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Hymenoptera of Canada
expand article infoAndrew M.R. Bennett, Cory S. Sheffield§, Jeremy R. deWaard|
‡ Canadian National Collection of Insects, Ottawa, Canada
§ Royal Saskatchewan Museum, Regina, Canada
| University of Guelph, Guelph, Canada
Open Access

Abstract

A summary of the numbers of species of the 83 families of Hymenoptera recorded in Canada is provided. In total, 8757 described species are recorded compared to approximately 6000 in 1979, which is a 46% increase. Of the families recognized in 1979, three have been newly recorded to Canada since the previous survey: Anaxyelidae (Anaxyleoidea), Liopteridae (Cynipoidea), and Mymarommatidae (Mymarommatoidea). More than 18,400 BINs of Canadian Hymenoptera are available in the Barcode of Life Data Systems (Ratnasingham and Hebert 2007) implying that nearly 9650 undescribed or unrecorded species of Hymenoptera may be present in Canada (and more than 10,300 when taking into account additional species that have not been DNA barcoded). The estimated number of unrecorded species is very similar to that of 1979 (10,637 species), but the percentage of the fauna described/recorded has increased from 36% in 1979 to approximately 45% in 2018. Summaries of the state of knowledge of the major groups of Hymenoptera are presented, including brief comments on numbers of species, biology, changes in classification since 1979, and relevant taxonomic references.

Keywords

ants, bees, biodiversity assessment, Biota of Canada, DNA barcodes, Hymenoptera, survey, wasps

Introduction

Hymenoptera constitutes one of the most speciose orders in Canada and the world (Forbes et al. 2018). During the last assessment of this order in Canada (Masner et al. 1979), 6028 species were reported, although the approximately 80 species in the family Eurytomidae were inadvertently omitted and the numbers of species of Platygastroidea, Ceraphronoidea, Bethylidae, Cynipoidea and Pompilidae were overestimated as it appears that undescribed species were included. Thus, the known richness in 1979 was approximately 6000 species. The most comprehensive faunal inventory of Hymenoptera in Canada is the Catalog of the Hymenoptera in America North of Mexico (Krombein et al. 1979) which listed general distributions of species up to 1972 to 1976 (the cut-off date depending on the superfamily). No complete distributional survey of the species of Hymenoptera in North America (or Canada) has followed, although the species lists and distributions on which the analysis in this manuscript is based will be published in a forthcoming series of checklists of the Hymenoptera of Canada, Alaska, and Greenland (A Bennett unpubl. data). Nonetheless, a tremendous amount of data has been produced since 1979 on the taxonomy, nomenclature, and distribution of particular groups of Hymenoptera, including the presence of species in Canada. Some of the most important sources are noted in the respective sections on major taxa (see below as well as in Table 1). In some instances in which major references were not included in Masner et al. (1979), these references have also been included in Table 1.

Census of Hymenoptera in Canada.

Taxon1 No. species reported in Masner et al. (1979) No. species currently known from Canada No. BINs2 available for Canadian species Est. no. undescribed or unrecorded species in Canada3 General distribution by ecozone3A Information sources4
Sawflies (previously suborder Symphyta) Smith 1979a, Goulet 1987, Taeger et al. 2010, Blank et al. 2012
Superfamily Anaxyleoidea
Anaxyelidae 0 1 0 0 Western Interior Basin Goulet 1992
Superfamily Cephoidea
Cephidae 16 12 8 0 all except Arctic Ries 1937, Smith 1986, Smith and Solomon 1989, Smith and Schiff 2005
Superfamily Orussoidea5
Orussidae 3 5 4 0 south of Boreal Cordillera and Taiga ecozones, except Atlantic Maritime Middlekauff 1983, Skvarla et al. 2015
Superfamily Pamphilioidea6
Pamphiliidae 47 54 33 0 all except Arctic Middlekauff 1958, 1964, Eidt 1969
Superfamily Siricoidea5
Siricidae 14 20 25 4 all except Arctic Schiff et al. 2012, Goulet et al. 2015
Xiphydriidae 5 7 3 0 all except Arctic Smith 1976
Superfamily Tenthredinoidea
Argidae 14 27 17 0 all except Arctic Smith 1969a, 1971a, 1989
Cimbicidae 4 8 15 7 southern Arctic and south Smith 1979a
Diprionidae 22 25 14 0 southern Arctic and south Ross 1955, Smith 1974
Pergidae 5 3 4 1 Mixedwood Plains, Atlantic Maritime Smith 2006
Tenthredinidae 300 532 528 200* all ecozones Goulet 1986, 1996, Smith 1969b, c, 1971b, 1979b
Superfamily Xyeloidea6
Xyelidae 13 16 11 0 all except Arctic Burdick 1961, Smith 1977, Smith and Schiff 1998
Total Sawflies 443 710 662 212*
Suborder Apocrita
Superfamily Ichneumonoidea
Braconidae 830 1165 3411 2246 all ecozones Marsh 1979a, b, Wharton et al. 1997, Yu et al. 2016
Ichneumonidae 2001 3037 4748 1705* all ecozones Carlson 1979c, Dasch 1979, 1984, 1988, 1992, Townes 1983, Townes et al. 1992, Schwarzfeld 2014, Yu et al. 2016
Total Ichneumonoidea 2831 4202 8159 3951*
Superfamily Diaprioidea7
Diapriidae 8 150 177 760 583 southern Arctic and south Masner 1991, 1993a, Johnson 1992, Masner and García 2002, Sharkey 2007, Sharkey et al. 2012
Diapriidae 8 ? 8 3 0 south of Arctic and Boreal Cordillera Masner 1976a
Total Diaprioidea 150 185 763 583
Superfamily Platygastroidea9
Platygastridae 10 250 160 2287 2127 southern Arctic and south Masner 1976b, 1983a, b, Townes and Townes 1981, Ritchie and Masner 1983, Johnson 1992, 2018
Superfamily Proctotrupoidea11
Heloridae 2 2 2 0 all except Arctic Townes 1977b, Johnson 1992
Pelecinidae 1 1 1 0 Boreal Shield, Mixedwood Plains, Atlantic Maritime Johnson 1992, Johnson and Musetti 1999
Proctotrupidae 60 67 63 0 southern Arctic and south Townes and Townes 1981, Johnson 1992
Roproniidae 2 2 1 0 Boreal Shield, Mixedwood Plains Townes 1948, Johnson 1992
Vanhorniidae 1 1 0 0 Boreal Shield, Mixedwood Plains Townes and Townes 1981, Johnson 1992
Total Proctotrupoidea 66 73 67 0
Superfamily Chalcidoidea Gibson et al. 1997, Noyes 2017
Aphelinidae 30 37 130 93 all ecozones Gordh 1979, Rosen and Debach 1979, Schauff et al. 1996, Shirley et al. 2017
Azotidae 12 ? 1 1 0 Mixedwood Plains Jarvis 1907, Heraty et al. 2013
Chalcididae 22 37 18 0 southern Arctic and south Burks 1979d, Halstead 1990,Delvare and Bouček 1992
Encyrtidae 65 102 265 163 all ecozones Trjapitzin and Gordh 1978a, b, Gordh 1979, Noyes and Woolley 1994
Eucharitidae 10 7 6 0 all except Arctic Burks 1979h, Heraty 1985, 2002
Eulophidae 13 173 373 1373 1000 all ecozones Burks 1979h, Yoshimoto 1983, Peck 1985, Schauff 1991, LaSalle 1994, Triapitsyn and Headrick 1995, Hansson 1987, 1988, 1989, 1994a, b, 1995, 1996a, b
Eupelmidae 14 29 24 0 south of Taiga ecozones Burks 1979g, Gibson 1989, 1995, 2002, 2010, 2011
Eurytomidae 14 0 90 107 17 southern Arctic and south Bugbee 1967, Burks 1979c, Zhang et al. 2017
Leucospidae 1 1 1 0 Pacific Maritime, Western Interior Basin, Prairies, Boreal Shield, Mixedwood Plains, Atlantic Maritime Bouček 1974, Burks 1979e
Megastigmidae 15 ? 21 2 0 southern Arctic and south Milliron 1949, Burks 1979c, Janšta et al. 2017
Mymaridae 15 93 369 276 all ecozones Burks 1979i, Huber and Fidalgo 1998, Huber and Lin 2000, Huber 2004, 2012, Triapitsyn et al. 2007, Triapitsyn 2017
Ormyridae 5 9 11 2 south of Taiga Cordillera and Arctic Burks 1979b, Hanson 1992
Perilampidae 11 20 25 5 south of Taiga ecozones Burks 1979b, Darling 1983, 1999, Darling and Miller 1991
Pteromalidae 110 293 697 404 all ecozones Burks 1979b, Heydon 1989, 1995, Heydon and LaBerge 1988, Darling 1991, Bouček 1993, Gibson and Vikberg 1998, Gibson 2000, 2003, 2009, Gibson and Floate 2001,
Signiphoridae 4 1 7 6 Mixedwood Plains Gordh 1979, Woolley 1988
Tetracampidae 2 4 4 0 Montane Cordillera, Boreal Shield, Mixedwood Plains, Atlantic Maritime Yoshimoto 1978, Bouček 1993
Torymidae 32 58 148 90 southern Arctic and south Grissell 1976, 1979, 1995, 2000, Janšta et al. 2017
Trichogrammatidae 6 34 113 79 southern Arctic and south Burks 1979j, Pinto 1999, 2004
Total Chalcidoidea 500 1210 3301 2135
Superfamily Mymarommatoidea16
Mymarommatidae 16 0 2 1 0 Boreal Shield, Mixedwood Plains, Atlantic Maritime Gibson et al. 2007, Huber et al. 2008
Superfamily Ceraphronoidea
Ceraphronidae 35 26 275 249 southern Arctic and south Dessart 1975, Muesebeck 1979, Dessart and Cancemi 1987, Johnson and Musetti 2004
Megaspilidae 35 21 101 80 southern Arctic and south Muesebeck 1979, Dessart 1981, 1987, Dessart and Cancemi 1987, Johnson and Musetti 2004
Total Ceraphronoidea 70 47 376 329
Superfamily Cynipoidea
Cynipidae 17 110 62 133 71 southern Arctic and south Burks 1979a, Shorthouse and Ritchie 1984, Ritchie 1993, Melika and Abrahamson 2002, Ronquist et al. 2015
Figitidae 18 36 60 620 560 all ecozones Burks 1979a, Ferrer-Suay et al. 2012, 2014, Ritchie 1993, Ros-Farré and Pujade-Villar 2009, 2011, 2013
Ibaliidae 4 4 2 0 all except Arctic Burks 1979a, Liu and Nordlander 1992, 1994, Ritchie 1993
Liopteridae 0 1 0 0 Mixedwood Plains Burks 1979a, Ritchie 1993, Liu et al. 2007
Total Cynipoidea 150 127 755 631
Superfamily Evanioidea
Aulacidae 17 18 6 0 southern Arctic and south Townes 1950, Carlson 1979a, Smith 2001, Deans et al. 2018
Evaniidae 4 4 2 0 Mixedwood Plains Townes 1949b, Carlson 1979a, Deans 2005, Deans et al. 2018
Gasteruptiidae 10 8 8 0 all except Arctic Townes 1950, Deans et al. 2018
Total Evanioidea 31 30 16 0
Superfamily Stephanoidea
Stephanidae 19 2 2 0 0 Pacific Maritime, Western Interior Basin, Mixedwood Plains Townes 1949a, Carlson 1979d, van Achterberg 2002, Aguiar 2004
Superfamily Trigonaloidea
Trigonalidae 20 4 4 4 0 Pacific Maritime, Boreal Shield, Mixedwood Plains, Atlantic Maritime Townes 1956, Carlson 1979b, Carmean 1995, Carmean and Kimsey 1998
Aculeata
Superfamily Chrysidoidea21
Bethylidae 35 27 83 56 all except Arctic Evans 1978, Krombein 1979a,
Chrysididae 22 33 84 101 17 all except Arctic Krombein 1979a, Bohart and Kimsey 1982, Kimsey and Bohart 1991
Dryinidae 35 50 150 100 all except Arctic Krombein 1979a, Olmi 1984
Embolemidae 2 2 0 0 south of Taiga ecozones Krombein 1979a, Olmi 1995
Total Chrysidoidea21 105 163 334 173
Superfamily Apoidea
Apoidea: Apiformes Hurd 1979, Sheffield et al. 2017
Andrenidae 250 182 153 20–40* southern Arctic and south Bouseman and LaBerge 1979, LaBerge 1969, 1973, 1977, 1980, 1986, 1987, 1989, LaBerge and Bouseman 1970, LaBerge and Ribble 1972, 1975, Donovan 1977, Ribble 1968, 1974
Apidae 23 189 206 213 50* all ecozones Cockerell 1903, LaBerge 1956a, b, 1961, Mitchell 1962, Daly 1973, Rightmyer 2008, Williams et al. 2014
Colletidae 45 54 50 5–10* all ecozones Stephen 1954, Mitchell 1960, Snelling 1966a, b, 1970
Halictidae 110 200 184 15* all ecozones Mitchell 1960, Roberts 1972, 1973, McGinley 1986, Gibbs 2010, Dumesh and Sheffield 2012, Gibbs et al. 2013, Heron and Sheffield 2015
Megachilidae 150 210 209 20* All ecozones Sandhouse 1939; Michener 1938a, b, c, 1939, 1947; Timberlake 1943; Mitchell 1962; Rightmyer et al. 2010; Sheffield et al. 2011; Gonzalez and Griswold 2013
Melittidae 2 3 3 0 Montane Cordillera, Prairies, Mixedwood Plains, Atlantic Maritime Michez and Patiny 2005, Michez and Eardley 2007, Payette 2013, Sheffield and Heron 2018
Total Apiformes 746 855 812 110–135*
Apoidea: Spheciformes Bohart and Menke 1976, Krombein 1979d, Pulawski 2018, Sann et al. 2018
Ammoplanidae 24 ? 5 4 0 Western Interior Basin, Prairies, Mixedwood Plains Smith 2008, 2009
Ampulicidae 24 ? 2 0 0 Boreal Shield, Mixedwood Plains Kohl 1893, Rohwer 1917, Bradley 1934, Buck 2004
Astatidae 24 ? 19 12 0 southern Arctic and south Parker 1962, 1969, 1972, Steiner 1973, Finnamore 1982, 1997, Buck 2004, Ratzlaff 2016, Sheffield 2017
Bembicidae 24 ? 81 56 0 southern Arctic and south Parker 1917, Bradley 1920, Bohart and Horning 1971, Finnamore 1982
Crabronidae s. str.24 ? 181 127 0 southern Arctic and south Finnamore 1982, 1997, Bohart 1976, Leclercq 2000, 2006, 2008, 2012
Mellinidae 24 ? 2 2 0 Prairies, Mixedwood Plains Strickland 1947, Finnamore 1982, Buck 2004
Pemphredonidae 24 ? 53 58 0 southern Arctic and south Strickland 1947, Finnamore 1982, Buck 2004
Philanthidae 24 ? 56 26 0 southern Arctic and south Scullen 1965, 1968, Bohart 1966, Bohart and Grissell 1975
Psenidae 24 ? 34 27 7 southern Arctic and south Finnamore 1980, 1982, 1983, Buck 2004, Ratzlaff 2016
Sphecidae 225 64 50 0 southern Arctic and south Menke 1965
Total Spheciformes 225 497 362 7
Total Apoidea 971 1352 1174 117–142*
Vespoidea s. lat.25
Superfamily Formicoidea
Formicidae 139 212 302 90 southern Arctic and south Francoeur 1997, Glasier and Acorn 2014, Glasier et al. 2013, 2016, Canadian Endangered Species Conservation Council 2016, AntWeb 2018, Bolton 2018; J Heron pers. comm.
Superfamily Pompiloidea26
Mutillidae 27 30 26 13 0 south of Boreal Cordillera and Taiga ecozones except Pacific Maritime Krombein 1979b, Pitts 2007, Williams et al. 2012, Brothers and Lelej 2017
Pompilidae 150 107 107 0 southern Arctic and south Evans 1950, 1951a, b, Townes 1957, Krombein 1979c, Wasbauer and Kimsey 1985, Finnamore 1997, Paiero et al. 2010
Sapygidae 6 7 9 2 all except Arctic Krombein 1979b, Kurzenko 1996
Superfamily Scolioidea28
Scoliidae 2 4 2 0 Western Interior Basin, Mixedwood Plains Krombein 1979b, MacKay 1987
Superfamily Tiphioidea26
Sierolomorphidae 1 2 4 2 Pacific Maritime, Boreal Cordillera, Boreal Plains, Prairies Mixedwood Plains Evans 1961, Krombein 1979b, Finnamore 1997, Buck et al. 2005, Lelej and Mokrousov 2015
Tiphiidae 29 25 31 13 0 south of Boreal Cordillera and Taiga ecozones Allen 1966, 1968, 1971, Krombein 1979b, Kimsey and Wasbauer 2006
Superfamily Thynnoidea26
Chyphotidae 30 ? 1 0 4* Western Interior Basin, Prairies Mickel 1967, Krombein 1979b
Thynnidae 29 ? 3 3 2* south of Taiga ecozones except Pacific Maritime Pate 1947, Krombein 1979b, Kimsey 2009
Superfamily Vespoidea s. str.
Rhopalosomatidae 2 1 0 2* Mixedwood Plains Townes 1977a, Lohrmann et al. 2012
Vespidae 100 96 102 6 southern Arctic and south MacLachlan 1980, Buck et al. 2008, Buck et al. 2012, Kimsey and Carpenter 2012, Canadian Endangered Species Conservation Council 2016, R Longair pers. comm.
Total Vespoidea s. lat.25 455 490 555 108
Total Aculeata 1531 2005 2063 398–423
Total Hymenoptera 6028 8757 18,454 10,366–10,391

