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ARTICLE
A checklist of the bees (Hymenoptera: Apoidea) of
Manitoba, Canada
Jason Gibbs1* , Emily Hanuschuk1
,
2†, Reid Miller1, Melanie Dubois2, Massimo Martini1xx,
Steve Robinson1, Phoenix Nakagawa1‡, Cory S. Sheffield3, Sophie Cardinal4, and
Thomas M. Onuferko5
1Department of Entomology, University of Manitoba, 12 Dafoe Road, Winnipeg, Manitoba, R3T 2N2, Canada, 2Brandon
Research and Development Centre, Agriculture and Agri-Food Canada, 2701 Grand Valley Road, Brandon, Manitoba,
R7C 1A1, Canada, 3Royal Saskatchewan Museum, 445 Albert Street, Regina, Saskatchewan, S4P 4W7, Canada, 4Canadian
National Collection of Insects, Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue, Ottawa,
Ontario, K1A 0C6, Canada, and 5Beaty Centre for Species Discovery, Canadian Museum of Nature, P.O. Box 3443,
Station D, Ottawa, Ontario, K1P 6P4, Canada
*Corresponding author. Email: jason.gibbs@umanitoba.ca
(Received 19 July 2022; accepted 11 October 2022)
Abstract
We record 392 species or morphospecies of bees (Hymenoptera: Apoidea) for Manitoba, Canada, which is
154 more species than reported in 2015 and includes five new generic records since 2015 (Ashmeadiella,
Brachymelecta,Eucera, Neolarra, and Triepeolus). Thirteen new records reported here are new for Canada:
Calliopsis (Nomadopsis)australior Cockerell, Perdita (Perdita)tridentata Stevens, Brachymelecta
interrupta (Cresson), Diadasia (Dasiapis)ochracea (Cockerell), Melissodes bidentis Cockerell, Nomada
crawfordi crawfordi Cockerell, Nomada fuscicincta Swenk, Nomada sphaerogaster Cockerell, Nomada
xantholepis Cockerell, Triepeolus cf. grindeliae Cockerell, Dianthidium (Dianthidium)parvum
(Cresson), Coelioxys (Xerocoelioxys)nodis Baker, and Megachile (Megachiloides)dakotensis Mitchell.
We remove the following species from the list of Manitoba bees based on re-examination of voucher
material: Andrena (Ptilandrena)geranii Robertson, Andrena (Rhacandrena)robertsonii Dalla Torre,
Andrena (Simandrena)nasonii Robertson, Andrena (Trachandrena)ceanothi Viereck, Andrena
(Trachandrena)quintilis Robertson, Lasioglossum (Hemihalictus)pectoraloides (Cockerell),
Lasioglossum (Lasioglossum)forbesii (Robertson), and Dianthidium (Dianthidium)concinnum
(Cresson). We propose that Nomada alpha paralpha Cockerell, 1921 and N. alpha dialpha Cockerell,
1921 are junior synonyms of N. alpha Cockerell, 1905. Nomada arenicola Swenk, 1912 is considered a
junior synonym of N. fervida Smith, 1854. Protandrena albertensis (Cockerell) and Neolarra mallochi
Michener are recognised as valid species. We provide additional notes on taxonomy, nomenclature,
and behaviour for select species in the list.
Subject editor: Jeremy deWaard
†Current address: Canadian Grain Commission, 303 Main Street, Winnipeg, Manitoba, R3C 3G8, Canada
‡Current address: Department of Soil Science, University of Manitoba, 13 Freedman Crescent, Winnipeg, R3T 2N2,
Manitoba, Canada
XXCurrent address: Nature Conservation and Landscape Ecology, University of Freiburg, Tennenbacher Str. 4, 79106
Freiburg, Germany
© The authors and His Majesty, the King, in right of Canada, as represented by the Minister of Agriculture and Agri-Food Canada, 2023.
Published by Cambridge University Press on behalf of The Entomological Society of Canada. This is an Open Access article, distributed
under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted
re-use, distribution and reproduction, provided the original article is properly cited.
The Canadian Entomologist (2023), 155, e3, 1–38
doi:10.4039/tce.2022.45
Published on
behalf of the
Entomological
Society of
Canada
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
Introduction
Bees (Hymenoptera: Apoidea: Anthophila) are a monophyletic group within the
hymenopteran superfamily Apoidea (Melo 1999; Danforth et al. 2006; Debevec et al. 2012;
Hedtke et al. 2013; Branstetter et al. 2017). Seven families of bees are currently recognised
(Michener 2007), with six occurring in Canada: Andrenidae, Apidae, Colletidae, Halictidae,
Megachilidae, and Melittidae. Some authors prefer to recognise a single bee family (Moure
et al. 2007), but this practice is not widely accepted. Globally, more than 20 500 bee species
have been described (Ascher and Pickering 2022), and more than 900 bee species have been
documented in Canada (Sheffield et al. 2017; Sheffield 2019). Bees in Canada are divided into
52 genera, although generic limits differ among authors (Mitchell 1960,1962; Michener 2007;
Dorchin et al. 2018; Ascher and Pickering 2022). Bees display a range of anatomical and
behavioural adaptations that contribute to their biodiversity and aesthetic quality.
Most bees are solitary and live in underground burrows or pre-existing cavities
(Krombein 1967; Cane et al. 2007; Michener 2007; Cane and Neff 2011; Danforth et al. 2019).
Less commonly, nests are excavated in wood and constructed externally on substrates or
under rocks. Honey bees (Apis mellifera Linnaeus) and bumble bees (genus Bombus) are
unusual for making wax structures in larger hollows in trees or, often the case among bumble
bees, abandoned rodent burrows (Michener 1974; Plowright and Laverty 1984). Social
behaviour in Apis and Bombus is well known, but it also commonly occurs in the Halictidae.
Social behaviour in halictids is much more plastic and variable than in Apis or Bombus
(Michener 1974; Packer et al. 1989; Eickwort et al. 1996; Field 1996; Mueller 1996; Richards
et al. 2003; Gibbs et al. 2012b).
Bees are known for their close connection with flowers, which extends back to the early
Cretaceous, approximately 123 Ma (Cardinal and Danforth 2013). Bees typically harvest
pollen and nectar from flowers to provision their nests (Portman et al. 2019). Flowering
plants can exploit this behaviour for pollination. Some bees are specialist floral visitors
(oligoleges), using only a small subset of available plants (Robertson 1926; Wcislo and
Cane 1996; Cane and Sipes 2006). Polylectic bees use a wide range of flowers, a trait that may
have aided the early diversification of bees (Murray et al. 2018). Many bees –approximately
13% –do not collect pollen but act as brood parasites in other bee nests (Michener 2007;
Danforth et al. 2019). This strategy has originated multiple times in bees (Smith et al. 2007;
Cardinal et al. 2010; Gibbs et al. 2012a; Litman et al. 2013). In general, bees are considered
among the most important animal pollinators (Kevan and Baker 1983; Ollerton et al. 2011).
Over the last two decades, there has been substantial interest in the status of pollinators (Allen-
Wardell et al. 1998; Kevan and Phillips 2001; Marlin and LaBerge 2001; Biesmeijer et al. 2006;
Berenbaum et al. 2007; Potts et al. 2010; Winfree 2010; Colla et al. 2012; Bartomeus
et al. 2013; Lebuhn et al. 2013; Senapathi et al. 2015). With the possible exception of bumble
bees (Cameron et al. 2011; Kerr et al. 2015), few wild bee taxa have been sufficiently well
documented in North America to provide effective baseline data to reliably measure
conservation status. Museum data can provide some insight into historical trends, but
analysing these data can be challenging due to the inconsistent and usually unknown
sampling methods applied over time (Bartomeus et al. 2013,2019). Statistical modelling can
provide predictions of how land-use changes may affect bees (Koh et al. 2016), but these are a
poor replacement for empirical studies of actual bees. Several published and unpublished
checklists have become available for states and provinces in recent years for North America
(Donovall and VanEngelsdorp 2010; Jean 2010; Scott et al. 2011; Canadian Endangered
Species Conservation Council 2015; Dibble et al. 2017; Gibbs et al. 2017; Kilpatrick
et al. 2020). A consistent trend that emerges in these studies is how limited basic inventories
or checklists of bees are for most of the continent (Jamieson et al. 2019). Without these
baseline data, efforts to monitor trends in bees are quixotic (Tepedino et al. 2015).
2 Gibbs et al.
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
Historical bee collection in Manitoba, Canada
Entomological research in Manitoba, Canada was strongly influenced by Norman Criddle
(1875–1933). Criddle and his siblings were avid collectors and natural historians
(Criddle 1975). The homestead near Treesbank on which he was raised, referred to on
collection labels as Aweme, and now the Criddle/Vane Homestead Provincial Park, is an
important historical site for entomology (Roughley 2000). Aweme as a locality may be a much
broader area than just the Criddle homestead, based on the breadth of species, including
habitat specialists, labelled with this location. In 1913, Criddle was employed by the Division
of Entomology of the Canadian Dominion government’s Experimental Farms Branch (then a
division of the Dominion of Canada’s Department of Agriculture; now under Agriculture and
Agri-Food Canada) as an entomological field officer for Manitoba and, beginning in 1919, as
an entomologist (Gibson and Crawford 1933). Criddle began the first federal entomology lab
at Aweme in 1915, which was the centre for early entomological research in the province. The
Criddle home, known as St. Albans, was a meeting spot for Criddle’s friends and colleagues.
Although Criddle’s professional work was focused on crop protection, particularly against
grasshoppers, many of our earliest records of bees and many other insects for Manitoba stem
from his collections (Gibson 1914,1915,1916,1917; Gibson and Criddle 1920), including
specimens used in the description of new bees (Sladen 1916a).
Ralph Durham Bird (1901–1972), a native Manitoban, worked at Aweme under the direction
of Criddle from 1924 to 1926. He left Manitoba for several years but returned in 1933 to head the
Federal Entomology Laboratory following Criddle’s death (Bird 1975). The laboratory at that time
moved to Brandon, Manitoba. He came to reside in Winnipeg, Manitoba with the founding of the
Dominion’s Department of Agriculture Research Station to head the entomology section and later
its crop protection section. Bird’s research career, like Criddle’s, focused primarily on agricultural
pests, but he collected several early bee records for the province.
