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Bees of the Genus Dufourea Lepeletier (Hymenoptera: Halictidae: Rophitinae) of Canada*

Sheila Dumesh1**

Corey S. Sheffield2

1Department of Biology, York University 4700 Keele St., Toronto, ON M3J 1P3 [email protected]
2Royal Saskatchewan Museum, 2445 Albert Street, Regina, SK S4P 4W7 [email protected]

* This paper is contribution #13 from the Canadian Pollination Initiative. **Corresponding Author.

Bees of the Genus Dufourea Lepeletier (Hymenoptera: Halictidae: Rophitinae) of Canada*

Bees of the Genus Dufourea Lepeletier (Hymenoptera: Halictidae: Rophitinae) of Canada*

Sheila Dumesh1**

Corey S. Sheffield2

1Department of Biology, York University 4700 Keele St., Toronto, ON M3J 1P3 [email protected]
2Royal Saskatchewan Museum, 2445 Albert Street, Regina, SK S4P 4W7 [email protected]

* This paper is contribution #13 from the Canadian Pollination Initiative. **Corresponding Author.

Abstract

Dufourea Lepeletier is the only representative of the subfamily Rophitinae in Canada. An illustrated key to females and males of the eight Canadian species is provided, and each is described; the male of D. fimbriata, and females of D. holocyanea and D. maura are described for the first time. Most Dufourea are floral specialists, so their presence within habitats is determined by that of their host. Floral hosts and known distributions of Dufourea in Canada are provided.