The survey of Masner et al. (1979) gave estimates on the number of undescribed or unrecorded species in Canada, with the caveat “Such estimates are especially difficult (because so many families of Hymenoptera are inadequately known)…” The same caveat applies to the current survey; however, the use of Barcode Index Numbers (BINs) (Ratnasingham and Hebert 2013) based on 2% or greater sequence divergence of DNA barcodes in the Barcode of Life Data System (BOLD) provides us with new tools to help estimate hitherto unrecorded species diversity for some groups. For the purposes of this assessment, rather than add a somewhat subjective value for each family, the numbers of unrecorded species for most families were calculated based on the number of known BINs minus the number of described recorded species. This method is used for all families except those for which ongoing revisionary studies or faunal surveys have indicated that the number of BINs is not a good estimate of the total number of unrecorded species in Canada (these values noted in Table 1 with an asterisk). The BIN totals are current as of July 2018, but are likely underestimated for most superfamilies (see discussion at end of Faunal Analysis section).

The distribution of Hymenoptera families across ecozones in Canada (Rankin et al. 2011) is incompletely known. The species data in the checklists on which Table 1 is based were sorted by political unit, not ecozone, and it was not possible to go back and determine precise ecozones based on specimen locality data for all taxa (although this was done for the smaller families). In some cases, such as for the families of sawflies, knowledge of host plant distributions helped us make judgements on the ecozone ranges. For most families, there is no supporting information such as host distributions, habitats, or climatic ranges that can help discern whether range gaps are due to lack of sampling or whether a species is actually absent from an ecozone. Therefore, subjective decisions were made concerning whether to extrapolate the known range to encompass areas where there are sampling gaps. Most large Hymenoptera families span all the southern ecozones of Canada and some of them also range into the Arctic.

The information sources from which the data in Table 1 were taken is not exhaustive but instead contains the most important sources. Additional sources are noted for many taxa in the main body of text. In addition to the literature, specimens of all families in the Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Ottawa were also examined, providing a rich source of data to aid completion of Table 1. Family and superfamily level classification mainly follows Goulet and Huber (1993), except as noted in the footnotes of Table 1.

Overview of Hymenoptera diversity

The current study reports 8757 described species compared to approximately 6000 in 1979 (once omissions and overestimates in Masner et al. (1979) are taken into account). The approximately 2750 new records of Canadian Hymenoptera added since 1979 represent a 46% increase and an average of 71 new records/year. These figures indicate that Hymenoptera is one of the most diverse and relatively least known orders of insects in Canada. Masner et al. (1979) estimated 10,637 undescribed/unrecorded species which, when combined with the 6000 recorded species totalled 16,637 species for Canada. This means that in 1979, approximately 36% of the total estimated species were described/recorded. Currently, we estimate 10,366–10,391 undescribed/unrecorded species (Table 1) in an estimated total fauna of 19,148 species, of which approximately 46% are described/recorded.

Sawflies (previously suborder Symphyta)

All sawflies are herbivorous as larvae, except for members of the superfamily Orussoidea which are parasitoids of larval wood-boring Coleoptera and Hymenoptera (Middlekauff 1983, Goulet 1993). Sawflies occupy a paraphyletic grade at the base of the phylogeny of Hymenoptera, and Xyeloidea is consistently recovered as the sister group to all other Hymenoptera (Sharkey et al. 2012, Peters et al. 2017). Global estimates for species richness range between 8000 and 8300 species (Taeger et al. 2010, Huber 2017). Species of all seven superfamilies of sawflies and 12 of the 14 extant families, except Blasticotomidae and Megalodontesidae, are recorded in Canada. Masner et al. (1979) recorded eleven families in Canada; Anaxyelidae (Syntexis libocedrii Rohwer) was subsequently recorded from southern British Columbia (Goulet 1992). Furthermore, Masner et al. (1979) recorded 443 described species of sawflies, whereas the current survey has 710, an increase of 60.3% (Table 1). As in most parts of the world (especially the northern hemisphere), the sawfly fauna of Canada is dominated by the family Tenthredinidae, representing 75% or more of the total species.

Masner et al. (1979) estimated only 131 undescribed/unrecorded sawfly species, which was a significant underestimate as more than twice as many species (267) have been subsequently recorded from Canada. This large increase was a result of extensive work by many authors, most notably David R. Smith, who authored the Nearctic catalogue (Smith 1979a) and many revisions (e.g., Smith 1979b, 1989), and Henri Goulet (e.g., Goulet 1986, 1996). In addition, extensive collecting and faunal surveys (e.g., Goulet 1987) have contributed to our knowledge. A catalogue of world species (Taeger et al. 2010) and an accompanying searchable, electronic taxonomic database ECatSym (Blank et al. 2012) is also a rich source of information about all sawflies, including the Canadian fauna.

Examining DNA barcode data alone, it may appear that most of the diversity of Canadian sawflies has now been discovered as the number of BINs is lower than recorded species in all families except Siricidae, Cimbicidae, and Pergidae, all of which have fewer than 30 described species. This is likely misleading as surveys of Tenthredinidae in northern Europe, which has been surveyed much more intensively than northern North America, reveal a much greater diversity than northern Canada (e.g., Prous et al. 2017), suggesting that the Canadian fauna includes many more species than currently known. Ongoing revisions indicate that at least 200 undescribed species of Tenthredinidae are present in Canada, mostly in the subfamilies Nematinae and Tenthredininae (H Goulet pers. comm.) which emphasizes the fact that more collecting and DNA barcoding of Canadian sawflies is required.

Apocrita

Ichneumonoidea

In terms of described species, Ichneumonoidea is the largest superfamily of Hymenoptera, both in Canada (4202 species) (Table 1) and the world (47,177 species) (Yu et al. 2016). It is also the largest superfamily of insects in Canada comprising an impressive 10.8% of the 38,925 described insect species recorded (Langor 2019). There are two families, Ichneumonidae and Braconidae. Almost all ichneumonoids are parasitoids of other insects (Wahl and Sharkey 1993), the exceptions being a few genera that are parasitoids of spiders or prey on arachnid eggs (Townes 1969) and a few that are known to be phytophagous (e.g., Marsh 1991). The electronic catalogue of Ichneumonoidea (Yu et al. 2016) is an invaluable resource for accessing knowledge of the superfamily, including taxonomy, nomenclature, distribution, biology, references, etc.

