systematics 34, 1-26. (Pubitemid 38171411)
53. GOULSON, D. 2010. Bumblebees, Behaviour, Ecology, and Conservation, 2nd edition. Oxford University Press, Oxford.
54. GREENE, J.W. 1858. II. Descriptions of several new Hymenopterous insects from the north west coast of America. Annals of the Lyceum of Natural History of New York 7, 11-12.
55. HARRIS, M. 1776. An exposition of English insects, with curious observations and remarks, wherein each insect is particularly described; its parts and properties considered; the different sexes distinguished, and the natural history faithfully related. London.
56. HEBERT, P.D.N., RATNASINGHAM, S. & DEWAARD, J.R. 2003. Barcoding animal life: cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society of London (B) 270, S96-S99. (Pubitemid 36960325)
57. HEBERT, P.D.N., PENTON, E.H., BURNS, J.M., JANZEN, D.H. & HALLWACHS, W. 2004a. Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences USA 101, 14812-14817. (Pubitemid 39410757)
58. HEBERT, P.D.N., STOECKLE, M.Y., ZEMLAK, T.S. & FRANCIS, C.M. 2004b. Identification of birds through DNA barcodes. PLoS Biology 2, e312.
59. HOBBS, G.A. 1968. Ecology of species of Bombus (Hymenoptera: Apidae) in southern Alberta. VII. Subgenus Bombus. Canadian Entomologist 100, 156-164.
60. ICZN 1996. Opinion 1828. Apis terrestris Linnaeus, 1758, A. muscorum Linnaeus, 1758 and A. lucorum Linnaeus, 1761 (currently Bombus terrestris, B. muscorum and B. lucorum) and Bombus humilis Illiger, 1806 (Insecta, Hymenoptera): specific names conserved, and neotypes designated for B. terrestris and B. muscorum. Bulletin of Zoological Nomenclature 53, 64-65.
61. ICZN 1999. International Code of Zoological Nomenclature, 4th edition. International Commission on Zoological Nomenclature, London.
62. INGS, T.C., SCHIKORA, J.&CHITTKA, L. 2005. Bumblebees, humble pollinators or assiduous invaders? A population comparison of foraging performance in Bombus terrestris. Oecologia 144, 508-516. (Pubitemid 41399462)
63. INOUE, M.N., YOKOYAMA, J. &WASHITANI, I. 2008. Displacement of Japanese native bumblebees by the recently introducedBombus terrestris (L.) (Hymenoptera: Apidae). Journal of Insect Conservation 12, 135-146. (Pubitemid 351325561)
64. ITO,M.&SAKAGAMI, S.F. 1980. The bumblebee fauna of theKurile Islands (Hymenoptera: Apidae). LowTemperature Science (B) 38, 23-51.
65. KAILA, L. & STAHLS, G. 2006. DNA barcodes: evaluating the potential of COI to differentiate closely related species of Elachista (Lepidoptera: Gelechioidea: Elachistidae) from Australia. Zootaxa 2006, 1-26. (Pubitemid 43926373)
66. KERZHNER, I.M. 1972. [Historical survey of studies of the insect fauna of theMongolian People's Republic]. Nasekomye Mongolii, Leningrad 1, 57-112.
67. KIMURA, M. 1980. A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution 16, 111-120. (Pubitemid 11166431)
68. KIRBY, W. 1837. Part 4. The insects. In: RICHARDSON, J., Ed., Fauna boreali-Americana; or the zoology of the northern parts of British America: containing descriptions of the objects of natural history collected on the late Northern Land expeditions, under command of Captain Sir John Franklin, R.N. Norwich, pp. xxxix + 325.
69. KNOWLES, L.L. & CARSTENS, B.C. 2007. Delimiting species without monophyletic gene trees. Systematic Biology 56, 887-895. (Pubitemid 350129639)
70. KRÜGER, E. 1951. Phaenoanalytische Studien an einigen Arten der Untergattung Terrestribombus O. Vogt (Hymen. Bomb.). I. Teil. Tijdschrift voor Entomologie 93, 141-197.