John Braithwaite Wallis (1876–1962) was a friend and colleague of R.D. Bird and the Criddle
family. Wallis’s interests were primarily in other insect orders: he wrote a monograph on tiger
beetles (Wallis 1961). Wallis was hired by the Department of Entomology of the Manitoba
Agricultural College to build an insect collection. In 1983, the collection was renamed the
J.B. Wallis Museum of Entomology in his honour (Galloway et al. 2010). The department
itself was founded in about 1920 by Alvin Valentine Mitchener (1888–1962), who was the
first entomologist at the University of Manitoba (then the Manitoba Agriculture College),
where he worked until 1954. Mitchener’s work with bees was largely limited to honey bees,
although his long-term data on pollen and nectar plants are of value to those interested in
wild bees (Mitchener 1948).
Bee collections in the province were largely haphazard in the first half of the 20th century. One
exception is a study of the province’s bumble bees by Felix Neave (1901–1986), which recognised
23 species (Neave 1933). Alexander Jardine Hunter (1868–1940) was a medical doctor and
missionary in Teulon, Manitoba (Mitchell 1940). His collections provide an early record of
the Interlake fauna between Lake Manitoba and Lake Winnipeg. Interest in native pollinators
began to emerge in the 1940s and 1950s out of the Field Crop Insect Laboratory in Brandon,
based on specimen records and publications (Cole 1955; Stephen 1955). Several scientists at
Brandon collected bees at least occasionally, including R.D. Bird (above), Walter Askew
(1929–2000), and Clifford Francis Barrett (1925–2017). William Procuronoff Stephen
(1927–2016) and Thomas Victory Cole (1918–1999), of the Brandon lab, both conducted
research on alfalfa pollination (Cole 1955,1957; Stephen 1955; Bird 1963). Stephen, although
a native Manitoban, spent little of his professional career in the province, working at Brandon
only from 1947 to 1952 before taking a position at Oregon State University, Corvallis,
Oregon, United States of America (Bird 1963). Nevertheless, his taxonomic revision of Colletes
had a lasting impact on North American melittology and included numerous records from the
The Canadian Entomologist 3
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province (Stephen 1954). His subsequent introduction of blue vane traps for bee collecting
(Stephen and Rao 2005) has also changed the way collectors survey bees globally. Beginning
in 1953, Cole led research on insect pollination of alfalfa at the field station in Wanless,
Manitoba, which resulted in long series of bees, especially Bombus and Megachile. Cole
studied leafcutters and created artificial nests for B. terricola Kirby (Cole 1957; Bird 1963). He
also developed methods, which were decades ahead of their time, for habitat management to
improve pollinators in areas adjacent to field crops (Cole 1955). Pollinator research during
this time also included a focus on sunflower crops, Helianthus annuus Linnaeus (Compositae)
(Barrett 1955), which resulted in a series of Protandrena from Altona, Manitoba. In the 1950s,
Arthur Robinson Brooks (1917–1962) and Leonard Alexander Kelton (born Konotopetz;
1923–2011) collected hundreds of bee records while surveying insects of the Prairie Provinces.
Brooks had been a dipterist and Kelton focused on Hemiptera, but Brooks later undertook
studies on numerous prairie insects (Riegert 1990; Henry and Gill 2016). Even though neither
researcher was a hymenopterist, Brooks collected the only known specimens of Holcopasites
stevensi Crawford from the province. In 1961, Herbert Edward Milliron (1923–1981) made a
significant collection of bumble bees from the province, many of which are deposited at the
Canadian National Collection, Ottawa, Ontario, Canada. He subsequently published a series
on the bumble bees of the Western Hemisphere (Milliron 1971,1973a, 1973b). Most bee
research in Manitoba focused on two managed exotic species –the European honey bee and
the alfalfa leafcutter bee (Megachile rotundata Fabricius). Honey bee research in the
Department of Entomology, University of Manitoba, was led during this time by Stanley
Cameron Jay (Holliday and Currie 2009). One of his students, Robert Christopher Plowright,
completed his doctorate on Bombus domestication and caste differentiation (Plowright 1966;
Plowright and Jay 1966,1968). Applied studies of pollinators, although valuable, have not
always provided much information on the broader bee fauna of the province.
General collections of bees in Manitoba were few during the 1960s, but in 1977, the
Department of Entomology, University of Manitoba, hired Terry D. Galloway. Although
Galloway’s research was largely in veterinary entomology, he demonstrated a healthy interest
in wild bees in the mid-1970s through the 1980s, based on more than 2000 specimen records.
During the mid-1980s, William James Turnock (1929–2008) collected bees, although his
research focus was on pest control. Turnock et al. (2006) published records of 15 bumble bee
species captured inadvertently in baited traps for bertha armyworm, Mamestra configurata
Walker (Noctuidae), in Manitoba canola fields. Also during this period, David Harvey
Pengelly (1922–2004) was active in the province, primarily in the area of Erickson where he
retired after a career as professor at the Ontario Agricultural College, University of Guelph
(Guelph, Ontario, Canada; Marshall 2004). Pengelly’s interest was primarily in the genus
Megachile, which was the basis of his doctoral studies at Cornell University (Ithaca, New
York, United States of America; Pengelly 1955). During his time, Pengelly inspired several
young entomologists, including a future curator of the J.B. Wallis Museum of Entomology,
Robert Edward Roughley (1950–2009). The museum expanded greatly during Roughley’s
tenure as curator. Roughley was an expert in aquatic beetles, but he also studied the fauna of
grassland insects and conducted surveys at Aweme. He participated in a nationwide project to
study Canadian pollinators, CANPOLIN (Galloway et al. 2010), which resulted in thousands
of specimen records (Patenaude 2007). In 2011, the museum was rededicated as the
J.B. Wallis/R.E. Roughley Museum of Entomology.
Bee collecting in Manitoba in the 2000s
A large survey of prairie bees was undertaken by one of Roughley’s students, Andrea
Patenaude. She collected extensively during her thesis work at Spruce Woods Provincial Park
(Patenaude 2007). Patenaude’s specimens were included in taxonomic studies of the genus
4 Gibbs et al.
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
Lasioglossum (Gibbs 2010). During CANPOLIN, bees were collected at Canadian Airforces Base
Shilo neighbouring Spruce Woods Provincial Park. Several of these bees were sequenced as part of
an effort to DNA barcode Canadian bees (Sheffield et al. 2017). More recent theses on wild bees in
grasslands were conducted by Sarah Semmler and Reid Miller in the Tall Grass Prairie Preserve
(Semmler 2015; Miller 2021), by Marika Olynyk in fragmented grasslands of southwestern
Manitoba (Olynyk 2017; Olynyk et al. 2021), and by Emily Hanuschuk across multiple
landscape in southern Manitoba (Hanuschuk 2021), resulting in more than 30 000 specimen
records. These studies include novel records for the province, including the first provincial
records of the genus Dianthidium (Semmler et al. 2018). Other surveys of wild bee diversity
across multiple habitat types in southern Manitoba, including thesis research by Massimo
Martini (2022), have contributed to the provincial records reported herein. In a survey of
prairie bees in Canada, Sheffield et al. (2014) listed 218 species for the prairie region of
Manitoba. A slightly larger list of 236 Manitoba bees was made available through a recent
report on the status of species in Canada (Canadian Endangered Species Conservation
Council 2015), and an online checklist of approximately 250 species was released by
Sheffield (2019).
Relatively few of the studies on wild bees in Manitoba have resulted in peer-reviewed
publications (but see Neave 1933; Turnock et al. 2006; Semmler et al. 2018; Robson
et al. 2019; Gibbs et al. 2021; Olynyk et al. 2021). In 2017, research on wild bees at the
University of Manitoba, Winnipeg, began in earnest, the results of which included the
discovery of many new provincial records (Semmler et al. 2018; Gardner and Gibbs 2021;
Onuferko et al. 2021; Satyshur et al. 2021; Wrigley et al. 2021). Among these were new
national records and new species, as well as erroneous published records based on
misidentified material. Hence, the purpose of the current study is to present these discoveries
and provide a more accurate and complete checklist of the bees of Manitoba, which has
changed considerably and is therefore warranted. This checklist is intended to stimulate
renewed interest and support research on the Manitoba bee fauna and other areas of the
Prairie Provinces and the north–central region of the United States of America. Due to the
need for taxonomic revisions in several key bee groups (e.g.,Nomada and Sphecodes), this
checklist is necessarily preliminary.
Methods
This study is restricted to the geographic boundaries of the Province of Manitoba, Canada.
Manitoba is situated between the provinces of Saskatchewan to the west and of Ontario to the
east, the territory of Nunavut to the north, and the states of North Dakota and Minnesota,
United States of America, to the south. The Manitoban climate has extreme seasonality, with
temperatures in the south ranging from –40 °C to 38 °C between winter and summer. There
are six broad ecozones (Smith et al. 1998). The Taiga Shield, Southern Arctic, and the
Hudson Plains occur in the north. The town of Churchill occurs near the northern limit of
the Hudson Plains. The middle latitudes of the province to the southeastern corner are of the
Boreal Shield Ecozone. The Boreal Plains Ecozone occurs to the southwest of the Boreal
Shield. The southwestern part of the province is largely Prairies Ecozone, significant portions
of which have been converted to large-scale agriculture.