Braconidae is the second largest family of Hymenoptera in Canada (1165 described species recorded; Table 1) and the world (21,221 described species; Yu et al. 2016). The current total is a 40.4% increase over the 830 species reported by Masner et al. (1979). Masner et al. (1979) estimated that there were 3200 undescribed/unrecorded braconid species in Canada. The number of BINs of Braconidae recorded in Canada in BOLD is 3411 (2246 more BINs than the number of described species recorded in Canada). Therefore, even though the estimate of undescribed Canadian braconid species by Masner et al. (1979) may appear to be a slight overestimate, studies on the percentage of undescribed microgastrine braconids in Canada and elsewhere in the world (e.g., Rodriguez et al. 2013) indicate that the number of undescribed braconids in Masner et al. (1979) may be accurate or even conservative. Good progress has been made on Canadian Braconidae since Masner et al. (1979) including a catalogue of all Nearctic species (Marsh 1979a, b) and keys to all New World genera (Wharton et al. 1997).

There are 25,285 described species of Ichneumonidae worldwide (Yu et al. 2016) and the actual fauna is estimated to be greater than 100,000 species (Gauld et al. 2002). Based on described, recorded species, Ichneumonidae is the most speciose family in Canada with 3037 species (Table 1) which represents approximately 35% of all described species of Hymenoptera recorded in Canada and 7.8% of all described species of insects (Langor 2019). The number of described, recorded species of ichneumonids reported in Masner et al. (1979) was 2001, including one species of “Pachylommatidae”, now called Hybrizontinae, that is considered part of Ichneumonidae (Sharkey and Wahl 1992). Since 1979, the number of described species of ichneumonids recorded in Canada has increased by 1036 (51.8%). Most of this increase was because of major revisions of Nearctic taxa (e.g., Dasch 1979, 1984, 1988, 1992, Townes 1983, Townes et al. 1992) as well as increased distributional knowledge via faunal surveys (e.g., Schwarzfeld 2014).

Masner et al. (1979) estimated that there were 5000 undescribed or unrecorded species of ichneumonids in Canada, but no discussion was provided to justify the estimate. There are 4748 known BINs for Canadian Ichneumonidae, ~1700 more BINs than recorded species (Table 1). Comparing the number of BINs to the estimated total number of species from Masner et al. (1979) (2001 known + 5000 anticipated = 7001), the current study has a shortfall of 2253 species. Whether an additional 2253 species of Canadian Ichneumonidae await discovery is unclear, but certainly, considering the very high diversity of Ichneumonidae in northern latitudes (e.g., 97 morphospecies recorded from Ellesmere Island, Nunavut; Timms et al. 2013), it is likely that many more species of Ichneumonidae remain to be collected and DNA barcoded in the less well-sampled regions of Canada (especially the North).

Diaprioidea

Historically, Diaprioidea was grouped within Proctotrupoidea (e.g., Masner et al. 1979, Muesebeck 1979); however, Sharkey (2007) found that Proctotrupoidea was polyphyletic and therefore removed Diapriidae and related families to a new superfamily. Diaprioidea includes four families (Sharkey 2007, Sharkey et al. 2012) of which Diapriidae is the most speciose, with 2048 species reported by Huber (2017), the other three families accounting for an additional 61 species. Two families are known in Canada, Diapriidae and Diapriidae (Table 1). Masner et al. (1979) considered Diapriidae part of Diapriidae, but the phylogenetic analysis of Sharkey et al. (2012) raised the subfamily Ismarinae to family status within Diaprioidea. Almost all Diaprioidea, for which the biology is known, are parasitoids of Diptera, although Diapriidae are hyperparasitoids of Dryinidae (Hymenoptera) parasitizing leafhoppers (Hemiptera: Cicadellidae) (Masner 1993a).

Masner et al. (1979) recorded 150 described species of Diapriidae (including Diapriidae) in Canada. The current study records 177 described species of Diapriidae and eight Diapriidae, which together is a 23.3% increase from 1979. The ratio of BINs to described species is 4.1 (763 BINs) implying that many undescribed/unrecorded species of diaprioids occur in Canada. Despite the relatively poor state of knowledge of Diaprioidea in Canada, there are some valuable resources on the group including keys to the New World genera of Diapriinae (Masner and García 2002) and a world catalogue with species distributions by biogeographical region (Johnson 1992). All information on Diapriidae has been updated and placed by N Johnson and colleagues on Hymenoptera Online (various contributors 2018), including additional distributional information and relevant literature. Masner (1976a) revised the world species of Diapriidae.

Platygastroidea

Masner et al. (1979) and Muesebeck (1979) classified Platygastridae and Scelionidae within Proctotrupoidea, but later classifications (e.g., Masner and Huggert 1989) separated these two families from Proctotrupoidea to form Platygastroidea. Sharkey (2007) synonymized the two on the basis of paraphyly of Scelionidae with respect to Platygastridae. Almost all known Platygastroidea are egg parasitoids of a variety of insect orders as well as of spiders (Masner 1993b). Huber (2017) indicated that there are 5385 known species worldwide.

Masner et al. (1979) recorded 150 described species of Scelionidae and 100 Platygastridae in Canada. The current study records only 160 described species combined, which means that the numbers reported by Masner et al. (1979) included undescribed species. Masner et al. (1979) estimated that there may be up to 300 unrecorded species of Platygastroidea in Canada, i.e., more unrecorded species than recorded. The ratio of BINs to recorded species in the current study supports the fact that Platygastroidea are very poorly known in Canada and, in fact, this value (14.3) is the highest of any Hymenoptera superfamily in Canada (2287 BINs). Based on this, there may be more than 2,100 undescribed/unrecorded species of Platygastroidea in Canada, making it the third largest superfamily of Hymenoptera in the country (after Ichneumonoidea and Chalcidoidea), and easily the most poorly known. Despite the apparent dearth of knowledge of the group, there have been many studies of Platygastroidea since 1979, such as a world revision of Platygastridae s. str. (Townes and Townes 1981), the world catalogue of Johnson (1992) that included species in the former Scelionidae (but not Platygastridae s. str.), keys to world genera of Scelioninae (Masner 1976b) and many revisions of Nearctic genera (e.g., Masner 1983a, b, Ritchie and Masner 1983). All information on Platygastroidea is available on an extensive website devoted to the systematics of the superfamily (Johnson 2018) and much of this knowledge has also been uploaded into Hymenoptera Online (various contributors 2018).

Proctotrupoidea

Proctotrupoidea is comprised of eight families (Huber 2017), of which five are present in Canada (Table 1). Huber (2017) recorded 448 described species worldwide. Masner et al. (1979) placed Pelecinidae within its own superfamily but it is now classified within Proctotrupoidea (Johnson and Musetti 1999). Conversely, Masner et al. (1979) included Platygastridae, Scelionidae, and Diapriidae in Proctotrupoidea but the first two families now comprise Platygastroidea (Masner 1993b) and the latter is placed in Diaprioidea (Sharkey 2007).

All proctotrupoids are parasitoids. Species of Proctotrupidae have been reared from Coleoptera and Diptera (Masner 1993a). Pelecinidae parasitizes Scarabaeidae (Coleoptera) (Johnson and Musetti 1999), Heloridae has been reared from Chrysopidae (Neuroptera) (Townes 1977b), Roproniidae from a sawfly (Masner 1993a), and Vanhorniidae from Eucnemidae (Coleoptera) (Deyrup 1985).

There are 73 described species of Proctotrupoidea in Canada, compared to 66 reported in Masner et al. (1979) (Table 1). Proctotrupidae is the largest family with 67 species and the other four families have one or two species each. The ratio of BINs to described species for the superfamily is 0.92. The world catalogue by Johnson (1992) summarizes the species and regional distributions, and additional references on the superfamily can be found on Hymenoptera Online (various contributors 2018).

Chalcidoidea and Mymarommatoidea

Chalcidoidea is comprised of 23 extant families (Heraty et al. 2013, Janšta et al. 2017), of which 18 are present in Canada (Table 1). Masner et al. (1979) included Mymarommatidae (as Mymarommidae) as a family within Chalcidoidea, but Gibson (1986) removed Mymarommatidae from Chalcidoidea, and Noyes and Valentine (1989) were the first to treat the taxon as a superfamily. Chalcidoidea have been reared as parasitoids from a wide variety of insect orders as well as some Arachnida and the nematode family Anguinidae, but a few are predators (using more than one host to complete development) and some are phytophagous (Gibson 1993). The biology of Mymarommatoidea is unknown although one has been reared from a bracket fungus and most are collected in shady, moist areas such as deciduous forests (Gibson et al. 2007, Huber et al. 2008).

Chalcidoidea is one of the world’s most diverse superfamilies of organisms. More than 22,700 species are described (Huber 2017), but Heraty et al. (2013) estimated that there may be as many as 500,000 species worldwide. Masner et al. (1979) recorded 16 families of Chalcidoidea in Canada (not including Mymarommatidae) but neglected to include Eurytomidae in their treatment. Other differences include Elasmidae (Elasmus Westwood), now classified within Eulophidae (Gauthier et al. 2000), Azotidae (Ablerus Howard), now classified in its own family instead of within Aphelinidae (Heraty et al. 2013), and Megastigminae removed from Torymidae and classified as Megastigmidae (Janšta et al. 2017). Masner et al. (1979) recorded 500 described species of Chalcidoidea in Canada, whereas the current survey records 1210 (a 142% increase which is the largest percentage increase of new species records over that time period for any Hymenoptera superfamily in Canada). The great increase in the number of recorded Chalcidoidea is a reflection of the large amount of work that has been done on this group (see Table 1). Chapters in the Nearctic catalogue were provided by Burks 1979b-j, Gordh 1979, and Grissell 1979, including distributional ranges in Canada. A key to the genera of Nearctic Chalcidoidea was published by Gibson et al. (1997). These publications have facilitated biological and faunal distributional studies for many taxa. All of the literature and taxonomic information to date is freely available in an online catalog, Universal Chalcidoidea Database (Noyes 2017). Despite the great amount of progress made on Canadian Chalcidoidea since 1979, the number of BINs (3301) is 2.7 times the number of recorded species and, based on this number, it is estimated that an additional 2135 undescribed/unrecorded species occur in the country (Table 1). The most speciose families in Canada based on BINs are Eulophidae (1373), Pteromalidae (697), and Mymaridae (369).

Mymarommatoids are very small wasps with a body length less than 1 mm (Gibson 1993). Huber (2017) reported ten described species worldwide. In Canada, they are only recorded in the east (Gibson et al. 2007), but they are also known in Montana (Hatten et al. 2010); therefore their range likely spans Canada from west to east. Masner et al. (1979) recorded no described species of Mymarommatidae in Canada, but predicted one unrecorded species to be present. The current survey records two species in Canada (Table 1) and Huber et al. (2008) provided keys to the described Nearctic species. There is one BIN for Mymarommatoidea from Canada in BOLD.

Ceraphronoidea

Ceraphronoidea is comprised of two families, Ceraphronidae and Megaspilidae, and there are 603 described species worldwide (Huber 2017). All ceraphronoids are parasitoids, most usually of Diptera, or hyperparasitoids of Hymenoptera, but they have also been associated with Hemiptera, Thysanoptera, Lepidoptera, Neuroptera and Mecoptera (Masner 1993c). There are 47 recorded Canadian species of Ceraphronoidea (Table 1), but this is one of the most poorly studied groups of Hymenoptera which is reflected in a BIN to recorded species ratio of 8.0 (10.6 for Ceraphronidae and 4.8 for Megaspilidae). This implies that there are over 375 species of Ceraphronoidea in Canada, of which most (329) remain to be described/recorded. Masner et al. (1979) recorded 70 described species in Canada (35 for each family), but this number included undescribed species based on their knowledge of the literature and examination of specimens in the CNC. Muesebeck (1979) provided a catalogue for the Nearctic species with Canadian distributions and Johnson and Musetti (2004) published a world catalog with distributions by region. Dessart and Cancemi (1987) provided keys to genera.

