Hymenoptera: Apoidea, Apiformes) of the Prairies Ecozone with Comparisons to Other Grasslands of Canada
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427 Chapter 11 The Bees (Hymenoptera: Apoidea, Apiformes) of the Prairies Ecozone with Comparisons to other Grasslands of Canada Cory S. Sheffield Royal Saskatchewan Museum, 2340 Albert Street, Regina, SK, S4P 2V7 Email: [email protected] S. Danae Frier Royal Saskatchewan Museum, 2340 Albert Street, Regina, SK, S4P 2V7 University of Regina, Department of Biology, 3737 Wascana Parkway, Regina, SK, S4S 0A2 Sheila Dumesh York University, Biology Department, 4700 Keele Street, Toronto, ON, M3J 1P3 Abstract. The prairie grasslands of Canada are home to at least 387 bee species. This represents almost half (i.e., 48%) of the bee taxa reported for Canada and over 90% of the total recorded from the three Prairie Provinces of Alberta, Saskatchewan, and Manitoba. The Prairies Ecozone thus represents one of the most diverse and important areas supporting bee diversity in the country, with 23% of the bee species listed not found anywhere else in Canada, a value similar to that of the tallgrass savanna of the Mixedwood Plains Ecozone (southern Ontario and Québec) and only surpassed by that of the bunchgrass/sagebrush grasslands of the Western Interior Basin Ecozone (south central British Columbia) with respect to endemism. However, the Canadian Prairies support a higher proportion of bee species with dietary specialization; over half of these are oligoleges of the Asteraceae. Over 70% of the native prairie grasslands were lost before 1990. Therefore, it is likely that many of the endemic floral specialists have also become much less common, though little baseline data exist for rigorous comparison. Much of the Canadian Prairies have not been surveyed extensively for bees, and several new records are reported here. A full taxonomic review of the bees of the prairie grasslands is presented, including bionomic summaries. Résumé. Les prairies canadiennes abritent au moins 386 espèces d’abeilles, soit près de la moitié (48 %) de tous les taxons de ce groupe répertoriés au Canada et plus de 90 % du total des taxons signalés dans les trois provinces des Prairies —Alberta, Saskatchewan et Manitoba. L’écozone des prairies constitue une des régions les plus importantes pour le maintien de la diversité des abeilles au Canada : 23 % des espèces d’abeilles recensées n’existent nulle part ailleurs dans le pays, un pourcentage d’endémisme semblable à celui établi pour les prairies à herbes hautes de l’écozone des plaines à forêts mixtes (sud de l’Ontario) et qui n’est surpassé que dans les peuplements de graminées cespiteuses ou d’armoises de l’écozone du bassin intérieur de l’ouest (centre-sud de la Colombie-Britannique). Les prairies herbeuses abritent cependant une proportion plus élevée d’abeilles affichant une spécialisation alimentaire, la moitié étant des espèces oligolectiques strictement inféodées aux Asteraceae. Plus de 70 % des peuplements de prairies indigènes étaient déjà disparus en 1990. Il est donc probable que plusieurs des espèces endémiques oligolectiques de ces habitats soient devenues beaucoup plus rares, même si l’on dispose de peu de données de référence permettant une comparaison rigoureuse. Seule une infime partie de l’écozone des prairies a fait l’objet de recensements complets des abeilles, et ce chapitre présente plusieurs nouvelles mentions. Il propose en outre un examen taxonomique complet des abeilles des prairies herbeuses, accompagné de résumés bionomiques. Sheffield, C. S., S. D. Frier, and S. Dumesh. 2014. The Bees (Hymenoptera: Apoidea, Apiformes) of the Prairies Ecozone, with Comparisons to other Grasslands of Canada. In Arthropods of Canadian Grasslands (Volume 4): Biodiversity and Systematics Part 2. Edited by D. J. Giberson and H. A. Cárcamo. Biological Survey of Canada. pp. 427-467. © 2014 Biological Survey of Canada. ISBN 978-0-9689321-7-9 doi: http://dx.doi.org/10.3752/9780968932179.ch11 428 C. S. Sheffield, S. D. Frier, and S. Dumesh Introduction No previous review of the bee species occurring in Canada or any of its ecozones has been completed, though distributional data from taxonomic revisions provide information for some taxa (see taxonomic treatments listed below). Previous published estimates of the number of bee species in Canada range from 520 (Hurd 1979: catalogue) to over 970 (Danks 1979: general estimate). Sheffield et al. (2011a) reported 803 species and provided totals for each province and territory, but subsequent work verifying occurrence, new synonymies, and taxonomic clarification resulted in a more conservative tally of 797 confirmed species (CSS, unpublished data). This number represents about 4% of the current world total of just over 20,000 species (Ascher and Pickering 2014). Globally, bee diversity