An initial checklist was compiled using taxonomic literature (Mitchell 1935a, 1935b;
Sandhouse 1939; Stephen 1954; Hurd and Michener 1955; Timberlake 1960;
LaBerge 1961,1967,1971,1973,1977,1980,1986,1987,1989; Ordway 1966;
Ribble 1967,1968; Shinn 1967; Milliron 1971,1973a, 1973b; LaBerge and Ribble 1972,1975;
Daly 1973; Donovan 1977; Bouseman and LaBerge 1979; McGinley 1986,2003;
The Canadian Entomologist 5
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Broemeling 1988; Rightmyer 2008; Gibbs 2010,2011; Rightmyer et al. 2010; Sheffield et al. 2011;
Gibbs et al. 2013,2017; Williams et al. 2014; Onuferko 2017,2018) and ecological studies
(Cole 1955,1957; Stephen 1955; Plowright 1966; Patenaude 2007; Olynyk 2017; Robson
et al. 2019). Published records were confirmed, where possible, by examining material in
relevant collections, particularly the J.B. Wallis/R.E. Roughley Museum of Entomology and the
Canadian National Collection of Insects, Arachnids, and Nematodes (records digitised at
https://www.cnc.agr.gc.ca/taxonomy/TaxonMain.php). The historical bee collections at the
J.B. Wallis/R.E. Roughley Museum were re-examined and databased to be deposited on
Canadensys (https://www.canadensys.net) and the University of Manitoba Dataverse
(https://doi.org/10.34990/FK2/55PV3G). Material was examined from the Brandon Research
and Development Centre, Agriculture and Agri-Food Canada, Brandon, the Canadian
Museum of Nature, and the Illinois Natural History Survey, Prairie Research Institute,
Champaign, Illinois, United States of America. Additional records come from other
well-digitised collections, such as the American Museum of Natural History, New York,
New York, United States of America. New collections were made as part of ongoing research
projects and to fill gaps in our knowledge of Manitoban bees. These included records from
student thesis projects (Emily Hanuschuk, Reid Miller, and Massimo Martini) and samples
from the Manitoba Conservation Data Centre, Winnipeg, Manitoba. Some effort was made to
collect potential species with specialised floral associations that were known from
neighbouring jurisdictions. For example, specialist bees of Amorpha (Fabaceae) are known to
occur in Minnesota, so targeted collection from this host plant took place. We also checked
iNaturalist (www.iNaturalist.org) and Bumble Bee Watch (www.bumblebeewatch.org) and
provide the unique code for observations supporting new records for the province.
Classification largely follows Michener (2007), except that we recognise subfamilies of Apidae
supported by recent phylogenetic studies (Cardinal et al. 2010; Bossert et al. 2019), and subgenera
of Andrena and Lasioglossum follow recent studies (Gibbs et al. 2012b, 2013; Pisanty et al. 2022).
Michener (2000,2007) recognised Pterosarus and Heterosarus as subgenera of Protandrena, but
they have also been treated as subgenera or synonyms of Pseudopanurgus (Timberlake 1967;
Ascher 2004; Ascher and Pickering 2020). Phylogenetic studies seem to suggest that a broadly
defined Pseudopanurgus is paraphyletic (Bossert et al. 2021; Ramos et al. 2022). No
Pseudopanurgus sensu stricto occur in Manitoba, but we implicitly use Protandrena as an
umbrella genus for all North American Protandrenini. The following literature was used to
identify specimens and determine taxon concepts: Andrena Fabricius: Mitchell (1960);
LaBerge (1967,1969,1973,1977,1980,1986,1989); Ribble (1967,1968,1974); LaBerge and
Bouseman (1970); LaBerge and Ribble (1972,1975); Bouseman and LaBerge (1979); Calliopsis
Smith: Rozen (1958); Mitchell (1960); Shinn (1967); Perdita Smith: Timberlake
(1954,1958,1960,1968); Mitchell (1960); Protandrena Cockerell: Mitchell (1960);
Timberlake (1967,1973,1975); Scott et al. (2011); Bombus Latreille:
Milliron (1971,1973a, 1973b); Laverty and Harder (1988); Williams et al. (2008,2014,2019);
Ghisbain et al. (2020); Brachymelecta Linsley: Hurd and Linsley (1951); Mitchell (1962);
Onuferko et al. (2021); Diadasia Patton: Timberlake (1941); Adlakha (1969); Snelling (1994);
Eucera: Timberlake (1969); Melissodes Latreille: LaBerge (1956a, 1956b, 1961);
Mitchell (1962); Holcopasites Ashmead: Hurd and Linsley (1972); Epeolus Latreille:
Brumley (1965); Onuferko (2017,2018); Triepeolus Robertson: Rightmyer (2008); Neolarra
Ashmead: Michener (1939a); Shanks (1977); Nomada Scopoli: Cockerell
(1903,1905a, 1905b, 1908); Mitchell (1962); Broemeling (1988); Broemeling and
Moalif (1988); Alexander and Schwarz (1994); Schwarz and Gusenleitner (2004); Droege et al.
(2010); Epeoloides Giraud: Mitchell (1962); Packer et al. (2007); Ceratina Latreille:
Mitchell (1962); Daly (1973); Rehan and Richards (2008); Rehan and Sheffield (2011);
6 Gibbs et al.
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Colletes Latreille: Stephen (1954); Mitchell (1960); Hylaeus Fabricius: Mitchell (1960);
Snelling (1966,1968,1970); Oram (2018); Augochlorella Sandhouse: Mitchell (1960);
Ordway (1966); Coelho (2004); Lasioglossum Curtis: McGinley (1986,2003);
Gibbs (2010,2011); Gibbs et al. (2013); Sphecodes Latreille: Mitchell (1960); M. Arduser
(unpublished data); Dufourea Lepeletier: Dumesh and Sheffield (2012); Gibbs et al. (2014);
Anthidium Fabricius: Gonzalez and Griswold (2013); Dianthidium Cockerell:
Mitchell (1962); Grigarick and Stange (1968); Stelis Panzer: Cockerell (1898); Sladen (1916b);
Popov (1938); Mitchell (1962); Coelioxys Latreille: Mitchell (1962,1980); Baker (1975);
de Silva (2012); da Rocha Filho and Packer (2016); Megachile Latreille:
Mitchell (1934,1935a, 1935b, 1936,1937a, 1937b, 1962); Parker (1978); Sheffield et al. (2011);
Byzdk (2012); Ashmeadiella Cockerell: Michener (1939b); Hurd and Michener (1955);
Mitchell (1962); Rowe (2017); Hoplitis Klug: Michener (1947); Mitchell (1962); Rowe (2017);
Osmia Panzer: Sandhouse (1939); White (1952); Mitchell (1962); Rightmyer et al. (2010);
and Macropis Panzer: Mitchell (1960); Michez and Patiny (2005). Some identifications were
confirmed by colleagues with expertise in the relevant taxon. Zach Portman, University of
Minnesota, confirmed the identification of Perdita fallax Cockerell from photographs.
Karen Wright, Texas A&M University, College Station, Texas, United States of America,
identified a synoptic set of Melissodes for Manitoba, which improved our taxon concepts.
Terry Griswold, United States Department of Agriculture, Agriculture Research Service,
verified and corrected identifications of a synoptic set of Stelis.
The checklist (Table 1) is ordered alphabetically by family, subfamily, tribe, genus, subgenus
(where applicable), and species. Some species are flagged as provisional when plausible literature
records could not be confirmed by examining voucher specimens or uncertain if the identification
is doubtful. Material examined is provided for new Canadian and Manitoban records in
Supplementary material S1. In some cases, “new”records are difficult to define because
specimens may be recorded outside of the traditional scientific literature, either in theses
(Patenaude 2007; Semmler 2015; Olynyk 2017) or within online databases. In addition, the
collaborative nature of our research has meant that some bees first identified as part of this
study, including new generic records for the province and new Canadian records, were
released early to benefit other studies (Gardner and Gibbs 2021; Onuferko et al.2021;
Satyshur et al. 2021; Wrigley et al. 2021) or were being worked on simultaneously as part of
graduate theses (Hanuschuk 2021; Miller 2021; Martini 2022). Our goal is to provide
accessible, verifiable data for interesting records, even if there may be an earlier record outside
of peer-reviewed publications. For this reason, we list as new records any species recorded
during our research since the recent publication of a checklist of Canadian bees (Canadian
Endangered Species Conservation Council 2015) but acknowledge their occurrence in other
data sources. When many distinct collectioneventsexistforanewspecies,only
municipalities are provided, followed by the number of examined specimens. Intertegular
spans (Cane 1987) for common species in southern Manitoba are provided for reference
(Supplementary material S2). Measurements were taken using an ocular micrometer or a
microscope-mounted camera and calculated using NIS-Elements (Nikon Instruments Inc.,
Melville, New York, United States of America). Historical records were georeferenced using
online gazetteers and Google Earth (https://earth.google.com). The following abbreviations are
used below in material examined sections: CFB, Canadian Forces Base; PF, provincial forest;
PP, provincial park; and WMA, wildlife management area. The abbreviations for collections
referred to below are as follows: AAFC, Brandon Research and Development Centre,
Agriculture and Agri-Food Canada; AMNH, American Museum of Natural History;
CMNC, Canadian Museum of Nature; CNC, Canadian National Collection of Insects,
Arachnids, and Nematodes; INHS, Illinois Natural History Survey; MCDC, Manitoba
Conservation Data Centre; and WRME, J.B. Wallis/R.E. Roughley Museum of Entomology.
The Canadian Entomologist 7
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Table 1. Checklist of the bees of Manitoba. Data sources are indicated by superscripts (see Methods). New provincial
records are indicated in bold text. New Canadian records are preceded by an asterisk. Exotic species are indicated
using a superscript “E”. The following abbreviations are used for behavioural data: sol., solitary; comm., communal;
sub., subsocial; eus., eusocial; par., cleptoparasitic or socially parasitic; G, ground; W, wood; S, stems; H, hives; E, exterior
surfaces; C, cavities; oligo., oliglolectic; and poly., polylectic. No lecty information is provided for parasites, although they
may have limited pollen use by virtue of their hosts. The host column is used for plant taxa for oligolectic bees and bee taxa
for brood parasites. It is likely that some oligolectic bees have host preferences for related taxa not included in the list.
Similarly, records of a single bee host do not preclude additional undocumented hosts for cleptoparasites. Data sources are
as follows: 1primary literature, including taxonomic revisions and other studies found in peer-reviewed journals; 2WRME
(J.B. Wallis/R.E. Roughley Museum of Entomology) material –specimen confirmed; 3unpublished theses and secondary
sources (e.g., nonpeer-reviewed literature). These are primarily theses (Patenaude 2007; Semmler 2015; Olynyk 2017)or
country-wide checklists (Canadian Endangered Species Conservation Council 2015; Sheffield 2019)–only recorded when it
does not occur in the primary literature; 4AAFC (Agriculture and Agri-Food Canada) Brandon material –specimen
confirmed; 5CNC (Canadian National Collection) material –specimen confirmed; 6CMNC (Canadian Museum of Nature
collection) material –specimen confirmed; 7recorded in a public database and housed at an external institution –
specimen not examined; 8images available on iNaturalist; and 9INHS (Illinois Natural History Survey) material –specimen
confirmed. ? indicates uncertainty.