71. KRÜGER, E. 1954. Phaenoanalytische Studien an einigen Arten der Untergattung Terrestribombus O. Vogt (Hymenoptera, Bombidae). II. Teil. Tijdschrift voor Entomologie 97, 263-298.
72. KRÜGER, E. 1956. Phaenoanalytische Studien an einigen Arten der Untergattung Terrestribombus O. Vogt (Hymenoptera, Bombidae). II. Teil. Tijdschrift voor Entomologie 99, 75-105.
73. KRÜGER, E. 1958. Phaenoanalytische Studien an einigen Arten der Untergattung Terrestribombus O. Vogt (Hymenoptera, Bombiidae). III. Teil. Tijdschrift voor Entomologie 101, 283-344.
74. KUHLMANN, M., ELSE, G.R., DAWSON, A. & QUICKE, D.L.J. 2007. Molecular, biogeographical and phenological evidence for the existence of three western European sibling species in the Colletes succinctus group (Hymenoptera: Apidae). Organic Diversity and Evolution 7, 155-165. (Pubitemid 47031721)
75. KULLENBERG, B., BERGSTRöM, G. & STÄLLERG-STENHAGEN, S. 1970. Volatile components of the cephalic marking secretion of male bumble bees. Acta Chemica Scandinavica 24, 1481-1483.
76. LEVCHENKO, T.V. 2010. Fauna and ecology of bees (Hymenoptera: Apoidea) of the Moscow region [Abstract]. PhD thesis, Lomonosov Moscow State University.
77. LINNAEUS, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, Holmiae.
78. LINNAEUS, C. 1761. Fauna Suecica sistens animalia Suecica regni: Mammalia, Aves, Amphibia, Pisces, Insecta, Vermes. Distributa per classes & ordines, genera & species, cum differentiis specierum, synonymis auctorum, nominibus incolarum, locis natalium, descriptionibus insectorum, Stockholmiae.
79. LøKEN, A. 1966. Notes on Fabrician species of Bombus Latr. and Psithyrus Lep., with designations of lectotypes (Hym., Apidae). Entomologiske Meddelelser 34, 199-206.
80. LøKEN, A. 1973. Studies on Scandinavian bumble bees (Hymenoptera, Apidae). Norsk entomologisk Tidsskrift 20, 1-218.
81. LøKEN, A., PEKKARINEN, A. & RASMONT, P. 1994. Case 2638. Apis terrestris Linnaeus, 1758, A. muscorum Linnaeus, 1758 andA. lucorum Linnaeus, 1761 (currently Bombus terrestris, B. muscorum and B. lucorum) and Bombus humilis Illiger, 1806 (Insecta, Hymenoptera): proposed conservation of usage of the specific names. Bulletin of Zoological Nomenclature 51, 232-236.
82. MACFARLANE, R.P. & GURR, L. 1995. Distribution of bumble bees in New Zealand. New Zealand Entomologist 18, 29-36.
83. MADDISON, W.P. & MADDISON, D.R. 2001. MacClade analysis of phylogeny and character evolution Version 4. Sinauer Associates Inc., Sunderland, MA.
84. MADDISON, W.P. & KNOWLES, L.L. 2006. Inferring phylogeny despite incomplete lineage sorting. Systematic Biology 55, 21-30.
85. MAGNACCA, K.N. & BROWN, M.J.F. 2010. Mitochondrial heteroplasmy and DNA barcoding in Hawaiian Hylaeus (Nesoprosopis) bees (Hymenoptera: Colletidae). BMC Evolutionary Biology 10, 1-16.
86. MALLET, J. 1995. A species definition for the modern synthesis. Trends in Ecology and Evolution 10, 294-299.
87. MALLET, J. 1997. What are species? Trends in Ecology and Evolution 12, 453-454.
88. MATSUMURA, C., YOKOYAMA, J. & WASHITANI, I. 2004. Invasion status and potential impacts of an invasive alien bumblebee, Bombus terrestris L. (Hymenoptera: Apidae) naturalized in southern Hokkaido, Japan. Global Environment Research 8, 51-66.
89. MEIER, R., SHIYANG, K.,VAIDYA, G. & NG, P.K.L. 2006. DNA barcoding and taxonomy in Diptera: a tale of of high intraspecific variability and low identification success. Systematic Biology 55, 715-728.