is highest in semi-arid environments (Michener 1979, 2007), and this pattern probably holds relatively well within Canada. However, we do not have a complete picture of the bees for Canada as a whole or within many of the country’s ecozones. Additional surveys, including application of genetic methods to reveal cryptic species (e.g., Gibbs 2009; Sheffield et al. 2009; Rehan and Sheffield 2011), should increase the confirmed number of bee species in Canada. Bees are dependent on pollen and nectar (and in rarer cases, floral oils). Therefore, they require habitats rich in food plants to complete their life cycle. Bees also require suitable nesting sites (see Table 1), and for some species, nesting materials (Westrich 1996). Therefore, grasslands of Canada should be ideal habitats for most bees, as grasslands are dominated by herbaceous vegetation and have some of the best soil for plant growth in the country (Federal, Provincial and Territorial Governments of Table 1. Review of the literature on bee nesting behaviour and ecology for some of the bee species occurring in the Prairies Ecozone. Family Reference and Bee Species COLLETIDAE Batra 1980: Colletes inequalis, C. thoracicus, C. validus Rajotte 1979: Colletes validus Torchio et al. 1988: Colletes kincaidi Fye 1965: Hylaeus annulatus (as H. ellipticus), H. basalis, H. verticalis Torchio 1984: Hylaeus leptocehpalus (as H. bisinuatus) ANDRENIDAE Batra 1990: Andrena alleghaniensis Johnson 1981: Andrena dunningi Miliczky 1988: Andrena erythrogaster, A. cressonii, Miliczky 2008: Andrena prunorum Miliczky and Osgood 1995: Andrena vicina, A. regularis Miliczky et al. 1990: Andrena nivalis Norden and Scarbrough 1979: Andrena miserabilis Rozen 1973: Andrena accepta Schrader and LaBerge 1978: Andrena carlini, A. regularis Miliczky 1991: Calliopsis coloradensis Eickwort 1977b: Perdita halictoides, P. octomaculata The Bees (Hymenoptera: Apoidea, Apiformes) of the Prairies Ecozone 429 with Comparisons to other Grasslands of Canada Family Reference and Bee Species HALICTIDAE Eickwort 1981: Agapostemon spp. LaBerge and Ribble 1966: Agapostemon sericeus (as L. radiatus), A. splendens Packer and Knerer 1986: Halictus ligatus Soucy 2002: Halictus rubicundus Yanega 1990; Halictus rubicundus Batra 1987: Lasioglossum laevissimum Bohart and Youssef 1976: Lasioglossum lusorium (as Evylaeus galpinsiae), L. abberans LaBerge and Isakson 1963: Lasioglossum zephyrum McGinley 2003: Lasioglossum subgenus Sphecodogastra s.s. Packer and Owen 1989: Lasioglossum cooleyi Packer et al. 1989a: Augochlora aurata, Lasioglossum cinctipes, L. laevissimum Packer 1992: Lasioglossum laevissimum Packer 1994: Lasioglossum tenax MELITTIDAE Rozen and Jacobson 1980: Macropis nuda Megachilidae Custer and Hicks 1927: Anthidiini Clement 1974: Dianthidium pudicum Michener and Michener 1999: Dianthidium curvatum Fye 1965: Hoplitis albifrons, H. spoliata, Osmia atriventris, O. proxima, Megachile inermis, M. relativa, M. gemula Eickwort et al. 1981: Megachile texana Hobbs 1954: Megachile in prairies Jenkins and Matthews 2004: Megachile frigida Neff and Simpson 1991: Megachile fortis Sheffield et al. 2011b: Megachile in Canada Cane et al. 2007: Osmia spp. review Cane 2012: Osmia integra Parker and Tepedino 1982: Osmia sculleni Rightmyer et al. 2013: Osmia spp. review APIDAE Hobbs et al. 1961; Anthophora occidentalis Torchio 1971: Anthophora peritomae Hobbs 1964, 1965a, 1965b, 1966a, 1966b, 1967, 1968: Bombus spp. Rau 1928: Ceratina calcarata Rehan and Richards 2010: Ceratina calcarata Vickruck et al. 2011: Ceratina calcarata, C. dupla Eickwort 1977a: Diadasia diminuta Linsley and MacSwain 1957: Diadasia spp. Linsley et al. 1963: Eucera speciosa Miliczky 1985: Eucera hamata Linsley 1943: brief account of Habropoda cineraria Cameron et al. 1996: Melissodes druriella (as M. rustica) Clement 1973: Melissodes druriella (as M. rustica) Miliczky 2000: Melissodes microsticta Parker et al. 1981: Melissodes agilis 430 C. S. Sheffield, S. D. Frier, and S. Dumesh Canada 2010). These conditions have also resulted in these habitats being among the most heavily modified in the country (Scoggan 1978). Most grasslands in the three Prairie Provinces were converted to cropland prior to the 1930s (Federal, Provincial and Territorial Governments of Canada 2010), a phenomenon continuing today (Javorek and Grant 2011). Little of the original natural cover of the Canadian Prairies remains intact, and this region is now primarily an agricultural system (Riley et al. 2007; Shorthouse 2010). Globally, grassland systems are the most altered, and thus threatened, of all ecosystem types, yet they provide some of the most important ecological services (reviewed by Federal, Provincial and Territorial Governments of Canada 2010) and serve as regional and global “breadbaskets” for food production. Despite the key pollination