Dufourea monardae Photo by Cory S. Sheffield

Introduction

Rophitine bees (Rophitinae) are a relatively small (214 species, as per Michener (2007); 257 species listed by Ascher and Pickering (2012)), basal subfamily of the Halictidae (Pesenko 1999) containing four tribes: the Rophitini (Old and New World members), Penapini (South America), Conanthalictini and Xeralictini (both restricted to south-western North America) (Patiny et al. 2007).  The Rophitinae are primarily Holarctic in distribution (Michener 1965; Rozen 1997; Niu et al. 2005) with about a dozen species occurring in the Southern Hemisphere: five in South America (Rozen 1997) and six or seven in Africa (Patiny and Michez 2006, 2007a, 2007b).  Rophitinae is the only subfamily of Halictidae in which all members are solitary (Pesenko and Astafurova 2006; Patiny et al. 2007; Danforth et al. 2008; Richards and Packer 2010).  Despite the paucity of species in the Southern Hemisphere, Patiny et al. (2007) suggest that the strictly South American tribe Penapini forms the sister group to the remaining Northern Hemisphere tribes, with all the Old World taxa having arisen from among the New World Rophitini.  The genus Dufourea Lepeletier in North America represents a dispersal event back from the Old World (Patiny et al. 2007).  As such, it is the only holarctic genus of Rophitinae (Michener 1965), and also the only one occurring in Canada.
Disagreements in higher level classification exist within the Rophitinae, particularly between classifications used in the Old and New World.  For instance, Niu et al. (2005) and Pesenko and Astafurova (2006) recognize 16 genera of Rophitinae, while Michener (2007) and Patiny et al. (2007) recognize 13, the latter two studies maintain Trilia Vachal as a subgenus of Dufourea, and Rhopitoides Schenck as a subgenus of Rophites Spinola, and place Flavodufourea Ebmer as a subgenus of Rophites Spinola.  Although Flavodufourea was originally described as a subgenus of Dufourea, a position supported by Patiny (2003), Michener (2007) believes its placement is still not decisive.  Michener (2007) also does not recognize subgenera within Dufourea.
Six genera of Rophitinae occur in North America (Michener 2007).  The largest genus is Dufourea (ca. 70 species), followed by Conanthalictus Cockerell (13 species), Micralictoides Timberlake (8 species), Protodufourea Timberlake (5 species), Sphecodosoma Crawford (3 species), and Xeralictus Cockerell (2 species).  Michener (1965) reviewed the Rophitinae of the Western Hemishpere; Snelling and Stage (1995) clarified the placement of the Xeralictini within the Rophitinae.
Dufourea are relatively uncommon bees in Canada, and no comprehensive account of the species has been published.  Morphologically, Dufourea is distinctive among the Canadian bee fauna, having antennae that arise low on the face, usually just above the upper margin of the clypeus.  Dufourea differs from other non-cleptoparasitic Halictidae in Canada in having the scopa restricted to the hind tibia (not present also on the hind femur as in other halictids), and in having a short clypeus which seldom is any longer than the labrum (Michener 1965).  Dufourea is also the only genus of Halictidae in Canada with two submarginal cells (the exceptions being a few species of Lasioglossum (Dialictus) and Sphecodes).  Keys in Mitchell (1960), Stephen et al. (1969); Michener et al. (1994), and Michener (2000; 2007) can be used to identify this genus in North America; Packer et al. (2007) provide an interactive web-based key for bee genera in eastern Canada.  Mitchell (1960) published keys to the four species found in the eastern Canada.
All Rophitinae, including Dufourea, nest in soil (Pesenko and Astafurova 2006), and construct a single nesting tunnel with lateral burrows which end in single cells, or a series of multiple cells (Michener 2007).  Rozen and McGinley (1976), Kukuk et al. (1985), and Rozen (1993) summarize the nesting biology of Rophitinae.  The nesting biology of three of the eight Canadian Dufourea have previously been studied: D. trochantera Bohart (Torchio et al. 1967); D. novaeangliae (Robertson) (Eickwort et al. 1983), D. holocyanea (Cockerell), including the description of the mature larvae (Rozen and Ozbek (2008).
Dufourea species are well known for their floral specialization (Lincoln 1981; Patiny et al. 2008).  Floral specialization has lead to morphological adaptations in many bees, including Dufourea; Alves-dos-Santos (2003) found that mouthparts of bees specializing on Pontederiaceae, including D. novaeangliae, are covered with hairs to remove pollen from the concealed anthers.  Each species of Dufourea in Canada specializes on a different plant family (Table 1).  Although Lincoln (1981) reported D. dilatipes Bohart on both Calochortus (Liliaceae) and Campanula (Campanulaceae) in Canada, it seems the latter floral record may actually be based on misidentification of specimens of D. maura (Cresson) as one of the two pollen samples examined was from Ontario (see Lincoln 1981), far outside the known range of D. dilatipes (Plate 1, Figure E).  Calochortus has a Canadian distribution restricted to southern AB and BC; only one species, C. apiculatus, is recorded from Waterton Lakes National Park, one of only a few known collections site of D. dilatipes in Canada (and one of the localities for the type series).  Most of the other species of Dufourea in Canada are also found in the west; D. holocyanea occurs in southern BC (Plate 3, Figure E), and D. trochantera occurs in BC and AB (Plate 8, Figure E).  Dufourea fimbriata (Cresson) has only been collected a few times in Canada, in AB (Edmonton), in NT (Yellowknife), the most northern record for this genus in North America, and in northern ON (Plate 2, Figure E).  Dufourea maura is the most widespread species, occurring from southern BC to ON (Plate 5, Figure E).  Dufourea marginata (Cresson) is also widespread in Canada, occurring in southern BC-ON.  Two species, D. monardae (Cresson) and D. novaeangliae occur in eastern Canada; D. monardae is known only from southern ON (Richards et al. 2011) (Plate 6, Figure E) and the adjacent United States (Bouseman 1976), D. novaeangliae has been collected in ON, QC (Payette 2001) and NS (Sheffield et al. 2003) (Plate 7, Figure E).
Our objectives are to provide an illustrated key to the eight species of Dufourea occurring in Canada, full species descriptions, images of male genitalia, distributional notes, and a general summary of floral relations.