Species Nests Substrate Lecty Floral or bee host
Andrenidae
Andreninae
Andrenini
Andrena (Andrena)clarkella (Kirby, 1802)1,2,5,8 sol. G oligo. Salix
Andrena (Andrena)frigida Smith, 18531,2,5,8,9 sol. G oligo. Salix
Andrena (Andrena)milwaukeensis
Graenicher, 19031,2,5,8
sol. G poly.
Andrena (Andrena)rufosignata Cockerell, 19021,2 sol. G poly.
Andrena (Andrena)thaspii Graenicher, 19031,2,5,9 sol. G poly.
Andrena (Callandrena s.l.) asteris Robertson, 1891
(simplex/solidiginis group)1,2,5,8
sol. G oligo. Solidago, Symphyotrichum
Andrena (Callandrena s.l.) helianthi Robertson, 1891
(helianthi group)1,2,5,8
sol. G oligo. Helianthus
Andrena (Callandrena s.l.) placata Mitchell, 1960
(simplex/solidiginis group)2
sol. G oligo. Solidago, Symphyotrichum
Andrena (Cnemidandrena)canadensis
Dalla Torre, 1896 (nubecula group)1,2,5
sol. G oligo. Solidago, Symphyotrichum
Andrena (Cnemidandrena)chromotricha
Cockerell, 1899 (chromotricha group)1,2,9
sol. G oligo. Compositae
Andrena (Cnemidandrena)hirticincta
Provancher, 1888 (hirticincta group)1,2
sol. G oligo. Solidago, Symphyotrichum
Andrena (Cnemidandrena)nubecula Smith, 1853
(nubecula group)1,2
sol. G oligo. Solidago, Symphyotrichum
Andrena (Cnemidandrena)parnassiae
Cockerell, 1902 (scutellinitens group)2
sol. G oligo. Parnassia
Andrena (Cnemidandrena)peckhami Cockerell, 1902
(chromotricha group)1
sol. G poly. Compositae
Andrena (Cnemidandrena)robervalensis
Mitchell, 1960 (scutellinitens group)1
sol. G poly. likely Compositae
Andrena (Cnemidandrena)runcinatae
Cockerell, 1906 (scutellinitens group)2,5
sol. G poly. likely Compositae
(Continued)
8 Gibbs et al.
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Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Andrena (Conandrena)bradleyi Viereck, 19072sol. G oligo. Ericaceae
Andrena (Gonandrena)persimulata Viereck, 19171,2,5 sol. G oligo. Cornus
Andrena (Holandrena)cressonii cressonii
Robertson, 18911,2
sol. G poly.
Andrena (Larandrena)miserabilis Cresson, 18721,2,3,5 sol. G poly.
Andrena (Leucandrena)barbilabris (Kirby, 1802)1,2,3.5,6 sol. G poly.
Andrena (Melandrena)carlini Cockerell, 19011,2,3,5 sol. G poly.
Andrena (Melandrena)commoda Smith, 18791,2 sol. G poly.
Andrena (Melandrena)dunningi Cockerell, 18981,2,5,8 sol. G poly.
Andrena (Melandrena)erythrogaster
(Ashmead, 1890)1,2,5,8,9
sol. G oligo. Salix
Andrena (Melandrena)lupinorum Cockerell, 19061,2,3,5 sol. G oligo. Fabaceae
Andrena (Melandrena)nivalis Smith, 18531,2,3,5 sol. G poly.
Andrena (Melandrena)regularis Malloch, 19171,2,3,5 sol. G poly.
Andrena (Melandrena)vicina Smith, 18531,2,3,5 sol. G poly.
Andrena (Micrandrena)illinoiensis Robertson, 1891
(illinoiensis group)1,2,5
sol. G poly.
Andrena (Micrandrena)melanochroa Cockerell, 1898
(piperi group)1,2,5
sol. G oligo. Rosaceae
Andrena (Micrandrena)nigrae Robertson, 1905
(illinoiensis group)2
sol. G oligo. Salix
Andrena (Micrandrena)salictaria Robertson, 1905
(illinoiensis group)1,2,3,5
sol. G oligo. Salix
Andrena (Micrandrena)ziziae Robertson, 1891
(piperi group)1,2,5,9
sol. G oligo. Zizia
Andrena (Parandrena)andrenoides
(Cresson, 1878)1,2,5
sol. G oligo. Salix
Andrena (Parandrena)wellesleyana
(Robertson, 1897)1,2,5,8
sol. G oligo. Salix
Andrena (Plastandrena)crataegi Robertson, 1893
(crataegi group)2,5
sol. G poly.
Andrena (Plastandrena)prunorum prunorum
Cockerell, 1896 (prunorum group)1,2,5,6
sol. G poly.
Andrena (Ptilandrena)algida Smith, 18531,2,3,5,9 sol. G poly.
Andrena (Ptilandrena)cf.campanulae
Viereck and Cockerell, 19142
sol. G oligo.
(?)
Andrena (Ptilandrena)nigrihirta (Ashmead, 1890)1,2,5,9 sol. G poly.
Andrena (Ptilandrena) aff. nigrihirta2,3 sol. G
Andrena (Scrapteropsis)alleghaniensis Viereck, 1907
(alleghaniensis group)1,2,8
sol. G poly.
Andrena (Scrapteropsis)imitatrix Cresson, 1872
(imitatrix group)1,2,5,9
sol. G poly.
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
Andrena (Simandrena)wheeleri Graenicher, 19041,2,5 sol. G poly.
Andrena (Taeniandrena)wilkella (Kirby, 1802)E,2,5,8 sol. G oligo. Fabaceae
Andrena (Thysandrena)medionitens
Cockerell, 19021,2,5
sol. G poly.
Andrena (Thysandrena)w-scripta Viereck, 19041,2,5 sol. G poly.
Andrena (Trachandrena)cyanophila
Cockerell, 1906 2,3
sol. G poly.
Andrena (Trachandrena)forbesii Robertson, 18911,2,5 sol. G poly.
Andrena (Trachandrena)hippotes
Robertson, 18951,2,5,8
sol. G poly.
Andrena (Trachandrena)mariae
Robertson, 18911,2,5,9
sol. G oligo. Salix
Andrena (Trachandrena)miranda Smith, 18791,2,5 sol. G poly.
Andrena (Trachandrena)sigmundi
Cockerell, 19021,2,5,9
sol. G oligo. Salix
Panurginae
Calliopsini
Calliopsis (Calliopsima)coloradensis Cresson, 18781,2 sol. G oligo. Compositae
Calliopsis (Calliopsis)andreniformis
Smith, 18532,3,9
sol. G poly.
*Calliopsis (Nomadopsis)australior
Cockerell, 18972,3,4
sol. G unk.
Perditini
Perdita (Cockerellia)albipennis Cresson, 1868
canadensis Crawford, 19121,2,6,8
sol. G oligo. Helianthus
Perdita (Perdita)bruneri Cockerell, 1897
(octomaculata group)1,2
sol. G oligo. Compositae
Perdita (Perdita)fallax Cockerell, 1896
(octomaculata group)6
sol. G oligo. Helianthus
Perdita (Perdita)halictoides Smith, 1853
(halictoides group)1,2,5
sol. G oligo. Physalis
Perdita (Perdita)maculigera Cockerell, 1896
maculipennis Graenicher, 1910
(octomaculata group)1,2,9
sol. G oligo. Salix
Perdita (Perdita)octomaculata (Say, 1824)
(octomaculata group)1
sol. G oligo. Solidago
Perdita (Perdita)perpallida perpallida
Cockerell, 1901 (octomaculata group)1,2,6,8
sol. G oligo. Dalea
Perdita (Perdita)swenki Crawford, 1915
(octomaculata group)1,2,6,8
sol. G oligo. Solidago
*Perdita (Perdita)tridentata Stevens, 1919
(octomaculata group)2,6
sol. G oligo. Helianthus
Protandrenini
Protandrena (Heterosarus)parva
(Robertson, 1892)2,3
sol. G poly.
(Continued)
10 Gibbs et al.
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Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Protandrena (Pterosarus)aestivalis
(Provancher, 1882)3,7
sol. G oligo. Compositae: Astereae
Protandrena (Pterosarus)albertensis
(Cockerell, 1937)2
sol. G oligo. Compositae: Cichorieae (?)
Protandrena (Pterosarus)albitarsis
(Cresson, 1872)2,3
sol. G oligo. Compositae: Heliantheae
Protandrena (Pterosarus)perlaevis
(Cockerell, 1896)2,3
sol. G oligo. Helianthus
Protandrena (Pterosarus)piercei
(Crawford, 1903)2,5
sol. G oligo. Helianthus
Protandrena (Pterosarus)renimaculata
(Cockerell, 1896)2,3
sol. G oligo. Grindelia
Protandrena (Pterosarus)rudbeckiae
(Robertson, 1895)1
sol. G oligo. Rudbeckia
Apidae
Anthophorinae
Anthophora (Clisodon)terminalis Cresson, 18691,2,5,8,9 sol. W poly.
Anthophora (Melea)bomboides Kirby, 1838
(bomboides group)1,2,8
sol. G poly.
Anthophora (Melea)occidentalis Cresson, 1869
(bomboides group)1,2
sol. G poly.
Anthophora (Mystacanthophora)walshii
Cresson, 1869 (montana group)2,3
sol. G poly.
Apinae
Apini
Apis (Apis)mellifera Linnaeus, 1758E,1,2,6,8 eus. H poly.
Bombini
Bombus (Alpinobombus)kirbiellus Curtis, 18351,2 eus. H poly.
Bombus (Alpinobombus)polaris Curtis, 18351,2,9 eus. H poly.
Bombus (Bombias)nevadensis Cresson, 18741,2,5,8 eus. H poly.
Bombus (Bombus)terricola Kirby, 18371,2,5,8.9 eus. H poly.
Bombus (Cullumanobombus)griseocollis
(De Geer, 1773)1,2,5,6,8,9
eus. H poly.
Bombus (Cullumanobombus)rufocinctus
Cresson, 18631,2,5,8,9
eus. H poly.