90. MEUSNIER, I., SINGER, G.A.C., LANDRY, J.-F., HICKEY, D.A., HEBERT, P.D.N. & HAJIBABAEI, M. 2008. A universal DNA mini-barcode for biodiversity analysis. BioMed Central Genomics 9, 214 [4 pp.].
91. MEYER, C.P. & PAULAY, G. 2005. DNA barcoding: error rates based on comprehensive sampling. PLoS Biology 3, e422.
92. MILLIRON, H.E. 1971. A monograph of the western hemisphere bumblebees (Hymenoptera: Apidae; Bombinae). I. The genera Bombus and Megabombus subgenus Bombias. Memoirs of the Entomological Society of Canada 82, 1-80.
93. MONAGHAN, M.T., BALKE, M., GREGORY, T.R. & VOGLER, A.P. 2005. DNA-based species delineation in tropical beetles using mitochondrial and nuclear markers. Philosophical Transactions of the Royal Society (B) 360, 1925-1933. (Pubitemid 44365053)
94. MONAGHAN, M.T., WILD, R., ELLIOT, M., FUJISAWA, T., BALKE, M., INWARD, D.J.G., LEES, D.C., RANAIVOSOLO, R., EGGLETON, P., BARRACLOUGH, T.G. & VOGLER, A.P. 2009. Accelerated species inventory on Madagascar using coalescentbased models of species delineation. Systematic Biology 58, 298-311.
95. MURRAY, T.E., FITZPATRICK, U., BROWN, M.J.F. & PAXTON, R.J. 2008. Cryptic species diversity in a widespread bumble bee complex revealed using mitochondrial DNA RFLPs. Conservation Genetics 9, 653-666. (Pubitemid 351579817)
96. NAYLOR, G.J.P. & BROWN, W.M. 1997. Structural biology and phylogenetic estimation. Nature 388, 527-528.
97. NYLANDER, W. 1848. Adnotationes in expositionem monographicam apum borealium. Meddelanden af Societatis pro fauna et flora fennica 1, 165-282.
98. PACKER, L., SHEFFIELD, C.S., GIBBS, J., DE SILVA, N., BEST, L.R., ASCHER, J., AYALA, R., MARTINS, D., ROBERTS, S.P.M., TADAUCHI, O., KUHLMANN, M.,WILLIAMS, P.H., EARDLEY, C., DROEGE, S. & LEVCHENKO, T.V. 2009. The campaign to barcode the bees of the world: progress, problems, prognosis. In: YURRITA, C.L., Ed., Memorias: VI Congresso Mesoamericano Sobre Abejas Nativas. Universidad de San Carlos de Guatemala, Centro de Estudios Conservacionistas, pp. 178-180.
99. PAMILO, P., VARVIO-AHO, S.-L. & PEKKARINEN, A. 1984. Genetic variation in bumblebees (Bombus, Psithyrus) and putative sibling species of Bombus lucorum. Hereditas 101, 245-251.
100. PAMILO, P., TENGö, J., RASMONT, P., PIRHONEN, K., PEKKARINEN, A. & KAARNAMA, E. 1997. Pheromonal and enzyme genetic characteristics of the Bombus lucorum species complex in northern Europe. Entomologica Fennica 7, 187-194.
101. PANFILOV, D.V. 1956. Contribution to the taxonomy of bumblebees (Hymenoptera, Bombinae), including the description of new forms. Zoologicheskii Zhurnal 35, 1325-1334.
102. PAPADOPOULOU, A.,BERGSTEN, J., FUJISAWA, T.,MONAGHAN,M.T., BARRACLOUGH, T.G. & VOGLER, A.P. 2008. Speciation and DNA barcodes: testing the effects of dispersal on the formation of discrete sequence clusters. Philosophical Transactions of the Royal Society (B) 363, 2987-2996.
103. PEDERSEN, B.V. 2002. European bumblebees (Hymenoptera: Bombini) phylogenetic relationships inferred from DNA sequences. Insect Systematics and Evolution 33, 361-386. (Pubitemid 36171585)
104. PEKKARINEN, A. 1979. Morphometric, colour and enzyme variation in bumblebees (Hymenoptera, Apidae, Bombus) in Fennoscandia and Denmark. Acta Zoologica Fennica 158, pp. 60.