Cynipoidea

Cynipoidea (gall wasps and allies) is another understudied group of Hymenoptera. There are approximately 3200 species described globally (Huber 2017). The biology of the superfamily is diverse, with Cynipidae being mostly phytophagous gall-makers (but also inquilines in galls of other insects), whereas species in other families are parasitoids (e.g., Ibaliidae on siricid and anaxyelid sawflies; eucoiline Figitidae on cyclorrhaphous Diptera) (Ritchie 1993, Ronquist 1999). The current study records 127 described species of Cynipoidea in Canada which is slightly fewer than the total (150) reported in Masner et al. (1979). The number of BINs of Cynipoidea is 755, which, if representative of the total number of species, means that there could be as many as 631 unrecorded species in Canada (Table 1). The BIN to described species ratio is 5.9 showing that Cynipoidea is the third most poorly known superfamily in Canada, after Platygastroidea and Ceraphronoidea.

The classification of the families of Cynipoidea was previously contentious, but appears to have been stabilized with recognition of five families worldwide (Ronquist 1999) of which four are recorded in Canada: Cynipidae, Figitidae (including the former Charipidae, Eucoilidae and Alloxystidae), Ibaliidae and Liopteridae (Ritchie 1993, Ronquist 1999). Liopteridae was not recorded from Canada in Masner et al. (1979), but one specimen (now lost) was collected near Hamilton, Ontario (Liu et al. 2007). There have been a few revisions and reviews since 1979 for Cynipidae (e.g., Melika and Abrahamson 2002, Ronquist et al. 2015). Figitidae is the largest family and has the most gaps in knowledge. Some subfamilies are well-studied, e.g., Aspiceratinae (Ros-Farré and Pujade-Villar 2009, 2011, 2013). A world catalogue is available for Charipinae (Ferrer-Suay et al. 2012) as are keys to Nearctic genera and a species checklist (Menke and Evenhuis 1991). In contrast, other subfamilies are lacking in revisions and literature, especially the diverse Eucoilinae. Up to date nomenclature and literature has been added to Hymenoptera Online (various contributors 2018) but the last Nearctic catalogue with distributional data for the entire superfamily was Burks (1979a).

Evanioidea

Evanioidea (ensign wasps and allies) is a small superfamily with 1130 species globally (Huber 2017) that, for Canada at least, appears to be relatively well-known. This is mainly because the group is mostly tropical and only a few genera and species have ranges that extend to northern latitudes. There are three families, all of which are present in Canada: Aulacidae, Evaniidae (ensign wasps), and Gasteruptiidae. Aulacidae are parasitoids of wood-boring Coleoptera and sawflies, Evaniidae lay their eggs in the oothecae of cockroaches, and Gasteruptiidae have been reared from nests of solitary bees or wasps where they prey on one or more eggs or larvae (Mason 1993). There are 30 described species of Evanioidea in Canada, compared to 31 reported by Masner et al. 1979 (Table 1). The ratio of BINs to recorded species is only 0.53 (16 BINs vs 30 recorded species) which shows that more DNA barcode sampling is required. This is most evident for Evaniidae for which only two Canadian DNA barcodes are present in BOLD, despite four recorded species. Few or no unrecorded Canadian species of Evanioidea are expected. In terms of literature, keys to the Nearctic species are available, for Aulacidae (Townes 1950), Gasteruptiidae (Townes 1950, Smith 1996) and Evaniidae (Townes 1949). Carlson (1979a) provided the Nearctic catalogue for Evanioidea, Deans (2005) updated the nomenclature for Evaniidae, and Smith (2001) published a world catalogue of Aulacidae. Up to date information about Evanioidea is available at Evanioidea Online (Deans et al. 2018).

Stephanoidea

Stephanoidea is a small, mostly tropical group of Hymenoptera comprised of one family, Stephanidae; 342 species are known globally (Huber 2017). They are long, slender insects (body length up to 2 cm) that parasitize wood-boring Coleoptera (Mason 1993). There are two species in Canada (Table 1), the same number reported by Masner et al. (1979). One species is in the west and one in the east. There are keys to the Nearctic species (Townes 1949a) and no additional species are expected in Canada. An updated key to world genera is provided by van Achterberg (2002) and Aguiar (2004) published a world catalog including distributions by country. A summary of literature on the family is found online (Aguiar 2005). Stephanidae was included in Ichneumonoidea by Masner et al. (1979) but this classification is no longer commonly accepted (Aguiar 2005).

Trigonaloidea

Trigonaloidea, comprised of one family, Trigonalidae, lay eggs on leaves which are eaten by caterpillars or sawfly larvae. Except for some extralimital species which are primary parasitoids of pergid sawflies (Raff 1934), eggs of most trigonalid larvae do not continue development following ingestion unless the host is parasitized by an ichneumonoid wasp or tachinid fly or is captured by a vespid wasp (Carmean 1995). Globally there are 92 known species (Huber 2017). There are four species of Trigonalidae recorded in Canada (Table 1), the same number reported by Masner et al. (1979) and also four BINs from Canadian specimens in BOLD. Townes (1956) provided keys to the four Nearctic species. It is unlikely that additional species will be recorded from Canada.

Aculeata

Aculeata is a demonstrably monophyletic group (Branstetter et al. 2017) comprised of the superfamilies Chrysidoidea, Apoidea, and the assemblage of families that previously comprised the Vespoidea (hereafter called Vespoidea s. lat.). Aculeata includes many of the most recognizable groups of Hymenoptera, including the bees, ants, and vespid wasps. There are 2005 described species of Aculeata recorded in Canada which represents 22.9% of all recorded described Hymenoptera species (Table 1). Except for the Chrysidoidea, the group is relatively well-known based on the ratios of BINs to recorded species.

Chrysidoidea

Chrysidoidea includes 6780 species worldwide (Huber 2017) classified into seven families (Gauld and Hanson 1995), of which four are present in Canada (Table 1). They are parasitoids (or occasionally kleptoparasites) of a wide range of insect orders including Coleoptera, Lepidoptera, Hymenoptera, Phasmatodea, Embioptera (for the extralimital Sclerogibbidae), and Hemiptera (Finnamore and Brothers 1993, Gauld and Hanson 1995). Based on molecular data, they are hypothesized to be the sister group to the rest of Aculeata (Heraty et al. 2011, Peters et al. 2017) or a paraphyletic grade of two clusters of families at the base of Aculeata (Branstetter et al. 2017).

There are 163 described species of Chrysidoidea recorded in Canada, compared to 105 reported by Masner et al. (1979), a 55.2% increase. All Nearctic families have been revised since 1979. For Chrysididae (cuckoo wasps), a Nearctic revision was published (Bohart and Kimsey 1982) as well as a world review with species checklists (Kimsey and Bohart 1991). Olmi (1984) published a world revision of Dryinidae, with a supplement (Olmi 1991). The other major family in Canada, Bethylidae, was revised for the Nearctic by Evans (1978), and it appears that the number of described species in Canada reported by Masner et al. (1979) (35) was a slight overestimate of the number of Canadian species currently known (27 based on Evans (1978) and material in the CNC). Finally, Olmi (1995) revised the small family Embolemidae, but his revision did not change the number of species recorded in Canada (two). In terms of undiscovered diversity in Canada, the proportion of BINs to described species is 2.04 for the superfamily, indicating that there may be as many as 173 undescribed/unrecorded species of Chrysidoidea present in Canada, most of which belong to Bethylidae and Dryinidae.

Apoidea

Just under 30,000 described species of Apoidea are known globally (Huber 2017), with approximately two thirds representing the bees (Michener 2007). In total, the number of Apoidea species recorded for Canada has increased by approximately 39% since 1979 (1352 vs 971). Within this superfamily, the Spheciformes grade (Sphecidae sensu Masner et al. 1979) is now regarded as multiple families (Sann et al. 2018). The Crabronidae s. lat. was until recently the largest of the families, with many more than 400 species in Canada. However, the recent splitting of Crabronidae (Sann et al. 2018) resulted in several subfamilies being raised to family level as follows: Astatidae, Bembicidae, Crabronidae s. str. (previously Crabroninae), Mellinidae, Pemphredonidae, Philanthidae, and Psenidae. In addition, the subtribe Ammoplanina (previously in Pemphredoninae) was also raised to family status. Collectively, these eight families are represented by 431 species in 68 genera (Table 1). In addition to the families in the former Crabronidae, the eleven genera of Sphecidae s. str. are represented by 64 species, with a BIN to recorded species ratio of 0.78. Finally, there are two species of Ampulicidae (in two genera) from Canada but neither have been barcoded yet and no other species are expected in Canada (the only other two Nearctic species known are both from the southern United States (Krombein 1979d).

Classification of bees (Apiformes) has also changed since Masner et al. (1979), specifically with the merging of the non-corbiculate apid families Anthophoridae and carpenter bees (i.e., Xylocopidae sensu Masner et al. 1979) with the corbiculate apids (i.e., bumble bees and honey bee) into the single family Apidae (Michener 2000, 2007), resulting in six families of bees in Canada: Andrenidae, Apidae, Colletidae, Halictidae, Megachilidae, and Melittidae. Sheffield et al. (2017) recently summarized the bees of Canada, indicating that there were 855 species, though the number is likely higher when unique BINs without accompanying species-level identification are considered, especially for the poorly studied taxa Sphecodes Latreille (Halictidae), Nomada Scopoli (Apidae), and Osmia Panzer (Megachilidae).

Both Spheciformes and Apiformes are relatively well known; for the former, a global catalogue of species and distributional information, based on published literature, is well-maintained (Pulawski 2018). This resource, in addition to works published since 1979 (e.g., Finnamore 1983, 1997, Buck 2004, Ratzlaff 2016), has increased our knowledge of sphecid wasps (Sphecidae s. lat.), and was used to provide the summaries in Table 1. For Apiformes, Sheffield et al. (2017) provided a recent summary of Canadian species, including information on DNA barcodes, and an online catalogue for species is also available (Sheffield 2018). For bees, many revisions have occurred since Masner et al. (1979), specifically for the Canadian fauna (e.g., Gibbs 2010, Sheffield et al. 2011, Dumesh and Sheffield 2012, Onuferko 2017), or those that have included Canada in their coverage (e.g., Gibbs 2011, Rehan and Sheffield 2011, Gibbs et al. 2013).

Vespoidea s. lat.

Vespoidea s. lat. is comprised of all Aculeata that do not belong to Chrysidoidea or Apoidea (i.e., all superfamilies listed below), and is globally represented by more than 29,000 species (Huber 2017). Historically, the monophyly of the group has been equivocal. The catalogue of Hymenoptera of America North of Mexico (Krombein et al. 1979) divided the group into separate superfamilies, as did the survey of Masner et al. (1979), although these two studies differed slightly in the composition of several superfamilies. Later, morphology-based, cladistic analyses either refuted Vespoidea’s monophyly (e.g., Rasnitsyn 1988), or supported it (Brothers and Carpenter 1993). With the introduction of molecular data and a re-evaluation of the way in which characters were divided into states and polarized in earlier morphological studies (e.g., Brothers and Carpenter 1993), a consensus appears to have been reached that Vespoidea is not monophyletic, and alternative classifications have been suggested (e.g., Pilgrim et al. 2008). More recent molecular phylogenetic analyses (Branstetter et al. 2017, Peters et al. 2017) have also refuted the monophyly of Vespoidea but ambiguity still exists about the relationships of the taxa and how they relate to Apoidea, in particular because of differences in taxon choice between analyses and differences in topology correlated with differing phylogenetic methods. Because of this, the classification used here follows the suggested arrangement of Pilgrim et al. (2008) with the exception of Myrmosidae which is considered a subfamily of Mutillidae (Brothers and Lelej 2017). There are 490 described species of Vespoidea s. lat. recorded in Canada.