Bombus (Psithyrus)bohemicus Seild, 1838/
ashtoni (Cresson, 1864)1,2,8,9
par. H NA Bombus (Bombus)
Bombus (Psithyrus)citrinus (Smith, 1854)1,2,5 par. H NA Bombus bimaculatus,
B. impatiens, B. vagans
Bombus (Psithyrus)flavidus Eversmann, 18521,2,5,8 par. H NA Bombus
Bombus (Psithyrus)insularis (Smith, 1861)1,2,5,6 par. H NA Bombus
Bombus (Psithyrus)suckleyi Greene, 18601,2 par. H NA Bombus
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
Bombus (Pyrobombus)bimaculatus Cresson, 18631,2,8 eus. H poly.
Bombus (Pyrobombus)centralis Cresson, 18641,2 eus. H poly.
Bombus (Pyrobombus)flavifrons Cresson, 18631,2,5,8 eus. H poly.
Bombus (Pyrobombus)frigidus Smith, 18541,2,5,8 eus. H poly.
Bombus (Pyrobombus)huntii Greene, 18601,2,8 eus. H poly.
Bombus (Pyrobombus)impatiens Cresson, 18633,E eus. H poly.
Bombus (Pyrobombus)jonellus (Kirby, 1802)1,2,5 eus. H poly.
Bombus (Pyrobombus)melanopygus
Nylander, 18481,2,8
eus. H poly.
Bombus (Pyrobombus)mixtus Cresson, 18781,2 eus. H poly.
Bombus (Pyrobombus)perplexus Cresson, 18631,2,5,8,9 eus. H poly.
Bombus (Pyrobombus)sandersoni Franklin, 19131,2,5,8 eus. H poly.
Bombus (Pyrobombus)sylvicola Kirby, 18371,2,8 eus. H poly.
Bombus (Pyrobombus)ternarius Say, 18371,2,5,6.8 eus. H poly.
Bombus (Pyrobombus)vagans vagans
Smith, 18541,2,5,8,9
eus. H poly.
Bombus (Pyrobombus)vancouverensis
Cresson, 18781,5
eus. H poly.
Bombus (Subterraneobombus)borealis
Kirby, 18371,2,5,6,8
eus. H poly.
Bombus (Thoracobombus)fervidus
(Fabricius, 1798)1,2,8
eus. H poly.
Bombus (Thoracobombus)pensylvanicus
(De Geer, 1773)1,5
eus. H poly.
Eucerinae
Emphorini
Diadasia (Coquillettapis)australis australis
(Cresson, 1878)1,2
sol. G oligo. Opuntia
Diadasia (Coquillettapis)diminuta
(Cresson, 1878)2
sol. G oligo. Sphaeralcea
*Diadasia (Dasiapis)ochracea (Cockerell, 1903)2sol. G oligo. Sphaeralcea and related genera
Eucerini
Eucera (Synhalonia)atriventris (Smith, 1854)2sol. G poly.
Eucera (Synhalonia)cf.chrysobotryae
(Cockerell, 1908)2
sol. G poly.
Eucera (Synhalonia)hamata (Bradley, 1942)2sol. G poly.
Melissodes (Eumelissodes)agilis Cresson, 18781,2,6 sol. G oligo. Helianthus
*Melissodes (Eumelissodes)bidentis
Cockerell, 19142
sol. G oligo. Compositae
Melissodes (Eumelissodes)confusus Cresson, 18781,2,5 sol. G oligo. Compositae
Melissodes (Eumelissodes)coreopsis
Robertson, 19052
sol. G oligo. Compositae
(Continued)
12 Gibbs et al.
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Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Melissodes (Eumelissodes)denticulatus
Smith, 18542
sol. G oligo. Vernonia
Melissodes (Eumelissodes)druriellus (Kirby, 1802)1,2,5 sol. G oligo. Solidago, Symphyotrichum
Melissodes (Eumelissodes)illatus Lovell and
Cockerell, 19061,2,5
sol. G oligo. Compositae
Melissodes (Eumelissodes)menuachus
Cresson, 18682
sol. G oligo. Compositae
Melissodes (Eumelissodes)pallidisignatus
Cockerell, 19052,3
sol. G oligo. Compositae
Melissodes (Eumelissodes)perlusus
Cockerell, 19251,2,5
sol. G oligo. Compositae
Melissodes (Eumelissodes)snowii Cresson, 18721,2,5,6 sol. G oligo. Compositae
Melissodes (Eumelissodes)subagilis
Cockerell, 19052
sol. G oligo. Compositae
Melissodes (Eumelissodes)subillatus
LaBerge, 19611,2,5
sol. G oligo. Compositae
Melissodes (Eumelissodes)trinodis
Robertson, 19012,8
sol. G oligo. Helianthus and related
Compositae
Melissodes (Eumelissodes)wheeleri Cockerell, 19062sol. G oligo. Compositae
Melissodes (Heliomelissodes)desponsus
Smith, 18542
sol. G oligo. Cirsium
Melissodes (Heliomelissodes)rivalis Cresson, 18722,3 sol. G oligo. Cirsium
Melissodes (Melissodes)bimaculatus bimaculatus
(Lepeletier, 1825)2,8
sol. G poly.
Nomadinae
Ammobatoidini
Holcopasites calliopsidis calliopsidis
(Linsley, 1943)2,8
par. G NA Calliopsis andreniformis
Holcopasites heliopsis (Robertson, 1897)2,5 par. G NA Calliopsis?
Holcopasites stevensi Crawford, 19152,5,6 par. G NA Calliopsis
Epeolini
Epeolus ainsliei Crawford, 19321,2,5,8 par. G NA likely Colletes susannae and
possibly C. wilmattae
Epeolus americanus (Cresson, 1878)1,2,5 par. G NA Colletes consors mesocopus
Epeolus compactus Cresson, 18781,2 par. G NA Colletes kincaidii
Epeolus gibbsi Onuferko, 20181,2 par. G NA Colletes
Epeolus interruptus Robertson, 19001,2,5 par. G NA Colletes aestivalis, C. brevicornis,
and C. willistoni are possible
hosts
Epeolus minimus (Robertson, 1902)1,2,5 par. G NA Colletes kincaidii is a likely host
Epeolus scutellaris Say, 18241,2 par. G NA likely Colletes simulans armatus
Triepeolus eliseae Rightmyer, 20172par. G NA possibly Melissodes
(Eumelissodes)
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
*Triepeolus cf. grindeliae Cockerell, 19072par. G NA possibly Melissodes
(Eumelissodes)
Triepeolus helianthi (Robertson, 1897)2par. G NA Melissodes (Eumelissodes) and
Nomiinae
Triepeolus michiganensis Mitchell, 19622par. G NA Likely Melissodes (Eumelissodes)
Triepeolus obliteratus Graenicher, 19112,8 par. G NA Likely Melissodes (Eumelissodes)
Triepeolus occidentalis (Cresson, 1878)2par. G NA Melissodes, possibly M.
menuachus
Triepeolus pectoralis (Robertson, 1897)2,3 par. G NA Melissodes druriellus
Triepeolus subalpinus Cockerell, 19102par. G NA Melissodes (Eumelissodes),
possibly M. agilis
Epeoloidini
Epeoloides pilosulus (Cresson, 1878)1,2 par. G NA Macropis
Melectini
Brachymelecta californica (Cresson, 1878)2par. G NA Anthophora
*Brachymelecta interrupta (Cresson, 1872)2par. G NA Anthophora walshii
Neolarrini
Neolarra (Phileremulus)vigilans Cockerell, 18952,6 par. G NA Perdita
Neolarra (Phileremulus)mallochi
(Crawford, 1912)2
par. G NA Perdita
Nomadini
Nomada alpha Cockerell, 1905 (ruficornis group)2par. G NA Andrena (?)
Nomada aquilarum Cockerell, 1903
(roberjeotiana group)1,2
par. G NA Andrena (Cnemidandrena) (?)
Nomada articulata Smith, 1854 (erigeronis group)2par. G NA Agapostemon
Nomada australis Mitchell, 1962 (erigeronis group)2par. G NA Agapostemon splendens (?)
Nomada banksi Cockerell, 1907 (ruficornis group)2par. G NA Andrena asteris (?)
Nomada composita Mitchell, 1962
(ruficornis group)
par. G NA
*Nomada crawfordi crawfordi Cockerell, 1905
(ruficornis group)2
par. G NA Andrena (?)
Nomada cressonii Robertson, 1893
(ruficornis group)1,2
par. G NA Andrena (Melandrena,
Plastandrena)
Nomada cuneata (Robertson, 1903)
(ruficornis group: bidentate mandible)1,2
par. G NA Andrena vicina
Nomada denticulata Robertson, 1902
(ruficornis group)2
par. G NA Andrena (?)
Nomada depressa Cresson, 1863 (ruficornis group)2par. G NA Andrena (?)
Nomada fervida Smith, 1854 (vegana group)2,8 par. G NA Andrena (?)
Nomada cf. florilega Lovell and Cockerell, 19052,3 par. G NA Andrena (?)
*Nomada fuscicincta Swenk, 1915
(ruficornis group: bidentate mandible)2
par. G NA Andrena ziziae (?)
(Continued)
14 Gibbs et al.
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Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Nomada gracilis Cresson, 1863 (ruficornis group)2par. G NA Andrena (?)
Nomada imbricata Smith, 1854 (ruficornis group)1,2,8 par. G NA Andrena (Melandrena)
Nomada integerrima Dalla Torre, 1896
(ruficornis group)2
par. G NA Andrena (?)
Nomada lehighensis Cockerell, 1903
(ruficornis group)2,3
par. G NA Andrena (?)
Nomada luteoloides Robertson, 1895
(ruficornis group)2
par. G NA Andrena (Melandrena)
Nomada maculata Cresson, 1863
(ruficornis group: bidentate mandible)2,3
par. G NA Andrena (Melandrena)
Nomada obliterata Cresson, 1863
(ruficornis group)2
par. G NA Andrena (?)
Nomada ovata (Robertson, 1903)
(ruficornis group: bidentate mandible)2,3
par. G NA Andrena (?)
Nomada quadrimaculata (Robertson, 1903)
(ruficornis group: bidentate mandible)2
par. G NA Andrena (?)