105. PÉREZ, J. 1895. Voyage de M. Ch. Alluaud aus 'îles Canaries (Novembre 1889-Juin 1890), 4emémoire (1). Hyménoptères. Annales de la Société entomologique de France 64, 191-201.
106. PÉREZ, J. 1905. Hyménoptères recueillis dans le Japon central, par M. Harmand, ministre plénipotentiaire de France à Tokio. Bulletin du Muséum national d'histoire naturelle 11, 23-39.
107. PITTIONI, B. 1939. Die Hummeln und Schmarotzerhummeln der Balkan-Halbinsel. II. Spezieller Teil. Izvestiya na Tsarskite prirodonauchni instituti v Sofiya 12, 49-115.
108. PLOWRIGHT, R.C. & STEPHEN, W.P. 1980. The taxonomic status of Bombus franklini (Hymenoptera: Apidae). Canadian Entomologist 112, 475-479.
109. PONS, J., BARRACLOUGH, T.G., GOMEZ-ZURITA, J., CARDOSO, A., DURAN, D.P., HAZELL, S., KAMOUN, S., SUMLIN, W.D. & VOGLER, A.P. 2006. Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology 55, 595-609.
110. POSADA, D. 2008. jModelTest: phylogenetic model averaging. Molecular Biology and Evolution 25, 1253-1256. (Pubitemid 351882013)
111. RASMONT, P. 1981. Redescription d'une espèceméconnue de bourdon d'Europe: Bombus lucocryptarum Ball, 1914 n. status. Bulletin et Annales de la Société Royale Belge d'Entomologie 117, 149-154.
112. RASMONT, P. 1983. Notes taxonomiques sur les bourdons (Hymenoptera, Apidae). Bulletin et Annales de la Société Royale Belge d'Entomologie 119, 167-170.
113. RASMONT, P. 1984. Les bourdons du genre Bombus Latreille sensu stricto en Europe occidentale et centrale (Hymenoptera, Apidae). Spixiana 7, 135-160.
114. RASMONT, P., COPPÉE, A., MICHEZ, D. & MEULEMEESTER, T.D. 2008. An overview of the Bombus terrestris (L. 1758) subspecies (Hymenoptera: Apidae). Annales de la Société entomologique de France 44, 243-250.
115. RASMONT, P., SCHOLL, A., DE JONGHE, R., OBRECHT, E. & ADAMSKI, A. 1986. Identité et variabilité des mâles de bourdons du genre Bombus Latreille sensu stricto en Europe occidentale et centrale (Hymenoptera, Apidae, Bombinae). Revue suisse Zoologique 93, 661-682.
116. RATNASINGHAM, S. & HEBERT, P.D.N. 2007. BOLD: The Barcode Of Life Data system (www.barcodinglife.org). Molecular Ecology Notes 2007, 1-10.
117. REINIG, W.F. 1930. Untersuchungen zur Kenntnis der Hummelfauna des Pamir-Hochlandes. Zoologische Ergebnisse der deutsch-russischen Alai-Pamir-Expedition der Notgemeinschaft der Deutschen Wissenschaft und der Akademie der Wissenschaften der U.d.S.S.R. Zeitschrift für Morphologie und ökologie der Tiere 17, 68-123.
118. RUBINOFF, D., CAMERON, S.A. & WILL, K. 2006. A genomic perspective on the shortcomings of mitochondrial DNA for "barcoding" identification. Journal of Heredity 97, 581-594. (Pubitemid 46033025)
119. SAKAGAMI, S.F. & ISHIKAWA, R. 1969. Note préliminaire sur la répartition géographique des bourdons japonais, avec descriptions et remarques sur quelques formes nouvelles ou peu connues. Journal of the Faculty of Science, Hokkaido University (Zoology) 17, 152-196.
120. SAKAGAMI, S.F. & ISHIKAWA, R. 1972. Note supplémentaire sur la taxonomie et répartition géographique de quelques bourdons Japonais, avec la description d'une nouvelle sousesp̀ ece. Bulletin of the National Science Museum, Tokyo 15, 607-616.