 

Materials and Methods

Specimens examined were from the following institutions: Canada: Packer Collection York University, Toronto, ON; Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, ON; Royal British Columbia Museum, Victoria, BC; Beaty Biodiversity Museum, University of British Columbia, Vancouver, BC; Royal Alberta Museum, Edmonton, AB; E.H. Strickland Entomological Museum, University of Alberta, Edmonton, AB; Brock University, St. Catharines, ON; Wallis-Roughley Museum of Entomology (formerly J.B. Wallis Museum), University of Manitoba, Winnipeg, MB; United States: United States Department of Agriculture Bee Biology and Systematics Laboratory, Logan, Utah; KU Biodiversity Institute, University of Kansas, Lawrence, Kansas.
Terminology used for the species descriptions includes: pd (puncture diameter); MOD (median ocellar diameter); IAD (interantennal distance); AOD (antennal-ocular distance); OVD ([median] ocellar-vertexal distance); OOD (ocellar-ocular distance); IOD (inter-ocellar distance); Tx (tergum x); Sx (sternum x).
Head breadth was measured as the greatest distance between outer most margins of compound eyes; head length was measured as the distance from the apex of the clypeus to the summit of the vertexal area.  In the descriptions, head breadth to length is given as a ratio (b:l).
Images were taken using a Visionary Digital BK Plus imaging system using a Canon EOS 40D digital SLR camera and processed with Adobe Photoshop.

Table 1.  Floral hosts of Dufourea (Halictidae) species occurring in Canada.

Dufourea species Floral Host Floral Host Family Reference
D. dilatipes Calochortus Liliaceae Lincoln 1981
D. fimbriata Potentilla Rosaceae Lincoln 1981
D. holocyanea Symphoricarpos Caprifoliaceae Lincoln 1981
D. maura Campanula Campanulaceae Lincoln 1981
D. marginata Helianthus Compositae Lincoln 1981
D. trochantera Phacelia Hydrophyllaceae Lincoln 1981
D. novaeangliae Pontedereria Pontederiaceae Bouseman 1986
D. monardae Monarda Labiatae Bouseman 1976

Species

Species List

Dufourea dilatipes
Dufourea fimbriata
Dufourea holocyanea
Dufourea marginata
Dufourea maura
Dufourea monardae
Dufourea novaeangliae
Dufourea trochantera

Acknowledgments

We thank Laurence Packer (York University, Toronto, ON), Jason Gibbs (Cornell University, Ithaca, NY), John Huber (Canadian National Collection, Ottawa, ON) and Terry Griswold (United States Department of Agriculture Bee Biology and Systematics Laboratory, Logan, UT) for helpful comments. We thank other donors of materials used in this study: Lincoln Best (York University), Andy Bennett (Canadian National Collection, Ottawa, ON), Rob Cannings (Royal British Columbia Museum, Victoria, BC), Karen Needham (Beaty Biodiversity Museum, University of British Columbia, Vancouver, BC), Matt Buck (Royal Alberta Museum, Edmonton, AB), Felix Sperling (E.H. Strickland Collection, University of Alberta, Edmonton, AB), Miriam Richards (Brock University, St. Catharines, ON), Terry Galloway (Wallis-Roughley Museum of Entomology, Winnipeg, MB), Sarah Semmler (University of Manitoba, Winnipeg, MB), Diana Bizecki Robson (Manitoba Museum, Winnipeg, MB), Terry Griswold, and Charles Michener (KU Biodiversity Institute, University of Kansas, Lawrence, KS). We also thank Claudia Ratti for her technical assistance with imaging and the web-based key. Funding for this study was provided by the Canadian Pollinator Initiative (NSERC-CANPOLIN), the Canadian Foundation for Innovation and Ontario Research Fund, both through Canadensys and a Discovery Grant to Laurence Packer.

References

Alves-dos-Santos, I. 2003. Adaptations of bee proboscides for collecting pollen from Pontederiaceae flowers. pgs. 257-263 in G.A.R. Melo and I. Alves-dos-Santos (eds). Apoidea Neotropica: Homenagem aos 90 Anos de Jesus Santiago Moure. Editoria UNESC, Criciúma.

Arduser, M.S. 1986. Records of Dufourea maura (Hymenoptera: Halictidae) from Isle Royale National Park, Michigan. Great Lakes Entomologist 19: 175-176.

Ascher, J. S. and J. Pickering. 2012. Bee species guide (Hymenoptera: Apoidea: Anthophila). http://www.discoverlife.org/mp/20q?guide=Apoidea_species

Bohart, G.E. 1948. New North American bees of the genus Dufourea (Hymenoptera: Halictidae) PART II. Annals of the Entomological Society of America 41(1):135.