Nomada rubicunda Olivier, 1811
(erigeronis group)2,6
par. G NA Agapostemon splendens
*Nomada sphaerogaster Cockerell, 1903
(ruficornis group)2
par. G NA Andrena (?)
Nomada valida Smith, 1854 (ruficornis group)2par. G NA Andrena (?)
Nomada vicina Cresson, 1863 (ruficornis group)2par. G NA Andrena (?)
Nomada vincta Say, 1837 (vincta group)1,2 par. G NA Andrena helianthi
*Nomada xantholepis Cockerell, 1911
(ruficornis group)2
par. G NA Andrena (?)
Xylocopinae
Ceratinini
Ceratina (Zadontomerus)calcarata
Robertson, 19001,2
sub. S poly.
Ceratina (Zadontomerus)dupla Say, 18371,2 sub. S poly.
Ceratina (Zadontomerus)mikmaqi
Rehan and Sheffield, 20112,6
sub. S poly.
Colletidae
Colletinae
Colletini
Colletes americanus Cresson, 1868
(americanus group)1,2,5
sol. G oligo. Compositae
Colletes andrewsi Cockerell, 1906
(aestivalis group)1,2,5,8
sol. G oligo. Heuchera
Colletes brevicornis Robertson, 1897
(willistoni group)1,2,5,8
sol. G oligo. Campanula
Colletes consors mesoscopus Swenk, 1907
(consors group)1,2
sol. G oligo. Phacelia
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
Colletes hyalinus hyalinus Provancher, 1888
(hyalinus group)2,5
sol. G poly.
Colletes impunctatus lacustris Swenk, 1906
(clypearis group)2
sol. G oligo. Gaylussacia
Colletes inaequalis Say, 1837 (inaequalis group)1,2,5,8 sol. G poly.
Colletes kincaidii Cockerell, 1898 (simulans group)1,2,5 sol. G poly.
Colletes nigrifrons Titus, 1900 (consors group)5,7 (AMNH) sol. G poly.
Colletes petalostemonis Swenk, 1906 (daleae group)1sol. G oligo. Dalea
Colletes phaceliae Cockerell, 1906 (hyalinus group)1sol. G poly.
Colletes robertsonii Dalla Torre, 1896
(robertsonii group)1,2,5
sol. G oligo. Fabaceae
Colletes rufocinctus Cockerell, 1929
(simulans group)1,2
sol. G oligo. Compositae
Colletes simulans armatus Patton, 1879
(simulans group)1,2,5
sol. G oligo. Solidago, Symphyotrichum
Colletes susannae Swenk, 1925
(americanus group)1,2,5,8
sol. G oligo. Dalea
Colletes validus Cresson, 1868 (inaequalis group)2,5 sol. G oligo. Ericaceae
Colletes willistoni Robertson, 1895
(willistoni group)1,2,5
sol. G oligo. Physalis
Colletes wilmattae Cockerell, 1904
(americanus group)1,2,5,6
sol. G oligo. Dalea
Hylaeinae
Hylaeini
Hylaeus (Cephalylaeus)basalis (Smith, 1853)1,2,5 sol. S poly. Rosaceae
Hylaeus (Hylaeus)annulatus (Linnaeus, 1758)
(annulatus group)2,3,5,8
sol. S poly. Rosaceae
Hylaeus (Hylaeus) aff. rudbeckiae
(Cockerell and Casad, 1895) (rudbeckiae group)2
sol. S poly.
Hylaeus (Hylaeus)fedorica (Cockerell, 1909)
(rudbeckiae group)2
sol. S poly.
Hylaeus (Hylaeus)leptocephalus
(Morawitz, 1871 [“1870”]) (leptocephalus group)E,2,3
sol. S poly. Fabaceae
Hylaeus (Hylaeus)mesillae cressoni (Cockerell, 1907)
(rudbeckiae group)2,3,5
sol. S poly.
Hylaeus (Hylaeus)rudbeckiae
Cockerell and Casad, 1895 (rudbeckiae group)2,5
sol. S poly.
Hylaeus (Hylaeus)saniculae (Robertson, 1896)
(rudbeckiae group)2
sol. S poly.
Hylaeus (Hylaeus)verticalis (Cresson, 1869)
(verticalis group)2,3,5
sol. S poly.
Hylaeus (Prosopis)affinis (Smith, 1853)1,2,8 sol. S poly.
Hylaeus (Prosopis) cf. gaigei (Cockerell, 1916)2sol. S poly.
(Continued)
16 Gibbs et al.
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Hylaeus (Prosopis)illinoisensis (Robertson, 1896) 2,3 sol. S poly.
Hylaeus (Prosopis)modestus modestus Say, 18372,3,8 sol. S poly.
Hylaeus (Prosopis)nelumbonis (Robertson, 1890)2,3 sol. S poly.
Hylaeus (Prosopis)sp.A
2sol. S poly.
Halictidae
Halictinae
Augochlorini
Augochlorella aurata (Smith, 1853)1,2,5,8 eus. G poly.
Halictini
Agapostemon (Agapostemon)sericeus (Forster, 1771)
(sericeus group)1,2,5,8
sol. G poly.
Agapostemon (Agapostemon)splendens
(Lepeletier, 1841) (splendens group)1,2,5,6
sol. G poly.
Agapostemon (Agapostemon)texanus Cresson, 1872
(splendens group)1,2,5,6,8
sol. G poly.
Agapostemon (Agapostemon)virescens
(Fabricius, 1775)1,2,8
com. G poly.
Halictus (Nealictus)parallelus Say, 18371,2 eus. G poly.
Halictus (Odontalictus)ligatus Say, 18371,2 eus. G poly.
Halictus (Protohalictus)rubicundus (Christ, 1791)1,2,5,8 eus. G poly.
Halictus (Seladonia)confusus confusus
Smith, 18531,2,8
eus. G poly.
Halictus (Seladonia)virgatellus Cockerell, 19011,2 sol. G poly.
Lasioglossum (Dialictus)absimile
(Sandhouse, 1924) (viridatum group)1,2
eus.? G poly.
Lasioglossum (Dialictus)admirandum
(Sandhouse, 1924) (viridatum group)1,2
eus.? G poly.
Lasioglossum (Dialictus)albipenne
(Robertson, 1890)1,2,6
eus.? G poly.
Lasioglossum (Dialictus)albohirtum
(Crawford, 1907)4
eus.? G poly.
Lasioglossum (Dialictus)cressonii
(Robertson, 1890)2,3
eus.? W poly.
Lasioglossum (Dialictus)dreisbachi
(Mitchell, 1960) (viridatum group)2
eus.? G poly.
Lasioglossum (Dialictus)ellisiae (Sandhouse, 1924)
(gemmatum group)2
sol.? G poly.
Lasioglossum (Dialictus)ephialtum Gibbs, 2010
(viridatum group)2
eus.? G poly.
Lasioglossum (Dialictus)imitatum (Smith, 1853)1,2 eus. G poly.
Lasioglossum (Dialictus)immigrans
Gardner and Gibbs, 20211,2
eus.? G poly
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
Lasioglossum (Dialictus)laevissimum
(Smith, 1853)1,2,8,9
eus. G poly.
Lasioglossum (Dialictus)leucocomus
(Lovell, 1908)2
eus.? G poly.
Lasioglossum (Dialictus)lineatulum
(Crawford, 1906)1,2
eus. G poly.
Lasioglossum (Dialictus)michiganense
(Mitchell, 1960) (platyparium group)2
par. G NA Lasioglossum (Dialictus)
Lasioglossum (Dialictus)nigroviride
(Graenicher, 1910)1,2,8
eus.? W poly.
Lasioglossum (Dialictus)novascotiae (Mitchell, 1960)
(viridatum group)2,3
eus.? G poly.
Lasioglossum (Dialictus)oblongum (Lovell, 1905)
(viridatum group)2
eus.? W poly.
Lasioglossum (Dialictus)occidentale (Crawford, 1902)
(anomalum group)1,2
eus.? G poly.
Lasioglossum (Dialictus)packeri Gibbs, 20101,5 eus.? G poly.
Lasioglossum (Dialictus)pavoninum (Ellis, 1913)1,5 eus.? G poly.
Lasioglossum (Dialictus)perpunctatum
(Ellis, 1913)1,2,9
eus.? G poly.
Lasioglossum (Dialictus)pictum (Crawford, 1902)1,2,6 eus.? G poly.
Lasioglossum (Dialictus)pilosum
(Smith, 1853)1,2,3,8,9
eus.? G poly.
Lasioglossum (Dialictus)planatum (Lovell, 1905)
(viridatum group)2
eus.? G poly.
Lasioglossum (Dialictus)prasinogaster Gibbs, 2010
(veganum group)2
eus.? G poly.
Lasioglossum (Dialictus)pruinosum
(Robertson, 1892)1,2
eus.? G poly.
Lasioglossum (Dialictus)rufulipes (Cockerell, 1938)
(testaceum group)1,2
sol.? G poly.
Lasioglossum (Dialictus)sagax (Sandhouse, 1924)
(viridatum group)1,2
eus.? G poly.
Lasioglossum (Dialictus)semicaeruleum
(Cockerell, 1895)1,2
eus.? G poly.
Lasioglossum (Dialictus)sheffieldi Gibbs, 2010
(perdifficile group)1,2
eus.? G poly.
Lasioglossum (Dialictus)sitocleptum Gibbs, 2010
(platyparium group)2
par. G NA Lasioglossum (Dialictus)
Lasioglossum (Dialictus)subversans
(Mitchell, 1960)1,2
eus.? G poly.
Lasioglossum (Dialictus)subviridatum
(Cockerell, 1938) (viridatum group)2
eus.? W poly.
Lasioglossum (Dialictus)succinipenne (Ellis, 1913)1,2 eus.? G poly.
Lasioglossum (Dialictus)taylorae Gibbs, 2010
(viridatum group)2
eus.? G poly.
(Continued)
18 Gibbs et al.
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Lasioglossum (Dialictus)tenax (Sandhouse, 1924)2sol. G poly.
Lasioglossum (Dialictus)timothyi Gibbs, 20101,2 eus.? G poly.
Lasioglossum (Dialictus)versans (Lovell, 1905)
(ruidosense group)1,2
eus.? G poly.
Lasioglossum (Dialictus)versatum
(Robertson, 1902)1,2
eus. G poly.