121. SALMON,M.A. 2000. The Aurelian Legacy. British Butterflies and their Collectors. Harley Books, Great Horkesley.
122. SCHMID-HEMPEL, P.,SCHMID-HEMPEL, R.,BRUNNER, P.C., SEEMAN, O.D. & ALLEN, G.R. 2007. Invasion success of the bumblebee, Bombus terrestris, despite a drastic genetic bottleneck. Heredity 99, 414-422. (Pubitemid 47482640)
123. SCHMIDT, B.C. & SPERLING, F.A.H. 2008.Widespread decoupling of mtDNA variation and species integrity in Grammia tiger moths (Lepidoptera: Noctuidae). Systematic Entomology 33, 613-634.
124. SCHOLL, A. & OBRECHT, E. 1983. Enzymelektrophoretische Untersuchungen zur Artabgrenzung im Bombus lucorum Komplex (Apidae, Bombini). Apidologie 14, 65-78.
125. SCHOLL, A., OBRECHT, E. & OWEN, R.E. 1990. The genetic relationship between Bombus moderatus Cresson and the Bombus lucorum auct. species complex (Hymenoptera: Apidae). Canadian Journal of Zoology 68, 2264-2268.
126. SJÖSTEDT, S. & HUMMEL, D. 1932. Schwedisch-chinesische wissenschaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sü Pingchang. Insekten gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927-1930. Einleitung. Arkiv för zoologi 25A, 9-34.
127. SKEVINGTON, J.H., KEHLMAIER, C. & STAHLS, G. 2007. DNA barcoding: mixed results for big-headed flies (Diptera: Pipunculidae). Zootaxa 2007, 1-26. (Pubitemid 46473096)
128. SKORIKOV, A.S. 1910. [Intraspecific forms of Bombus mendax Gerst.] (Hymenoptera, Bombidae). Russkoe éntomologicheskoe Obozrênie 9(1909), 328-330.
129. SKORIKOV, A.S. 1912. Neue Hummelformen (Hymenoptera, Bombidae). IV. Russkoe éntomologicheskoe Obozrênie 12, 606-610.
130. SKORIKOV, A.S. 1913. Neue Hummelformen (Hymenoptera, Bombidae). V. Russkoe éntomologicheskoe Obozrênie 13, 171-175.
131. SKORIKOV, A.S. [1923]. [Palaearctic bumblebees. Part I. General biology (including zoogeography)]. Izvestiya Severnoi oblastnoi stantsii zashchity rastenii ot vreditelei 4(1922), 1-160.
132. SKORIKOV, A.S. 1931. Die Hummelfauna Turkestans und ihre Beziehungen zur zentralasiatischen Fauna (Hymenoptera, Bombidae). In: LINDHOLM, V.A., Ed., Abhandlungen der Pamir-Expedition 1928, vol. 8. Academy of Sciences of the USSR, Leningrad, pp. 175-247.
133. SKORIKOV, A.S. 1933. Zur Hummelfauna Japans und seiner Nachbarl änder. Mushi 6, 53-65.
134. SMITH, F. 1852. Descriptions of some new and apparently undescribed species of hymenopterous insects from north China, collected by Robert Fortune, Esq. Transactions of the Entomological Society of London 2, 33-45.
135. SMITH, F. 1854. Catalogue of hymenopterous insects in the collection of the British Museum. Part II. Apidae. Trustees of the British Museum, London.
136. SMITH, F. 1861. Descriptions of new genera and species of exotic Hymenoptera. Journal of Entomology 1, 146-155.
137. SMITH, F. 1873. Descriptions of aculeate Hymenoptera of Japan, collected by Mr. George Lewis at Nagasaki and Hiogo. Transactions of the Entomological Society of London 1873, 181-206.
138. SONG, H., BUHAY, J.E., WHITING, M.F. & CRANDALL, K.A. 2008. Many species in one: DNA barcoding overestimates the number of species when nuclear mitochondrial pseudogenes are coamplified. Proceedings of the National Academy of Sciences USA 105, 13486-13491.