Formicoidea

Formicidae (the ants) was placed by itself in all molecular studies noted above, either as the sister group of Apoidea (Branstetter et al. 2017, Peters et al. 2017) or, in the preferred topology of Pilgrim et al. (2008), as sister group to Apoidea + Scolioidea. Formicidae is one of the great radiations of Hymenoptera with more than 16,000 described species (AntWeb 2018), but they are relatively poorly represented in Canada with only 212 described species recorded (Canadian Endangered Species Conservation Council 2016, J Heron pers. comm.), compared to 139 reported by Masner et al. (1979), a 52.5% increase (Table 1). In addition to the 2016 report on the conservation status of all Canadian species by province and territory, several regional checklists are available (Francoeur 1997 for the Yukon, Glasier and Acorn 2014 for the grasslands, Glasier et al. 2016 for Saskatchewan) as well as keys to workers of Alberta (Glasier et al. 2013). Our study records 302 BINs for ants (ratio to described species = 1.42), therefore there are likely ca. 90 additional species yet to be recorded in Canada. Considering the relatively good knowledge of ant taxonomy and distributional ranges, especially in northern latitudes, this is somewhat surprising, but it illustrates that even for supposedly well-known groups, our knowledge of the Canadian fauna is not complete. The ant taxonomic community is one of the most well-organized in entomology, with many resources including an online taxonomic and bibliographic catalogue (Bolton 2018) and an online database of specimen records, images and biological information (AntWeb 2018).

Pompiloidea

This group includes the velvet ants (Mutillidae), spider wasps (Pompilidae), and sapygid wasps (Sapygidae). Pompiloidea was not recognized by Masner et al. (1979). Instead, Pompilidae was placed with Vespidae in their Vespoidea s. str., and the other taxa were included in Scolioidea.

All pompiloids are parasitoids: Pompilidae on spiders (Day 1988), or in one case, a Phalangiidae (Opiliones) (Evans 1948); Mutillidae on other Aculeata, but also less commonly on Diptera, Lepidoptera, Coleoptera and Blattodea (Brothers and Finnamore 1993); and Sapygidae on bees and vespid wasps (Krombein 1979b). Some Pompilidae are kleptoparasitoids of other pompilids (Townes 1957).

The majority of Canadian diversity in this superfamily is in Pompilidae, with 107 of the 140 species (Table 1). Masner et al. (1979) listed 150 described species from Canada, but this appears to have been a slight overestimate. Pompilidae is a relatively poorly studied group in North America with only a few Nearctic faunal surveys since 1979 (e.g., Wasbauer and Kimsey 1985 for California, Finnamore 1997 for the Yukon, Sugar et al. 1999 for oak savannahs in southern Ontario). Nearctic identifications rely on the revisions of Evans (1950, 1951a, b) and Townes (1957). The most current Nearctic catalogue is Krombein (1979c), but many taxon names in this work are no longer valid and must be updated with reference to more recent, non-Nearctic catalogues (e.g., Wahis 1986, 2006). The ratio of BINs to recorded species is 1.0; however, as it is the second largest family in the Vespoidea s. lat. and it is relatively poorly studied, there could certainly be undescribed/unrecorded species.

Masner et al. (1979) recorded 30 described species of Mutillidae (including Chyphotidae) in Canada. The current study records 26 mutillids and one chyphotid (the latter now considered part of Thynnoidea). The number of recorded mutillids includes those of the subfamily Myrmosinae. This group was considered its own family by Pilgrim et al. (2008), but was moved back into Mutillidae by Brothers and Lelej (2017). There are only 13 BINs of Mutillidae from Canada on BOLD, and more sampling of this family is required. Mutillidae is a relatively well-studied family in North America with recent revisions of several major taxa (e.g., Pitts 2007, Williams et al. 2012); therefore, there is a good taxonomic foundation for surveying the Canadian fauna. Finally, Masner et al. (1979) recorded six species of Sapygidae and the current study has seven (Table 1), but there are nine BINs, therefore barcoded voucher specimens at Guelph need to be examined to determine which undescribed/unrecorded species may be present in Canada. Sapygidae is a relatively poorly studied family. Krombein (1979b) catalogued the Nearctic species including five Canadian species and Kurzenko (1996) provided a key to the Nearctic genera.

Scolioidea

Scolioidea, as defined by Pilgrim et al. (2008), consists of only one family, Scoliidae, in Canada (Krombein 1979b). Scolioidea of Masner et al. (1979) included six families (see footnote 28 in Table 1). Four species of Scoliidae are recorded from Canada, an increase from two species in Masner et al. (1979) (Table 1). All scoliids are parasitoids of Coleoptera, mostly Scarabaeoidea, but rarely Curculionoidea (Brothers and Finnamore 1993). Only two BINs have so far been recorded for Scoliidae from Canada, therefore more sampling is required. Historically, the classification of the family has been unstable, but there is now some consensus following publication of a world checklist (Osten 2005). There are 560 known species globally (Huber 2017). Very few studies on the Nearctic fauna have been done since the catalogue of Krombein (1979b), although MacKay (1987) treats the species of the southwestern US and has a key that includes all four species recorded in Canada.

Tiphioidea

Pilgrim et al. (2008) found that the family Tiphiidae was polyphyletic. The subfamilies Tiphiinae and Brachycistidinae clustered together, and therefore, these taxa were placed in Tiphiidae s. str. Their study related Tiphiidae s. str. to the monotypic family Sierolomorphidae, placing both families within Tiphioidea. Tiphioidea was not recognized by Masner et al. (1979). See Thynnoidea (below), for discussion of the placement of the other subfamilies previously belonging to Tiphiidae.

Tiphiidae are ectoparasitoids of Coleoptera (Brothers and Finnamore 1993). The current study records 31 species of Tiphiidae in Canada (Table 1). This compares to 25 species reported by Masner et al. (1979), a total which likely included one or more species that are now classified in Thynnidae. The ratio of BINs to recorded species is only 0.42, suggesting a need for more collecting and DNA barcoding. There have been no major revisions of Nearctic Tiphiinae since HW Allen’s efforts in the 1960s and 1970s (e.g., Allen 1966, 1971). Kimsey and Wasbauer (2006) provided a taxonomic checklist of the Brachycistidinae of the western Hemisphere.

Two species of Sierolomorphidae are currently recorded from Canada, up from one species reported by Masner et al. (1979); however, BOLD has four BINs from Canada for this family suggesting that undescribed/unrecorded species exist. Evans (1961) provided keys for the six Nearctic species. The hosts are unknown.

Thynnoidea

Phylogenetically, the five other subfamilies of Tiphiidae s. lat. (Kimsey 1991) clustered together in Pilgrim et al. (2008) and the valid name for this group is Thynnidae. Furthermore, Pilgrim et al. (2008) found that Thynnidae was the sister group of two subfamilies of Bradynobaenidae (Chyphotinae and Typhoctinae) which together, were raised to family status with the valid name Chyphotidae. Thynnoidea was not recognized by Masner et al. (1979).

In Canada, only three species of Thynnidae are recorded (Table 1), although a further two species are known (C Sheffield unpubl. data). Most Thynnidae are parasitoids of beetles (e.g., Methocinae on Cicindelinae), although one species of the extralimital subfamily Diamminae has been reared from mole crickets (Orthoptera: Gryllotalpidae) (Brothers and Finnamore 1993). Only three BINs are currently recorded for Thynnidae in Canada. Pate (1947) provided keys to the Nearctic genera.

One species of the family Chyphotidae is known from Canada (Mickel 1967) (Table 1). Little is known of the biology of Chyphotidae, but a species of the extralimital genus Typhoctes Ashmead has been found on immature Solifugae (Arachnidae) (Brothers and Finnamore 1993). Likely the Canadian species was included in Masner et al. (1979) as one of the species of Mutillidae recorded from Canada. There are no BINs for Chyphotidae from Canada. There are four species known from states bordering southwestern Canada (Mickel 1967), therefore more species are likely to occur in Canada.

Vespoidea s. str.

The analyses of Pilgrim et al. (2008) and Branstetter et al. (2017) found that Vespidae and Rhopalosomatidae are sister groups. In contrast, Vespoidea s. str. of Masner et al. (1979) was comprised of Vespidae and Pompilidae.

There are 96 species of Vespidae (yellow jackets, potter wasps, hornets, paper wasps, and allies) known from Canada (Canadian Endangered Species Conservation Council 2016, R Longair pers. comm.). The current number is slightly fewer than the number cited in the 2016 report (101) because the current list excludes several adventive species that are not considered to be established. Masner et al. (1979) recorded 100 species and the number of BINs is 102. The relative similarity of these totals illustrates the good level of knowledge that exists for Canadian Vespidae, especially for the northeastern Nearctic (Buck et al. 2008, 2012).

Rhopalosomatidae is a small family with only four genera worldwide (Brothers and Finnamore 1993). It has previously been proposed as the sister group of Pompilidae (Brothers 1999), related to Formicidae, Scoliidae, and Vespidae (Brothers and Carpenter 1993) or related to Mutillidae, Sapygidae, Scoliidae, Sierolomorphidae, and Tiphiidae (Masner et al. 1979). Very little is known of the biology of the family, and the only known hosts are crickets (Orthoptera: Gryllidae) (Townes 1977a). Only one species of Rhopalosomatidae is recorded from Canada, the brachypterous Olixon banksii (Brues) from southern Ontario (Lohrmann et al. 2012). Masner et al. (1979) listed two species from Canada, which we assume included Rhopalosoma nearcticum Brues, but we have not seen Canadian specimens of this species. It is recorded from Kentucky and Maryland, so its range could extend into Canada. A third genus, Liosphex Townes, is also recently recorded from Kentucky (L. boreus Lohrmann) (Lohrmann and Ohl 2010), therefore this genus may also be discovered in Canada in the future. There are no Canadian BINs for Rhopalosomatidae.

Faunal analysis

The results of the current survey have re-confirmed that Hymenoptera is one of the major constituents of biodiversity in Canada with 8,757 described species recorded (Table 1). The percentage of the Nearctic Hymenoptera fauna that is present in Canada cannot be determined precisely because Nearctic species totals have not been updated for some families since Krombein et al. (1979). However, total described species numbers for North America north of Mexico are known for two of the largest superfamilies, Ichneumonoidea and Chalcidoidea, which together comprise approximately two thirds of all described species recorded in Canada. Approximately 55% of Nearctic ichneumonoids are recorded in Canada (4202 of 7647) and approximately 34% of the chalcidoids (1210 of 3567). Together, 48.3% (5412 of 11,214) of these two superfamilies are recorded in Canada. If similar percentages exist for the remaining one third of species, then it can be estimated that roughly half of the described species of Hymenoptera in the Nearctic north of Mexico are recorded in Canada. At a global level, Canada has approximately 5.7% of the 153,410 described species of Hymenoptera in the world as tabulated by Huber (2017).

In terms of composition of the Hymenoptera of Canada, just over three quarters of the described, recorded species (77.3%) belong to three superfamilies: Ichneumonoidea (4202 species: 48.0%), Apoidea (1352 species: 15.4%) and Chalcidoidea (1210 species: 13.8%). The sawfly superfamily Tenthredinoidea is the fourth largest with 595 species (6.8% of total species) and Vespoidea s. lat. is fifth (490 species: 5.6%). The overall composition of Hymenoptera in Canada differs slightly if one considers total species (recorded species plus our estimates of unrecorded species). There are as many as 19,148 species with the following proportions: Ichneumonoidea (42.6%), Chalcidoidea (17.5%), Platygastroidea (12%), Apoidea (7.8%), and Tenthredinoidea (4.2%).