Lasioglossum (Dialictus)vierecki
(Crawford, 1904)1,2,6,8
sol. G poly.
Lasioglossum (Dialictus)viridatum Lovell, 1905
(viridatum group)1,2
eus.? G poly.
Lasioglossum (Dialictus)zephyrus (Smith, 1853)1,2,8 eus. G poly.
Lasioglossum (Evylaeus)cinctipes
(Provancher, 1888)1,2
eus. G poly.
Lasioglossum (Hemihalictus)foxii
(Robertson, 1895)2,6,8
sol. G poly.
Lasioglossum (Hemihalictus)inconditum
(Cockerell, 1916)1,2
sol. G poly.
Lasioglossum (Hemihalictus)macoupinense
(Robertson, 1895)1,2
sol. G poly.
Lasioglossum (Hemihalictus)nelumbonis
(Robertson, 1890)2
sol. G oligo.
Lasioglossum (Hemihalictus)pectorale
(Smith, 1853)1,2,6
sol. G poly.
Lasioglossum (Hemihalictus)swenki
(Crawford, 1906)1,2,6
sol. G poly.
Lasioglossum (Lasioglossum)acuminatum
McGinley, 1986 (forbesii group)2,9
sol. G poly.
Lasioglossum (Lasioglossum)athabascense
(Sandhouse, 1933)1,2
sol. G poly.
Lasioglossum (Lasioglossum)coriaceum
(Smith, 1853)1,2
sol. G poly.
Lasioglossum (Lasioglossum)paraforbesii
McGinley, 1986 (forbesii group)1,2,6
sol. G poly.
Lasioglossum (Leuchalictus)leucozonium
(Schrank, 1781)E,2,3,6
sol. G poly.
Lasioglossum (Leuchalictus)zonulus
(Smith, 1848)E,1,2,6,8.9
sol. G poly.
Lasioglossum (Sphecodogastra)aberrans
(Crawford, 1903) (lusorium group)1
sol. G oligo. Onagraceae
Lasioglossum (Sphecodogastra)boreale
Svensson, Ebmer and Sakagami, 1977
(fulvicorne/fratellum group)1,2
sol. G poly.
Lasioglossum (Sphecodogastra)comagenense
(Knerer and Atwood, 1964)
(fulvicorne/fratellum group)2
com. G poly.
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
Lasioglossum (Sphecodogastra)lusorium
(Cresson, 1872) (lusorium group)4
sol. G oligo. Onagraceae
Lasioglossum (Sphecodogastra)quebecense
(Crawford, 1907) (fulvicorne/fratellum group)2
sol. G poly.
Lasioglossum (Sphecodogastra)seillean
Gibbs and Packer, 2013 (fulvicorne/fratellum group)2
sol.? G poly.
Lasioglossum (Sphecodogastra)truncatum
(Robertson, 1901)2
eus. G poly.
Sphecodes atlantis Mitchell, 1956
(mandibularis group)2
par. G NA
Sphecodes banksii Lovell, 1909
(mandibularis group)2
par. G NA Lasioglossum vierecki
Sphecodes clematidis Robertson, 1897
(dichrous group)2,3
par. G NA
Sphecodes confertus Say, 1837
(confertus group)2,6
par. G NA
Sphecodes coronus Mitchell, 1956
(mandibularis group)2
par. G NA
Sphecodes cressonii (Robertson, 1903)
(mandibularis group)2
par. G NA
Sphecodes davisii Robertson, 1897
(mandibularis group)2,3
par. G NA
Sphecodes dichrous Smith, 1853 (dichrous group)2par. G NA
Sphecodes cf. galerus Lovell and Cockerell, 1907
(mandibularis group)2
par. G NA
Sphecodes illinoensis (Robertson, 1903)
(mandibularis group)2
par. G NA
Sphecodes johnsonii Lovell, 1909
(mandibularis group)2
par. G NA
Sphecodes levis Lovell and Cockerell, 1907
(mandibularis group)2
par. G NA
Sphecodes mandibularis Cresson, 1872
(mandibularis group)2
par. G NA
Sphecodes pecosensis pecosensis Cockerell, 1904
(ranunculi group)2
par. G NA
Sphecodes prosphorus Lovell and Cockerell, 1907
(dichrous group)2
par. G NA
Sphecodes pycnanthemi Robertson, 1897
(mandibularis group)2
par. G NA
Sphecodes smilacinae Robertson, 1897
(mandibularis group)2
par. G NA
Sphecodes solonis Graenicher, 1910
(dichrous group)2
par. G NA
Sphecodes townesi Mitchell, 1956
(mandibularis group)2
par. G NA
Nomiinae
Dieunomiini
(Continued)
20 Gibbs et al.
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Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Dieunomia (Dieunomia)heteropoda heteropoda
(Say, 1824)1,5
sol. G oligo. Helianthus ()
Rophitinae
Rophitini
Dufourea harveyi (Cockerell, 1906)2sol. G oligo. Potentilla
Dufourea marginata marginata (Cresson, 1878)1,2 sol. G oligo. Helianthus
Dufourea maura (Cresson, 1878)1,2,8 sol. G oligo. Campanula
Megachilidae
Megachilinae
Anthidiini
Anthidium (Anthidium)clypeodentatum
Swenk, 19141,2,5,6,8
sol. G poly. Fabaceae
Anthidium (Anthidium)manicatum manicatum
Linnaeus, 1758E, 2,8
sol. C poly.
Anthidium (Anthidium)tenuiflorae Cockerell, 19071,2 sol. G poly. Phacelia
*Dianthidium (Dianthidium)parvum
(Cresson, 1878)2,6
sol. E oligo.
Dianthidium (Dianthidium)pudicum pudicum
(Cresson, 1879)1,2
sol. E oligo.
Dianthidum (Dianthidium)simile (Cresson, 1864)1,2 sol. G oligo.
Stelis (Stelis)coarctatus Crawford, 19162,7,8 par. C NA Heriades,Hoplitis
Stelis (Stelis)foederalis Smith, 18542,7 par. C NA Hoplitis, Osmia
Stelis (Stelis)labiata (Provancher, 1888)1,2 par. C NA Hoplitis
Stelis (Stelis)lateralis Cresson, 18641,2 par. C NA Hoplitis
Stelis (Stelis) aff. interrupta Cresson, 18972par. C NA
Stelis (Stelis)nitida Cresson, 18782par. C NA
Stelis (Stelis)permaculata Cockerell, 18981,2 par. C NA Heriades carinata
Stelis (Stelis)subemarginata Cresson, 18781par. C NA Hoplitis, Osmia
Megachilini
Coelioxys (Boreocoelioxys)moestus Cresson, 18641,2,5 par. C NA Megachile
Coelioxys (Boreocoelioxys)octodentatus
Say, 18241,2,5,6
par. G NA Megachile
Coelioxys (Boreocoelioxys)porterae
Cockerell, 19001,2,5
par. C NA Megachile
Coelioxys (Boreocoelioxys)rufitarsis Smith, 18541,2,5,8 par. G NA Megachile
Coelioxys (Coelioxys)sodalis Cresson, 18781,2,5 par. G NA Megachile
Coelioxys (Cyrtocoelioxys)modestus Smith, 18542,8 par. C NA Megachile campanulae
Coelioxys (Paracoelioxys)funerarius Smith, 18541,2,5 par. G NA Megachile
Coelioxys (Synocoelioxys)alternatus Say, 18371,2,5 par. C NA Megachile pugnata
*Coelioxys (Xerocoelioxys)nodis Baker, 19752,6 par. C NA Megachile
(Continued)
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Table 1. (Continued )
Species Nests Substrate Lecty Floral or bee host
Megachile (Chelostomoides)campanulae
(Robertson, 1903) (exilis group)2,3
sol. C poly. Preference for Campanula
Megachile (Eutricharaea)rotundata
(Fabricius, 1787)E, 1,2
sol. C poly.
Megachile (Litomegachile)brevis Say, 18371,2 sol. C poly.
Megachile (Litomegachile)mendica Cresson, 18781,2 sol. C poly.
Megachile (Litomegachile)texana Cresson, 18781,2,5 sol. G poly.
Megachile (Megachile)centuncularis
(Linnaeus, 1758)1,2
sol. C poly.
Megachile (Megachile)inermis
Provancher, 18881,2,5,8,9
sol. C poly.
Megachile (Megachile)lapponica Thomson, 18721,2 sol. C poly.
Megachile (Megachile)montivaga Cresson, 18781,2,5 sol. S poly.
Megachile (Megachile)relativa Cresson, 18781,2.5,9 sol. C poly.
*Megachile (Megachiloides)dakotensis
Mitchell, 19262,6
sol. G poly.
Megachile (Megachiloides)wheeleri
Mitchell, 1927RSKM
sol. G poly.
Megachile (Phaenosarus)fortis Cresson, 18723sol. G poly.
Megachile (Sayapis)pugnata pugnata Say, 18371,2,8 sol. C oligo. Compositae
Megachile (Xanthosarus)circumcincta (Kirby, 1802)1,2 sol. G poly.
Megachile (Xanthosarus)frigida frigida
Smith, 18531,2,5,8,9
sol. C poly.
Megachile (Xanthosarus)gemula gemula
Cresson. 18781,2,9
sol. C poly.
Megachile (Xanthosarus)latimanus Say, 18231,2,5,8,9 sol. G poly.
Megachile (Xanthosarus)melanophaea melanophaea
Smith, 18531,2,5,6,8
sol. G poly.
Megachile (Xanthosarus)perihirta
Cockerell, 18981,2,5,6
sol. G poly.
Osmiini
Ashmeadiella (Ashmeadiella)bucconis bucconis
(Say, 1837)2,6,8
sol. C oligo. Compositae
Heriades (Neotrypetes)carinata Cresson, 18641,2,5,6 sol. S poly.
Heriades (Neotrypetes)variolosa variolosa
(Cresson, 1872)1,2
sol. S oligo. Compositae
Hoplitis (Alcidamea)albifrons albifrons
(Kirby, 1837)1,2,5,8
sol. S poly.
Hoplitis (Alcidamea)pilosifrons (Cresson, 1864)2,3,5,6 sol. S poly.
Hoplitis (Alcidamea)producta producta
(Cresson, 1864)1,2,5
sol. S poly.