139. SRIVATHSAN, A. & MEIER, R. 2011. On the inappropriate use of Kimura-2-parameter (K2P) divergences in the DNAbarcoding literature. Cladistics 27, 1-5.
140. STOUT, J.-C. & GOULSON, D. 2000. Bumble bees in Tasmania: their distribution and potential impact on Australian flora and fauna. Bee-World 81, 80-86.
141. TAMURA, K., DUDLEY, J., NEI, M. & KUMAR, S. 2007. MEGA4: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0. Molecular Biology and Evolution 24, 1596-1599. (Pubitemid 47236692)
142. THORP,R.W. 1970. The type locality of Bombus franklini and notes on putative Arizona records of other Bombini (Hymenoptera: Apidae). Pan-Pacific Entomologist 46, 177-180.
143. THORP, R.W. 2003. Bumble bees (Hymenoptera: Apidae): commercial use and environmental concerns. In: STRICKLER, K. & CANE, J.H., Ed., For Nonnative Crops, Whence Pollinators of the Future? Entomological Society of America, Lanham, MD, pp. 21-40.
144. THORP, R.W. & SHEPHERD, M.D. 2005. Profile: Subgenus Bombus Latreille, 1802 (Apidae: Apinae: Bombini). In: SHEPHERD, M.D., VAUGHAN, D.M. & BLACK, S.H., Eds., Red List of Pollinator Insects of North America. The Xerces Society for Invertebrate Conservation, Portland, OR, pp. 5.
145. TKALCU, B. 1962. Contribution à l'étude des bourdons du Japon (I) (Hymenoptera, Apoidea). Bulletin de la Société entomologique de Mulhouse 1962, 81-100.
146. TKALCU, B. 1967. Sur deux espèces de bourdons décrites par William Nylander (Hymenoptera, Apoidea: Bombus). Bulletin de la Société entomologique de Mulhouse 1967, 41-58.
147. TKALCU, B. 1969. Ergebnisse der Albanien-Expedition 1961 des Deutschen Entomologischen Institutes. 78. Beitrag. Hymenoptera: Apidae IV (Bombinae). Beiträge zur Entomologie 19, 887-916.
148. TKALCU, B. 1974a. Eine Hummel-Ausbeute aus dem Nepal- Himalaya (Insecta, Hymenoptera, Apoidea, Bombinae). Senckenbergiana biologica 55, 311-349.
149. TKALCU, B. 1974b. Bemerkenswerte Bienenfunde in der Tschechoslowakei (Hymenoptera, Apoidea). Acta Entomologica Bohemoslovaca 17, 205-208.
150. TKALCU, B. 1974c. Ergebnisse der 1. und 2. mongolischtschechoslowakischen entomologisch-botanischen Expedition in der Mongolei. Nr. 29: Hymenoptera, Apoidea, Bombinae. Sborník faunistickych prací Entomologického oddelení Národního musea v Praze 15, 25-57.
151. TOKORO, S., YONEDA, M., KUNITAKE, Y.K. & GOKA, K. 2010. Geographic variation in mitochondrial DNA of Bombus ignitus(Hymenoptera: Apidae). Applied Entomology and Zoology 45, 77-87.
152. TORRETTA, J.P., MEDAN, D. & ABRAHAMOVICH, A.H. 2006. First record of the invasive bumblebee Bombus terrestris (L.) (Hymenoptera, Apidae) in Argentina. Transactions of the American Entomological Society 132, 285-289. (Pubitemid 46448771)
153. TRAUTMANN, G. & TRAUTMANN, W. 1915. Beitrag zur Hummelkenntnis das europäischen arktischen Gebietes. Internationale Entomologische Zeitschrift 9, 189.
154. VOGT, O. 1908. Bombi (Hummeln). In: Wissenschaftliche Ergebnisse von Expedition Filchner nach China und Tibet 1903-1905, X. Band I. Teil, 1. Abschnitt: Zoologische Sammlungen, Berlin, pp. 100-101.
155. VOGT, O. 1909. Studien über das Artproblem. 1. Mitteilung. Ü ber das Variieren der Hummeln. 1. Teil. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin 1909, 28-84.