Canada’s Hymenoptera faunal structure is similar to other countries in northern latitudes. For example, Broad (2014) found the following proportions for described species recorded from Britain and Ireland: total species (7764), Ichneumonoidea (3913 species: 50.4 %), Chalcidoidea (1717 species: 22.1%), Tenthredinoidea (492 species: 6.4%), Apoidea (385 species: 4.9%) and Platygastroidea (362 species: 4.7%). The higher percentage of Apoidea recorded in Canada relative to Britain and Ireland is probably a reflection of greater diversity of habitats in Canada, especially hot, dry regions such as the Western Interior Basin, Prairies, and Mixedwood Plains ecozones which have a high diversity of Apoidea relative to cooler, more northern areas (Buck 2004, Sheffield et al. 2014). Relative to the whole world, Canada has a much higher percentage of described species of Ichneumonoidea (48.0% in Canada vs 30.8% for the whole world), slightly fewer Apoidea (15.4% vs 19.3%), approximately the same percentage of Chalcidoidea (13.8% vs 14.8%) and slightly more Tenthredinoidea (6.8% vs 4.7% worldwide). The higher percentage of Ichneumonoidea in northern latitudes compared to the tropics was discussed by previous authors (e.g., Janzen 1981, Gauld 1987), but more recent work on tropical ichneumonoids has demonstrated that this pattern is likely artefactual because of incomplete surveying of parasitoids in tropical areas of the world (Santos and Quicke 2011, Veijalainen et al. 2012, Timms et al. 2016). Apart from Ichneumonoidea, the other major difference between the composition of Hymenoptera in Canada compared to that of the entire world is the percentage of Vespoidea s. lat. (5.6% in Canada vs 19.0% in the entire world). Most vespoid families are predominantly tropical (Brothers and Finnamore 1993) and some, such as Chyphotidae, Rhopalosomatidae, Scoliidae, and Thynnidae have only one or a few species with ranges barely extending to southern Canada (see Table 1).

With respect to quantification of the number of introduced species of Hymenoptera in Canada, these numbers are available for some groups (e.g., sawflies, ants, bees and vespid wasps), but they are very poorly known for the parasitoid groups which encompass greater than 80% of the described species diversity of Hymenoptera in Canada. The reason for this lack of knowledge is a combination of poor distributional and taxonomic knowledge in many groups (both in Canada and elsewhere), as well as more than 100 years of well-meaning, but poorly documented, deliberate introductions of species for biological pest control that have obscured the native ranges of species in many groups. We can state that approximately 5% of sawflies appear to be introduced to Canada (H Goulet pers. comm.), and the Wild Species 2015 report (Canadian Endangered Species Conservation Council 2016) provided the following percentages: 7% of ants, just more than 2% of bees, and 5% of vespid wasps.

The 46% increase (8757 vs 6000) in recorded, described species since 1979 indicates that a great deal of work has been done in the last 39 years to document Canada’s Hymenoptera, but the high number of BINs (18,454) and estimated, unrecorded species (10,366–10,391) suggests that much more work is required as fewer than half of the total species are currently recorded. At the suprafamilial level, the following groups have had relatively few newly recorded species since 1979: Ceraphronoidea, Cynipoidea, Diaprioidea, Evanioidea, Platygastroidea, Proctotrupoidea, Stephanoidea, Trigonaloidea, bees (Apoidea: Apiformes), and Vespoidea s. lat. In contrast, other groups have had significant increases in the number of recorded species (values in parentheses are the percentage increases of recorded species in the current study compared to 1979): Chalcidoidea (142%), Apoidea: Spheciformes (121%), sawflies (60%), Chrysidoidea (55%), and Ichneumonoidea (48%). The great increase in the number of recorded species in these taxa indicates a relatively low level of taxonomic and distributional knowledge in 1979 coupled with a strong research effort since that time, especially publication of the Catalog of Hymenoptera of America North of Mexico (Krombein et al. 1979) and many revisionary studies in these groups (see references in Table 1).

Despite the great amount of research that has been performed on many groups of Canadian Hymenoptera, some groups require much more investigation, as indicated by high ratios of total estimated species (unrecorded plus recorded species) to recorded species: Platygastroidea (14.3), Ceraphronoidea (8.0), Cynipoidea (5.9), Diaprioidea (4.1), Chalcidoidea (2.7), Chrysidoidea (2.0), and Ichneumonoidea (1.9). All other suprafamilial taxa have ratios of approximately 1.2 or less, implying that they are relatively well documented in Canada. However, the ratio of total estimated species to recorded species is not the sole indicator of taxa in most need of taxonomic and survey work. Species richness must also be considered. For example, the ratio of total estimated species to recorded species for Ichneumonidae is 1.56 which is not even in the top ten ranking for families. However, in terms of the absolute number of unrecorded species estimated in this study, Ichneumonidae (1705 unrecorded species) ranks third behind only Braconidae (2246) and Platygastridae (2127).

Although we rely heavily on BIN data to estimate the number of undocumented species in most families, we realize that this approach may not provide good estimates of species richness in Canada for all families because of incomplete DNA barcoding libraries for some families and/or inability of DNA barcodes to distinguish all species correctly. Given the relatively short length of time that DNA barcoding has been in widespread use (Hebert et al. 2003), it is not surprising that some groups, especially those that are speciose in understudied regions, e.g., Ichneumonidae in the high Arctic (Timms et al. 2013), are not completely sampled and therefore are incompletely represented in the DNA barcode library. Also, there are taxa for which the DNA barcoding region of cytochrome oxidase I does not correctly distinguish all species. For example, 50–60% of 90 species of northwestern European sawflies of the genus Pristiphora Latreille could not be distinguished using DNA barcodes (Prous et al. 2017, and also see the general discussion on DNA barcoding of sawflies by Schmidt et al. 2017). Within the bees, Ceratina Latreille, Lasioglossum Curtis, and Bombus Latreille contain some problematic taxa in which multiple species share a single BIN (Sheffield et al. 2017), but these instances are rare and barcodes still permit identification to a sibling species pair or species group. In general, a large majority of hymenopteran species were able to be discriminated by barcoding in prior studies (e.g., 97.3% of European bees; Schmidt et al. 2015). In addition, hymenopteran specimens are notoriously difficult to barcode, exhibiting only a 65% recovery rate, roughly 30% lower than some orders like Lepidoptera and Diptera (Hebert et al. 2016). This poor barcode recovery is likely the product of their high adenine-thymine (AT) content (that complicates sequencing) and the demonstrated difficulties in PCR primer binding, both associated with the high rates of mitochondrial molecular evolution in Hymenoptera (Kaltenpoth et al. 2012). This low recovery rate compounds the challenge of comprehensively sampling the DNA of Hymenoptera across Canada, and thus underscores that estimation of the unknown Canadian fauna cannot rely on BINs alone. In summary, the percentage of the fauna that is documented (46%) may be under- or over-estimated; however, the actual percentage does not matter nearly as much as the stark fact that an enormous amount of work is required to document thousands of species that are hitherto unknown.

Acknowledgements

AMRB is grateful to colleagues at the CNC, especially D Barnes and E Maw for help collating data and references and G Gibson, H Goulet, J Huber, L Masner, S Cardinal, and R Longair for advice on species distributions and taxonomy. J Heron (BC Ministry of Environment) is thanked for inclusion of data from her Biota of Canada checklist of the ants of Canada. This chapter would not have been possible if not for the hard work of many summer students who helped search literature and format checklists during the past ten years. We are also grateful to the team at the Centre for Biodiversity Genomics (CBG, www.biodiversitygenomics.net) and the Barcode of Life Data Systems (BOLD, www.boldsystems.org) for sharing the infrastructure and platforms to acquire, access, and analyze DNA barcode data. Thanks also go to D Langor (Canadian Forest Service) for coordinating this very worthwhile project. Funding to AMRB was provided by internal operating grants by Agriculture and Agri-Food Canada. This chapter contributes to the University of Guelph’s Food from Thought research program supported by the Canada First Research Excellence Fund. CSS thanks colleagues at the Royal Saskatchewan Museum, especially R Oram for assisting in bee databasing initiatives, J Heron (BC Ministry of Environment and Climate Change Strategy) for Aculeata specimens from BC, and L Packer (York University); with funding in part from the Saskatchewan Ministry of Agriculture and the Canada-Saskatchewan Growing Forward 2 bi-lateral agreement, delivered by the Agriculture Council of Saskatchewan, and the Royal Saskatchewan Museum.