Hoplitis (Alcidamea)spoliata (Provancher, 1888)1,2 sol. S poly.
Hoplitis (Alcidamea)truncata (Cresson, 1878)2sol. S poly.
(Continued)
22 Gibbs et al.
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
Table 1. (Continu ed )
Species Nests Substrate Lecty Floral or bee host
Hoplitis (Formicapis)robusta (Nylander, 1848)1sol. S oligo. Potentilla
Osmia (Cephalosmia)subaustralis Cockerell, 19002sol. S oligo. Compositae
Osmia (Melanosmia)aquilonaria
Rightmyer et al., 20102
sol. unk. poly.
Osmia (Melanosmia)atriventris Cresson, 18641,2 sol. C poly.
Osmia (Melanosmia)bucephala Cresson, 18642,3,8 sol. C poly.
Osmia (Melanosmia)cyaneonitens Cockerell, 19062,3 sol. unk. unk.
Osmia (Melanosmia)distincta Cresson, 18641,2,6 sol. C oligo. Penstemon
Osmia (Melanosmia)cf.grindeliae Cockerell, 1910 sol. unk. unk.
Osmia (Melanosmia)illinoensis Robertson, 18972sol. C unk.
Osmia (Melanosmia)inermis (Zetterstedt, 1838)1,2 sol. E oligo. Vaccinium
Osmia (Melanosmia)inspergens
Lovell and Cockerell, 19072
sol. E poly.
Osmia (Melanosmia)integra Cresson, 18781,2,5,6 sol. E poly.
Osmia (Melanosmia)laticeps Thomson, 18721,2 sol. unk. oligo. Vaccinium
Osmia (Melanosmia)nearctica
Rightmyer et al., 20101,2
sol. unk. poly.
Osmia (Melanosmia)nigrifrons Cresson, 18783,5 sol. unk. unk.
Osmia (Melanosmia)nigriventris (Zetterstedt, 1838)1,2 sol. C unk.
Osmia (Melanosmia)paradisica Sandhouse, 19243sol. unk. unk.
Osmia (Melanosmia)proxima Cresson, 18642,5 sol. C unk.
Osmia (Melanosmia)simillima Smith, 18531,2,5 sol. C poly.
Osmia (Melanosmia)subarctica Cockerell, 19122sol. unk. poly.
Osmia (Melanosmia)tarsata Provancher, 1888RSKM sol. G poly.
Osmia (Melanosmia)tersula Cockerell, 19122,3 sol. C poly.
Osmia (Melanosmia)trevoris Cockerell, 18972sol. unk. poly.
Osmia (Melanosmia)virga Sandhouse, 19392sol. unk. oligo. Vaccinium
Osmia (Osmia)lignaria lignaria Say, 18371,2 sol. C poly.
Osmia (Osmia)lignaria Say, 1837 propinqua
Cresson, 18641
sol. C poly.
Melittidae
Melittinae
Macropidini
Macropis (Macropis)nuda (Provancher, 1882)1,2,5,8,9 sol. G oligo. Lysimachia
The Canadian Entomologist 23
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Results
We documented 392 species for Manitoba (Table 1), based on examination of more than
67 000 specimens. We found 154 new records for the province since 2015. Brachymelecta,
Eucera,Neolarra,Triepeolus,Ashmeadiella, and Dianthidium represent new generic records
for Manitoba during this time. Thirteen species are newly recorded for Canada: Calliopsis
(Nomadopsis)australior Cockerell, 1897; Perdita (Perdita)tridentata Stevens, 1919;
Brachymelecta interrupta (Cresson, 1872); Diadasia (Dasiapis)ochracea (Cockerell, 1903);
Melissodes bidentis Cockerell, 1914; Nomada crawfordi crawfordi Cockerell, 1905; Nomada
fuscicincta Swenk, 1915; Nomada sphaerogaster Cockerell, 1903; Nomada xantholepis
Cockerell, 1911; Triepeolus cf. grindeliae Cockerell; Coelioxys (Xerocoelioxys)nodis Baker, 1972;
Dianthidium (Dianthidium)parvum (Cresson, 1878); and Megachile (Megachiloides)dakotensis
Mitchell, 1926. We propose that Nomada alpha paralpha Cockerell, 1921 and N. alpha dialpha
Cockerell, 1921 are junior synonyms of N. alpha Cockerell, 1905, based on the type
localities all being within a small geographical area (Supplementary material S1). Nomada
arenicola Swenk, 1912 is considered a junior synonym of N. fervida Smith, 1854, due to a
lack of morphological or genetic separation (Supplementary material S1). Protandrena
albertensis (Cockerell, 1937) and Neolarra mallochi Michener, 1939 are recognised as valid
species (Supplementary material S1). Supporting information for new and interesting records
is provided in Supplementary material S1.
ANDRENIDAE
We document 74 andrenid bees in Manitoba, based on more than 4000 specimens, including
54 species of Andrena, three of Calliopsis, nine of Perdita, and eight of Protandrena for the
province. Perdita swenki Crawford was by far the most common species, with over 1400
records, which were mostly collected in pan traps. Four species –Andrena peckhami
Cockerell, Andrena robervalensis Mitchell, Perdita octomaculata (Say), and Protandrena
rudbeckiae (Robertson) –are provisional records included solely based on unconfirmed
literature accounts. Approximately half of the Andrena species (25) are oligolectic, and most
of the panurgines are oligoleges.
APIDAE
We document 113 apid species, based on more than 23 000 records, including four species of
Anthophora, Apis mellifera, 29 of Bombus, two of Brachymelecta, three of Ceratina, three of
Diadasia,Epeoloides pilosulus (Cresson), seven of Epeolus, three of Eucera, three of
Holcopasites,18ofMelissodes, two of Neolarra,29ofNomada, and eight of Triepeolus for the
province. Four apid genera were newly recorded for the province during this work: Eucera,
Triepeolus (see also Wrigley et al. 2021), Brachymelecta (see also Onuferko et al. 2021), and
Neolarra. Many new records come from the cleptoparasitic subfamily Nomadinae, including
Holcopasites calliopsidis (Linsley), H. heliopsis (Robertson), Neolarra vigilans Cockerell, and
N. mallochi, as well as 19 species of Nomada and eight of Triepeolus. The number of Nomada
species remains uncertain until the genus can be revised. Eucerinae is also well represented by
new records, including Eucera and six species of Melissodes.
COLLETIDAE
We document 33 colletid species, based on more than 2500 specimens, including 18 species of
Colletes and 15 of Hylaeus for the province. Three-quarters of the Colletes are oligolectic (14 of 18).
Colletes albescens Cresson is excluded from our verified list (see section on excluded bees in
Supplementary material S1). Colletes petalostemonis was not re-examined and is based on a
literature record (Sheffield et al. 2014). New records of Hylaeus include species that are
typically quite rare and not commonly recorded in Canada.
24 Gibbs et al.
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press
HALICTIDAE
We document 95 halictid species based on approximately 30 000 records, including four
species of Agapostemon, Augochlorella aurata (Smith), Dieunomia heteropoda (Say)¸ three
species of Dufourea, five of Halictus,62ofLasioglossum, and 19 of Sphecodes for the province.
Many new records are provided for Lasioglossum and Sphecodes. The latter is entirely
composed of brood parasites. Two brood parasitic Lasioglossum are recorded, the first
documented for the province. Several additional Lasioglossum are reported as new. Sphecodes
needs revision; therefore, many individuals are identified tentatively. Lasioglossum have been
revised for the region, but the viridatum group remains a challenge. Among the halictids, the
only oligoleges are Dieunomia heteropoda,L. aberrans (Crawford), L. lusorium (Cresson),
L. nelumbonis (Robertson), and the three Dufourea.
MEGACHILIDAE
We document 76 megachild species based on more than 6000 records, including three species
of Anthidium,Ashmeadiella bucconis (Say), nine species of Coelioxys, three of Dianthidium, two of
Heriades, six of Hoplitis,20ofMegachile,24ofOsmia, and eight of Stelis for the province. The
genus Ashmeadiella is newly recorded for the province. Dianthidium parvum is a new Canadian
record. These new records include one exotic species, Anthidium manicatum Linnaeus. Both the
nominal subspecies of Osmia lignaria and the subspecies O. lignaria propinqua are recorded for
the province. The latter has been introduced by commercial retailers of Osmia.
MELITTIDAE
We record one species, Macropis nuda (Provancher), based on 85 specimens (Gibbs
et al. 2021). Although M. ciliata Patton has appeared on some lists for the province, this was
apparently based on a misidentified specimen of M. nuda at the Illinois Natural History
Survey. Examination of the Illinois Natural History Survey specimen revealed it to have the
dark basitibial hairs and sculptured metapostnotum typical of M. nuda. The original
determination label also reads M. nuda.
Discussion
Our study documents a 64.7% increase in the known species richness since 2015 and a 15.6%
increase in the known generic richness of bees in Manitoba. The notable increase is the result of
recent intensive sampling of a historically undersampled region for bees and examination of large
numbers of specimens (including previously unidentified material) from museum collections. In
comparison, Sheffield and Heron’s(2019) recent checklist of the bees of British Columbia, which
has been better sampled historically for bees, documented only an 8.3% increase in the known
species richness for that province, bringing the total number to 483 species. That Manitoba
should contain upwards of 40% of the species of bees known to occur in Canada can be
explained by its geographic position at the longitudinal centre of Canada and as the transition
from eastern forests into western prairie. Manitoba thus marks the easternmost range for
multiple western species and the westernmost range for multiple eastern species.
The disparate habitats and floral communities present in Manitoba, from boreal forests to
prairies, provide for a more diverse bee fauna than was previously realised. Habitat conditions
may affect proportions of different plant syndromes and the prevalence of associated
pollinators (Robson et al. 2019). Earlier checklists documented fewer than 250 species
(Sheffield et al. 2014; Canadian Endangered Species Conservation Council 2015); however, we
record 392 bee species in the province. We would like to draw attention to three areas that
are of particular interest for future studies. First, we found records of many eastern species
that had previously gone undetected. It is likely that additional studies in the southeastern
corner of Manitoba, which has not been thoroughly sampled, will reveal more records of
The Canadian Entomologist 25
https://doi.org/10.4039/tce.2022.45 Published online by Cambridge University Press