156. VOGT, O. 1911. Studien über das Artproblem. 2. Mitteilung. Ü ber das Variieren der Hummeln. 2. Teil. (Schluss). Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin 1911, 31-74.
157. WATERS, J., DARVILL, B., LYE, G.C. & GOULSON, D. 2010. Niche differentiation of a cryptic bumblebee complex in theWestern Isles of Scotland. Insect Conservation and Diversity 2010, 1-7.
158. WERREN, J.H. 1997. Biology of Wolbachia. Annual Reviews of Entomology 42, 587-609.
159. WILL, K.W., MISHLER, B.D. & WHEELER, Q.D. 2005. The perils of DNA barcoding and the need for integrative taxonomy. Systematic Biology 54, 844-851.
160. WILLIAMS, P.H. 1991. The bumble bees of the Kashmir Himalaya (Hymenoptera: Apidae, Bombini). Bulletin of the British Museum (Natural History) (Entomology) 60, 1-204.
161. WILLIAMS, P.H. 1995. Phylogenetic relationships among bumble bees (Bombus Latr.): a reappraisal of morphological evidence. Systematic Entomology 19, 327-344.
162. WILLIAMS, P.H. 1998. An annotated checklist of bumble bees with an analysis of patterns of description (Hymenoptera: Apidae, Bombini). Bulletin of The Natural History Museum (Entomology) 67, 79-152 [updated at www.nhm.ac.uk/bombus/ accessed 2010].
163. WILLIAMS, P.H. 2000. Are Bombus lucorum and magnus separate species? BWARS Newsletter 2000, 15-17.
164. WILLIAMS, P.H. 2007. The distribution of bumblebee colour patterns world-wide: possible significance for thermoregulation, crypsis, and warning mimicry. Biological Journal of the Linnean Society 92, 97-118.
165. WILLIAMS, P.H., AN, J.-D. & HUANG, J.-X. 2011. The bumblebees of the subgenus Subterraneobombus: integrating evidence from morphology and DNA barcodes (Hymenoptera, Apidae, Bombus). Zoological Journal of the Linnean Society 163, 813-862.
166. WILLIAMS, P.H., AN, J.-D., HUANG, J.-X. & YAO, J. 2010. BBCI: A new initiative to document Chinese bumble bees for pollination research. Journal of Apicultural Research 49, 221-222.
167. WILLIAMS, P.H., CAMERON, S.A., HINES, H.M., CEDERBERG, B. & RASMONT, P. 2008. A simplified subgeneric classification of the bumblebees (genus Bombus). Apidologie 39, 46-74. (Pubitemid 351507455)
168. WILLIAMS, P.H., TANG, Y., YAO, J. & CAMERON, S. 2009. The bumblebees of Sichuan (Hymenoptera: Apidae, Bombini). Systematics and Biodiversity 7, 101-190.
169. WINTER, K., ADAMS, L., THORP, R.W., INOUYE, D.W., DAY, L., ASCHER, J. & BUCHMANN, S. 2006. Importation of non-native bumble bees into North America: potential consequences of using Bombus terrestris and other non-native bumble beesfor greenhouse crop pollination in Canada, Mexico, and the United States. AWhite Paper of the North American Pollinator Protection Campaign (NAPCC), San Francisco, pp. 33.
170. WOLF, S., ROHDE, M. & MORITZ, R.F.A. 2010. The reliability of morphological traits in the differentiation of Bombus terrestris and B. lucorum (Hymenoptera: Apidae). Apidologie 41, 45-53.
171. YAO, J. & WANG, S.-F. 2005. Hymenoptera, Apidae, Bombini. In: YANG, X., Ed., Insect Fauna of Middle-west Qiling Range and South Mountains of Gansu Province. Science Press, Beijing, pp. 890-896.
172. ZHAO, Y.-Z., AN, J.-D., ZHOU, Z.-Y., DONG, J.,XING, Y.-H. & QIN, J.-J. 2011. Pollination behavior of Apis mellifera ligustica and Bombus hypocrita (Hymenoptera, Apidae) and the influencing factors in peach greenhouse. Acta Entomologica Sinica 54, 89-96.