Bouseman, J.K. 1976. Dufourea monardae (Viereck) in Illinois and Michigan (Hymenoptera: Apoidea). Journal of the Kansas Entomological Society 49: 531-532.

Bouseman, J.K. 1986. Dufourea novaeangliae in Illinois, with confirmation of host plant (Hymenoptera: Halictidae). Great Lakes Entomologist 19: 203-204.

Cockerell, T.D.A. 1906. Descriptions and records of bees – VIII. Journal of Natural History Series 7 17(98): 222-230.

Cockerell, T.D.A. 1925. Bees in the collection of the California Academy of Sciences. Proceedings of the California Academy of Sciences 14: 185-215.

Cresson, E.T. 1878. Descriptions of new North American Hymenoptera in the collection of the American Entomological Society. Transcripts of American Entomological Society 7: 63-136.

Danforth, B.N, C. Eardley, L. Packer, K. Walker, A. Pauly, and F.J. Randrianambinintsoa. 2008. Phylogeny of Halictidae with an emphasis on endemic African Halictinae. Apidologie 39: 86-101.

Eickwort, G.C., P.R. Kukuk, and F.R. Wesley. 1986. The nesting biology of Dufourea novaeangliae (Hymenoptera: Halictidae) and the systematic position of the Dufoureinae based on behaviour and development. Journal of the Kansas Entomological Society 59: 103-120.

Kukuk, P.F., G.C. Eickwort, and F.R. Wesley. 1985.  Mate-seeking behavior of Dufourea novaeangliae (Hymenoptera: Halictidae: Dufoureinae): the effects of resource depletion. Journal of the Kansas Entomological Society 58: 142-150.

Lincoln, P.G. 1981. Phylogeny, Host Plant Relations and Speciation in Genus Dufourea (Hymenoptera; Apoidea). PhD Thesis, University of California, Santa Cruz. 307 pp.

Michener, C.D. 1937. Records and descriptions of North American bees. Journal of Natural History Series 10 19(112): 393-410.

Michener, C.D. 1951. In Muesebeck, C.F.W., K.V.Krombein, and H.K. Townes. Hymenoptera of America north of Mexico. United States Department of Agriculture, Agriculture Monograph 2: 1132.

Michener, C.D. 1965. A generic review of the Dufoureinae of the Western Hemishpere (Hymenoptera: Halictidae). Annals of the Entomological Society of America 58: 321-326.

Michener, C. D. 2000. The Bees of the World. Johns Hopkins University Press, Baltimore, MD. xiv + 913 pp.

Michener, C. D. 2007. The Bees of the World. 2nd Ed. Johns Hopkins University Press, Baltimore, MD. xvi + 953 pp.

Michener, C.D., R.J. McGinley, and B.N. Danforth. 1994. The Bee Genera of North and Central America (Hymenoptera: Apoidea). Smithsonian Institution Press, Washington, DC. vii + 209 pp.

Mitchell, T.B. 1960. Bees of the Eastern United States, Vol. 1. Technical Bulletin, North Carolina Agricultural Experiment Station No 141. 538 pp.

Niu, Z., Y. Wu, and D. Huang. 2005. A taxonomic study of the four genera of the subfamily Rophitinae from China (Hymenoptera: Halictidae). The Raffles Bulletin of Zoology 53: 47-58.

Packer, L., J.A. Genal arearo, and C.S. Sheffield. 2007. The bee genera of eastern Canada. Canadian Journal of Arthropod Identification 3: 1-33. (http://www.biology.ualberta.ca/bsc/ejournal/pgs03/pgs_03.html)

Patiny, S. 2003. Revision of the subgenus Dufourea (Flavodufourea) Ebmer, 1984 (Hymenoptera, Halictidae, Rophitinae) and description of a new species D. (Flavodufoureaulkenkalkana sp.nov. from Kazakhstan. Zootaxa 255: 1-8.

Patiny, S. and D. Michez. 2006. Phylogenetic analysis of the Systropha Illiger 1806
(Hymenoptera: Apoidea: Halictidae) and description of a new subgenus. Annales
de la Société Entomologique de France 41: 27-44.