References

  • Allen HW (1966) A revision of the Tiphiinae (Hymenoptera: Tiphiidae) of eastern North America. Transactions of the American Entomological Society 92: 231–356.
  • Allen HW (1968) A monographic study of the genus Paratiphia. Transactions of the American Entomological Society 94: 25–109.
  • Allen HW (1971) A monographic study of the genus Tiphia (Hymenoptera: Tiphiidae) of Western North America. Transactions of the American Entomological Society 97: 211–275.
  • Blank SM, Groll EK, Liston AD, Prous M, Taeger A (2012) ECatSym – electronic world catalog of Symphyta (Insecta, Hymenoptera). Program version 4.0 beta, data version 39 (18.12.2012). Digital entomological information, Müncheberg. http://sdei.senckenberg.de/ecatsym/ [accessed 3.III.2018]
  • Bohart RM (1966) A review of Aphilanthops and related genera (Hymenoptera: Sphecidae). Proceedings of the Entomological Society of Washington 68: 158–167.
  • Bohart RM (1976) A review of the Nearctic species of Crabro (Hymenoptera: Sphecidae). Transactions of the American Entomological Society 102: 229–287.
  • Bohart RM, Grissell EE (1975) California wasps of the subfamily Philanthinae (Hymenoptera: Sphecidae). Bulletin of the California Insect Survey 19: 1–92.
  • Bohart RM, Horning DS (1971) California bembicine sand wasps. Bulletin of the California Insect Survey 13: 1–49.
  • Bohart RM, Kimsey LS (1982) A synopsis of the Chysididae in America north of Mexico. Memoirs of the American Entomological Institute 33: 1–266.
  • Bohart RM, Menke ASE (1976) Sphecid wasps of the world. A generic revision. University of California Press, Berkeley, 695 pp.
  • Bouček Z (1974) A revision of the Leucospidae (Hymenoptera: Chalcidoidea) of the world. Bulletin of the British Museum (Natural History) (Entomology) Supplement 23: 1–241.
  • Bouseman JK, LaBerge WE (1979) A revision of the genus Andrena of the western hemisphere. Part IX. Subgenus Melandrena. Transactions of the American Entomological Society 104: 275–389. http://www.jstor.org/stable/25078228
  • Bradley JC (1934) Records of Ampulicidae (Hymenoptera: Sphecoidea). Entomological News 44: 32–34.
  • Branstetter MG, Danforth BN, Pitts JP, Gates MW, Kula RR, Brady SG (2017) Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27(7): 1019–1025. https://doi.org/10.1016/j.cub.2017.03.027
  • Brothers DJ, Carpenter JM (1993) Phylogeny of Aculeata: Chrysidoidea and Vespoidea (Hymenoptera). Journal of Hymenoptera Research 2: 227–304.
  • Brothers DJ, Finnamore AT (1993) Superfamily Vespoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, 161–278.
  • Brothers DJ, Lelej AS (2017) Phylogeny and higher classification of Mutillidae (Hymenoptera) based on morphological reanalyses. Journal of Hymenoptera Research 60: 1–97. https://doi.org/10.3897/jhr.60.20091
  • Buck M (2004) An annotated checklist of the spheciform wasps of Ontario (Hymenoptera: Ampulicidae, Sphecidae and Crabronidae). Journal of the Entomological Society of Ontario 134: 19–84.
  • Buck M, Cobb TP, Stahlhut JK, Hanner RH (2012) Unravelling cryptic species diversity in eastern Nearctic paper wasps, Polistes (Fuscopolistes), using male genitalia, morphometrics and DNA barcoding, with descriptions of two new species (Hymenoptera: Vespidae). Zootaxa 3502: 1–48.
  • Buck M, Marshall SA, Cheung DKB (2008) Identification atlas of the Vespidae (Hymenoptera, Aculeata) of the northeastern Nearctic region. Canadian Journal of Arthropod Identification 5: 1–492. https://doi.org/10.3752/cjai.2008.05
  • Buck M, Paiero SM, Marshall SA (2005) New records of native and introduced aculeate Hymenoptera from Ontario, with keys to eastern Canadian species of Cerceris (Crabronidae) and Eastern Nearctic species of Chelostoma (Megachilidae). Journal of the Entomological Society of Ontario 136: 37–52.
  • Burdick DJ (1961) A taxonomic and biological study of the genus Xyela Dalman in North America. University of California Publications in Entomology 17: 285–355.
  • Burks BD (1979a) Cynipoidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1045–1107.
  • Burks BD (1979b) Pteromalidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 768–834.
  • Burks BD (1979c) Eurytomidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 835–859.
  • Burks BD (1979d) Chalcididae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 860–873.
  • Burks BD (1979e) Leucospidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 874.
  • Burks BD (1979f) Eucharitidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 875–877.
  • Burks BD (1979g) Eupelmidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 878–889.
  • Burks BD (1979h) Eulophidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 967–1021.
  • Burks BD (1979i) Mymaridae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1022–1032.
  • Burks BD (1979j) Trichogrammatidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1033–1041.
  • Canadian Endangered Species Conservation Council (2016) Wild Species 2015: The General Status of Species in Canada. National General Status Working Group, 128 pp. https://www.wildspecies.ca/home [accessed 1.II.2018].
  • Carlson RW (1979a) Evanioidea. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press Washington, DC, 1109–1118.
  • Carlson RW (1979b) Trigonaloidea. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press, Washington, DC, 1197–1198.
  • Carlson RW (1979c) Ichneumonidae. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press, Washington, DC, 315–739.
  • Carlson RW (1979d) Stephanidae. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press, Washington, DC, 740–742.
  • Carmean D (1995) Trigonalyidae. In: Gauld ID, Hanson PE (Eds) The Hymenoptera of Costa Rica. Oxford University Press, Oxford, 187–192.
  • Cockerell TDA (1903) North American bees of the genus Nomada. Proceedings of the Academy of Natural Sciences of Philadelphia 55: 580–614.
  • Daly HV (1973) Bees of the genus Ceratina in America north of Mexico (Hymenoptera: Apoidea). University of California Publications in Entomology 74: 1–113.
  • Darling DC (1983) A review of the New World species of Euperilampus (Hymenoptera; Chalcidoidea), with notes about host associations and phylogenetic relationships. Quaestiones Entomologicae 19: 1–40.
  • Darling DC (1991) Revision of the world species of Spalangiopelta (Hymenoptera: Chalcidoidea: Pteromalidae: Ceinae). Royal Ontario Museum Life Sciences Contributions 155: 1–43. https://doi.org/10.5962/bhl.title.53481
  • Darling DC (1999) Life history and immature stages of Stephanolampus salicetum (Hymenoptera: Chalcidoidea: Perilampidae). Proceedings of the Entomological Society of Ontario 130: 3–14.
  • Darling DC, Miller TD (1991) Life history and larval morphology of Chrysolampus (Hymenoptera: Chalcidoidea: Chrysolampinae) in western North America. Canadian Journal of Zoology 69: 2168–2177. https://doi.org/10.1139/z91-303
  • Dasch CE (1979) Ichneumon-flies of America north of Mexico: 8. Subfamily Cremastinae. Memoirs of the American Entomological Institute 29: 1–702.
  • Dasch CE (1984) Ichneumon-flies of America north of Mexico: 9. Subfamilies Theriinae and Anomaloninae. Memoirs of the American Entomological Institute 36: 1–610.
  • Dasch CE (1988) Ichneumon-flies of America north of Mexico: 10. Subfamily Banchinae, tribe Glyptini. Memoirs of the American Entomological Institute 43: 1–644.
  • Dasch CE (1992) The Ichneumon-flies of America North of Mexico: Part 12. Subfamilies Microleptinae, Helictinae, Cylloceriinae and Oxytorinae. Memoirs of the American Entomological Institute 52: 1–470.
  • Day MC (1988) Spider wasps (Hymenoptera: Pompilidae). Handbooks for the identification of British insects 6(4): 1–60.
  • Deans AR (2005) Annotated catalog of the world’s ensign wasp species (Hymenoptera: Evaniidae). Contributions of the American Entomological Institute 34(1): 1–164.
  • Deans AR, Yoder MJ, Dole K (2018) Evanioidea. Online-catalog of information about evanioid wasps (Hymenoptera). http://www.evanioidea.info [accessed 1.II.2018]
  • Delvare G, Bouček Z (1992) On the New World Chalcidinae (Hymenoptera). Memoirs of the American Entomological Institute 53: 1–466.
  • Dessart P (1975) Material typique des microhymenoptera myrmécophiles de la Collection Wasmann déposé au Muséum Wasmannianum à Maastricht (Pays-Bas). Publicaties van het Natuurhistorisch Genootschap in Limburg 24(1–2): 1–94.
  • Dessart P (1981) Le genre Megaspilus Westwood, 1829, en Amérique du Nord (Hymenoptera, Ceraphronoidea, Megaspilidae) Bulletin de l’Institut royal des Sciences naturelles de Belgique 53: 1–9.
  • Dessart P (1987) Revision des Lagynodinae (Hymenoptera: Ceraphronoidea: Megaspilidae). Bulletin de l’Institut royal des Sciences naturelles de Belgique 57: 5–30.
  • Dessart P, Cancemi P (1987) Tableau dichotomique des genres de Ceraphronoidea (Hymenoptera) avec commentaires et nouvelles espèces. Frustula Entomologica. Nouvelles Séries VII–VIII (XX–XXI): 307–372.
  • Deyrup M (1985) Notes on the Vanhorniidae (Hymenoptera). Great Lakes Entomologist 18: 65–68.
  • Donovan BJ (1977) A revision of North American bees of the subgenus Cnemidandrena (Hymenoptera: Andrenidae). University of California Publications in Entomology 81: 1–107.
  • Dumesh S, Sheffield CS (2012) Bees of the genus Dufourea Lepeletier (Hymenoptera: Halictidae: Rophitinae) of Canada. Canadian Journal of Arthropod Identification 20: 1–36. https://doi.org/10.3752/cjai.2012.20
  • Eidt DC (1969) The life histories, distribution, and immature forms of the North American sawflies of the genus Cephalcia (Hymenoptera: Pamphiliidae). Memoirs of the Entomological Society of Canada 59: 1–56. https://doi.org/10.4039/entm10159fv
  • Evans HE (1948) Biological notes on two species of Anoplius (Hymenoptera: Pompilidae). Entomological News 59: 180–184.
  • Evans HE (1950) [1949] A taxonomic study of the Nearctic spider wasps belonging to the tribe Pompilini (Hymenoptera: Pompilidae). Part I. Transactions of the American Entomological Society 75: 133–270.
  • Evans HE (1951a) [1950] A taxonomic study of the Nearctic spider wasps belonging to the tribe Pompilini (Hymenoptera: Pompilidae). Part II: genus Anoplius Dufour. Transactions of the American Entomological Society 76: 207–361.
  • Evans HE (1951b) A taxonomic study of the Nearctic spider wasps belonging to the tribe Pompilini (Hymenoptera: Pompilidae). Part III. Transactions of the American Entomological Society 77: 203–330.
  • Evans HE (1961) A preliminary review of the Nearctic species of Sierolomorpha (Hymenoptera). Breviora, 140: 1–12.
  • Evans HE (1978) The Bethylidae of America north of Mexico. Memoirs of the American Entomological Institute 27: 1–332.
  • Ferrer-Suay M, Paretas-Martínez J, Selfa J, Pujade-Villar J (2012) Taxonomic and synonymic world catalogue of the Charipinae and notes about this subfamily (Hymenoptera: Cynipoidea: Figitidae). Zootaxa 3376: 1–92.
  • Ferrer-Suay M, Selfa J, Pujade-Villar J (2014) First records, new species and a key of the Charipinae (Hymenoptera: Cynipoidea: Figitidae) from the Nearctic region. Annals of the Entomological Society of America 107: 50–73. https://doi.org/10.1603/AN13077
  • Finnamore AT (1980) New species of Mimesa from North America (Hymenoptera: Pemphredonidae: Psenini). The Canadian Entomologist 112: 293–300. https://doi.org/10.4039/Ent112293-3
  • Finnamore AT (1982) The Sphecoidea of southern Quebec (Hymenoptera). Memoirs of the Lyman Entomological Museum and Research Laboratory 11: 1–348.
  • Finnamore AT (1983) Revision of the American species of Mimesa (Hymenoptera: Sphecidae: Pseninae). Memoirs of the Lyman Entomological Museum and Research Laboratory 12: 1–171.
  • Finnamore AT (1997) Aculeate wasps of the Yukon (other than Formicidae). In: Danks HV, Downes JA (Eds) Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods) Monograph Series 2, Ottawa, Ontario, 867–900.
  • Finnamore AT, Brothers DJ (1993) Superfamily Chrysidoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada Publication, 1894/E, Ottawa, Ontario, 130–160.
  • Forbes AA, Bagley RK, Beer MA, Hippee AC, Widmayer HA (2018) Quantifying the unquantifiable: why Hymenoptera – not Coleoptera – is the most speciose animal order. bioRxiv 274431. https://doi.org/10.1101/274431
  • Francoeur A (1997) Ants (Hymenoptera: Formicidae) of the Yukon. In: Danks HV, Downes JA (Eds) Insects of the Yukon. Biological Survey of Canada (Terrestrial Arthropods) Monograph Series 2, Ottawa, Ontario, 901–910.
  • Gauld ID, Hanson PE (1995) The chrysidoid families. In: Gauld ID, Hanson PE (Eds) The Hymenoptera of Costa Rica. Oxford University Press, Oxford, 464–503.