Patiny, S. and D. Michez. 2007a. New insights on the distribution and floral choices of
Systropha Illiger, 1806 in Africa (Hymenoptera: Apoidea), with description of a
new species from Sudan. Zootaxa 1461: 59-68.

Patiny, S. and D. Michez. 2007b. Biogeography of bees (Hymenoptera: Apoidea) in
Saraha and the Arabian deserts. Insect Systematics & Evolution 38: 19-34.

Patiny, S., D. Michez, and B.N. Danforth. 2008. Phylogenetic relationships and host-plant evolution within the basal clade of Halictidae (Hymenoptera, Apoidea). Cladistics 24: 255-269.

Payette, A. 2001. Première mention de l’abeille Dufourea novaeangliae (Robertson) (Hymenoptera : Halictidae) pour le Québec. Fabreries 26: 20-25.

Pesenko, Y.A. 1999. Phylogeny and classification of the family Halictidae revised
(Hymenoptera: Apoidea). Journal of the Kansas Entomological Society 72: 104–
123.

Pesenko, Y.A., and Y.V. Astafurova. 2006. Contributions to the Halictidae fauna of the Eastern Palaearctic Region: subfamily Rophitinae (Hymenoptera: Halictidae). Entomofauna 27: 317-356.

Richards, M.H., and L. Packer. 2010. Social behaviours in solitary bees: interactions among individuals in Xeralictus bicuspidariae Snelling (Hymenoptera: Halictidae: Rophitinae). Journal of Hymenoptera Research 19: 66-76.

Robertson, C. 1897. North American bees: descriptions and synonyms. Transactions of
the Academy of Sciences of St. Louis 7: 315-356.

Robertson, C. 1895. Notes of bees, with descriptions of new species. Transcations of
American Entomological Society 22(2): 115-128.

Rozen, J.G., Jr. 1997. South American rophitine bees (Hymenoptera: Halictidae:
Rophitinae). American Museum Novitates 3206: 1-27.

Rozen, J.G., Jr., and H.T. Özbek. 2008. Immatures of rophitine bees, with notes on their
nesting biology (Hymenoptera: Apoidea: Halictidae). American Museum
Novitates 3609: 1-35.

Scoggan, H.J. 1978. The Flora of Canada. Part 2. Pteridophyta, Gymnospermae,
Monocotyledoneae. National Museum of Natural Sciences, Ottawa, ON.

Sheffield, C.S., P.G. Kevan, R.F. Smith, S.M. Rigby, and R.E.L. Rogers. 2003.  Bee species of Nova Scotia, Canada, with new records and notes on bionomics and floral relations (Hymenoptera: Apoidea). Journal of the Kansas Entomological Society 76: 357-384.

Snelling, R.R., and G.I. Stage. 1995. Systematics and biology of the bee genus Xeralictus (Hymenoptera: Halictidae, Rophitinae). Natural History Museum of Los Angeles County Contributions in Science 451: 1-17.

Stephen, W.P., G.E. Bohart, and P.F. Torchio. 1969. The Biology and External Morphology of Bees with a Synopsis of the Genera of Northwestern America. Agricultural Experiment Station, Oregon State University, Corvallis. 140 pp.

Torchio, P.F., J.G. Rozen, Jr., G.E. Bohart, and M.S. Favreau. 1967. Biology of Dufourea and of its cleptoparasite, Neopasites (Hymenoptera: Apoidea). Journal of the New York Entomological Society 75: 132-146.

Viereck, 1904. A bee visitor to Pontederia (Pickerel Weed). Entomological News 15: 244-246.

Viereck, H.L. 1924. The identity of Conohalictoides noveangliae Robertson. Proceedings of the Entomological Society of Washington 26(1): 14-15.

Cite

Dumesh, S., Sheffield, C.S. 2012. Bees of the Genus Dufourea Lepeletier (Hymenoptera: Halictidae: Rophitinae) of Canada. Canadian Journal of Arthropod Identification No. 18, XX May 2012, available online at doi: 10.3752/cjai.2012.20