  • Gauld ID, Sithole R, Gómez JU, Godoy C (2002) The Ichneumonidae of Costa Rica. 4. Introduction and keys to species of the subfamilies: Metopiinae and Banchinae. Memoirs of the American Entomological Institute 66: 1–768.
  • Gauthier N, Lasalle J, Quicke DLJ, Godfray HCJ (2000) Phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with a reclassification of Eulophinae and the recognition that Elasmidae are derived eulophids. Systematic Entomology 25: 521–539. https://doi.org/10.1046/j.1365-3113.2000.00134.x
  • Gibbs J (2010) Revision of the metallic species of Lasioglossum (Dialictus) in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa 2591: 1–382.
  • Gibbs J (2011) Revision of the metallic Lasioglossum (Dialictus) of eastern North America (Hymenoptera: Halictidae). Zootaxa 3073: 1–216.
  • Gibbs J, Packer L, Dumesh S, Danforth BN (2013) Revision and reclassification of Lasioglossum (Evylaeus), L. (Hemihalictus) and L. (Sphecodogastra) in eastern North America (Hymenoptera: Apoidea: Halictidae). Zootaxa 3672: 1–117. https://doi.org/10.11646/zootaxa.3672.1.1
  • Gibson GAP (1986) Evidence for monophyly and relationships of Chalcidoidea, Mymaridae and Mymarommatidae (Hymenoptera: Terebrantes). The Canadian Entomologist 118: 205–240. https://doi.org/10.4039/Ent118205-3
  • Gibson GAP (1989) Phylogeny and classification of Eupelmidae, with a revision of the world genera of Calosotinae and Metapelmatinae (Hymenoptera: Chalcidoidea). Memoirs of the Entomological Society of Canada 149: 1–121. https://doi.org/10.4039/entm121149fv
  • Gibson GAP (1993) Superfamilies Mymarommatoidea and Chalcidoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 570–655.
  • Gibson GAP (1995) Parasitic wasps of the subfamily Eupelminae: classification and revision of world genera (Hymenoptera: Chalcidoidea, Eupelmidae). Memoirs on Entomology, International 5: 1–421.
  • Gibson GAP (2000) Differentiation of the species of Urolepis (Hymenoptera: Chalcidoidea: Pteromalidae), potential biocontrol agents of filth flies (Diptera: Muscidae). The Canadian Entomologist 132: 391–410. https://doi.org/10.4039/Ent132391-4.
  • Gibson GAP (2003) Phylogenetics and classification of Cleonyminae (Hymenoptera: Chalcidoidea: Pteromalidae). Memoirs on Entomology, International 16: 1–339.
  • Gibson GAP (2009) Revision of New World Spalangiinae (Hymenoptera: Pteromalidae). Zootaxa 2259: 1–159.
  • Gibson GAP (2010) Calosota Curtis (Hymenoptera, Chalcidoidea, Eupelmidae) – review of the New World and European fauna including revision of species from the West Indies and Central and North America. ZooKeys 55: 1–75. https://doi.org/10.3897/zookeys.55.490
  • Gibson GAP (2011) The species of Eupelmus (Eupelmus) Dalman and Eupelmus (Episolindelia) Girault (Hymenoptera: Eupelmidae) in North America north of Mexico. Zootaxa 2951: 1–97.
  • Gibson GAP, Floate K (2001) Species of Trichomalopsis (Hymenoptera: Pteromalidae) associated with filth flies (Diptera: Muscidae) in North America. The Canadian Entomologist 133: 49–85. https://doi.org/10.4039/Ent13349-1.
  • Gibson GAP, Huber JT, Woolley JB (Eds) (1997) Annotated keys to the genera of Nearctic Chalcidoidea (Hymenoptera). National Research Council Research Press, Ottawa, Ontario, 794 pp.
  • Gibson GAP, Read J, Huber JT (2007) Diversity, classification and higher relationships of Mymarommatoidea (Hymenoptera). Journal of Hymenoptera Research 16: 51–146.
  • Gibson GAP, Vikberg V (1998) The species of Asaphes Walker from America North of Mexico, with remarks on extralimital distributions and taxa (Hymenoptera: Chalcidoidea, Pteromalidae). Journal of Hymenoptera Research 7(2): 209–256.
  • Glasier JRN, Acorn JH (2014) Annotated list of ants (Hymenoptera: Formicidae) from the grasslands of Alberta and Saskatchewan. In: Giberson DJ, Cárcamo HA (Eds) Arthropods of Canadian Grasslands (Volume 4): Biodiversity and Systematics Part 2. Biological Survey of Canada, Ottawa, Ontario, 299–314.
  • Glasier JRN, Acorn JH, Nielsen SE, Proctor H (2013) Ants (Hymenoptera: Formicidae) of Alberta: a key to species based primarily on the worker caste. Canadian Journal of Arthropod Identification 22: 1–104. https://doi.org/10.3752/cjai.2013.22
  • Glasier JRN, Nielsen S, Acorn JH, Borysenko LH, Radtke T (2016) A checklist of ants (Hymenoptera: Formicidae) of Saskatchewan. The Canadian Field-Naturalist 130(1): 40–48. https://doi.org/10.22621/cfn.v130i1.1791
  • Gonzalez VH, Griswold TL (2013) Wool carder bees of the genus Anthidium in the Western Hemisphere (Hymenoptera: Megachilidae): diversity, host plant associations, phylogeny, and biogeography. Zoological Journal of the Linnean Society 168: 21–425. https://doi.org/10.1111/zoj.12017
  • Gordh G (1979) Encyrtidae. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 890–966.
  • Goulet H (1986) The genera and species of the Nearctic Dolerini (Symphyta: Tenthredinidae: Selandriinae): classification and phylogeny. Memoirs of the Entomological Society of Canada 118(S135): 1–208. https://doi.org/10.4039/entm118135fv
  • Goulet H (1987) Symphyta. In: The insects, spiders and mites of Cape Breton Highlands National Park. Agriculture Canada Biosystematics Research Centre Report 1, Ottawa, Ontario, 225–230.
  • Goulet H (1992) The genera and subgenera of the sawflies of Canada and Alaska: Hymenoptera: Symphyta. Insects and Arachnids of Canada Part 20. Agriculture Canada Publication 1876, Ottawa, Ontario, 235 pp.
  • Goulet H (1993) Superfamilies Cephoidea, Megalodontidae, Orussoidea, Siricoidea, Tenthredinoidea, and Xyeloidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada Publication 1894/E, Ottawa, Ontario, 101–129.
  • Goulet H (1996) Revision of the Nearctic species of the arcuata group of the genus Tenthredo with notes on the higher classification of the Tenthredinini (Hymenoptera, Symphyta, Tenthredinidae). Contributions of the American Entomological Institute 29(2): 1–135.
  • Goulet H, Boudreault C, Schiff NM (2015) Revision of the world species of Xeris Costa (Hymenoptera: Siricidae). Canadian Journal of Arthropod Identification 28: 1–127.
  • Goulet H, Huber J (1993) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 668 pp.
  • Grissell EE (1976) A revision of western Nearctic species of Torymus Dalman (Hymenoptera: Torymidae). University of California Publications in Entomology 79: 1–120.
  • Grissell EE (1979) Family Torymidae. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press, Washington, DC, 748–767.
  • Grissell EE (1995) Toryminae (Hymenoptera: Chalcidoidea: Torymidae): a redefinition, generic classification and annotated world catalogue of species. Memoirs on Entomology, International 2: 1–474.
  • Grissell EE (2000) A revision of New World Monodontomerus (Hymenoptera: Chalcidoidea: Torymidae). Contributions of the American Entomological Institute 32(1): 1–90.
  • Halstead JA (1990) Revision of Hockeria Walker in the Nearctic region with descriptions of males and five new species (Hymenoptera: Chalcididae). Proceedings of the Entomological Society of Washington 92(4): 619–640.
  • Hansson C (1987) Revision of the New World species of Chrysocharis Forster (Hymenoptera: Eulophidae). Entomologica Scandinavica (supplement) 31: 3–86.
  • Hansson C (1988) A revision of the genus Mestocharis and a review of the genus Grahamia (Hymenoptera, Eulophidae). Proceedings of the Entomological Society of Washington 90(1): 28–36.
  • Hansson C (1989) New World Species of Holcopelte and Ionympha (Hymenoptera: Eulophidae), with Descriptions of two new species. Proceedings of the Entomological Society of Washington 91(1): 59–65.
  • Hansson C (1994a) The classification of Chrysonotomyia Ashmead and Telopterus Silvestri (Hymenoptera: Eulophidae), with a review of the species in the Nearctic region. Proceedings of the Entomological Society of Washington 96: 665–673.
  • Hansson C (1994b) Re-evaluation of the genus Closterocerus Westwood (Hymenoptera: Eulophidae), with a revision of the Nearctic species. Entomologica Scandinavica 25: 1–25. https://doi.org/10.1163/187631294X00018
  • Hansson C (1995) Revision of the Nearctic species of Neochrysocharis Kurdjumov (Hymenoptera: Eulophidae). Entomologica Scandinavica 26: 27–46
  • Hansson C (1996a) Taxonomic revision of the Nearctic species of Omphale Haliday (Hymenoptera: Eulophidae). Entomologica Scandinavica Supplement 49: 1–78.
  • Hansson C (1996b) The status of the genera Asecodes Förster, Ionympha Graham and Teleopterus Silvestri (Hymenoptera: Eulophidae), with a review of Nearctic species. Entomologica Scandinavica 27: 159–167. https://doi.org/10.1163/187631296X00025
  • Hatten TD, Merz N, Johnson JB, Looney C, Ulrich T (2010) Note on occurrence of Mymaromella pala Huber and Gibson (Hymenoptera: Mymarommatidae) in Montana: a new state record. Western North American Naturalist 70: 567–569. https://doi.org/10.3398/064.070.0417 064.070.0417
  • Hebert PDN, Cywinska A, Ball SL, deWaard JR (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society B – Biological Sciences 270: 313–321. https://doi.org/10.1098/rspb.2002.2218
  • Hebert PDN, Ratnasingham S, Zakharov EV, Telfer AC, Levesque-Beaudin V, Milton MA, Pedersen S, Jannetta P, deWaard JR (2016) Counting animal species with DNA barcodes: Canadian insects. Philosophical Transactions of the Royal Society B – Biological Sciences 371(1702): 20150333. https://doi.org/10.1098/rstb.2015.0333
  • Heraty JM (1985) A revision of the Nearctic Eucharitinae (Hymenoptera: Chalcidoidea: Eucharitidae). Proceedings of the Entomological Society of Ontario 116: 61–103.
  • Heraty JM (2002) A revision of the genera of Eucharitidae (Hymenoptera: Chalcidoidea) of the world. Memoirs of the American Entomological Institute 68: 1–367.
  • Heraty JM, Burks RA, Cruaud A, Gibson GAP, Liljeblad J, Munro J, Rasplus J-Y, Delvare G, Janšta P, Gumovsky A, Huber J, Woolley JB, Krogmann L, Heydon S, Polaszek A, Schmidt S, Darling DC, Gates MW, Mottern J, Murray E, Dal Molin A, Triapitsyn S, Baur H, Pinto JD, van Noort S, George J, Yoder M (2013) A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics 29: 466–542. https://doi.org/10.1111/cla.12006
  • Heraty JM, Ronquist F, Carpenter JC, Hawks D, Schulmeister S, Dowling A, Murray D, Munro JB, Wheeler WC, Schiff N, Sharkey M (2011) Evolution of the hymenopteran megaradiation. Molecular Phylogenetics and Evolution 60: 73–88. https://doi.org/10.1016/j.ympev.2011.04.003
  • Heydon SL (1989) A review of the world species of Notoglyptus Masi (Hymenoptera: Pteromalidae). Proceedings of the Entomological Society of Washington 91: 112–123.
  • Heydon SL (1995) A review of the North American species of Thinodytes Graham and Mauleus Graham (Hymenotpera: Pteromalidae). Journal of Hymenoptera Research 4: 2–24.
  • Heydon SL, LaBerge WE (1988) A review of North American species of Sphegigaster north of Mexico and the biology of their hosts (Hymenoptera, Pteromalidae). Journal of the Kansas Entomological Society 61(3): 258–277.
  • Huber JT (2004) Review of the described Nearctic species of the crassicornis group of Anaphes s.s. (Hymenoptera: Mymaridae). Journal of the Entomological Society of Ontario 135: 3–86.
  • Huber JT (2012) Revision of Ooctonus (Hymenoptera: Mymaridae) in the Nearctic region. Journal of the Entomological Society of Ontario 143: 15–105.
  • Huber JT, Fidalgo P (1998) [1997] Review of the genus Stephanodes (Hymenoptera: Mymaridae). Proceedings of the Entomological Society of Ontario 128: 27–63.
  • Huber JT, Gibson GAP, Bauer LS, Liu H, Gates M (2008) The genus Mymaromella (Hymenoptera: Mymarommatidae) in North America, with a key to described extant species. Journal of Hymenoptera Research 17: 175–194.
  • Huber JT, Lin N-Q (2000) [1999] World review of the Camptoptera group of genera (Hymenoptera: Mymaridae). Proceedings of the Entomological Society of Ontario 130: 21–65.
  • Hurd PD (1979) Apoidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1741–2209.
  • Janšta P, Cruaud A, Delvare G, Genson G, Heraty J, Křížková B, Rasplus J-V (2017) Torymidae (Hymenoptera, Chalcidoidea) revised : molecular phylogeny, circumscription and reclassification of the family with discussion of its biogeography and evolution of life-history traits. Cladistics (2017): 1–25 [online]. https://doi.org/10.1111/cla.12228
  • Janzen DH (1981) The peak in North American ichneumonid species richness lies between 38 degrees and 42 degrees N. Ecology 62: 532–537. https://doi.org/10.2307/1937717
  • Jarvis TD (1907) A preliminary list of the scale insects of Ontario. Annual Report of the Entomological Society of Ontario 38: 50–72.
  • Johnson NF (1992) Catalog of world Proctotrupoidea excluding Platygastridae. Memoirs of the American Entomological Institute 51: 1–825.
  • Johnson NF, Musetti L (1999) Revision of the proctotrupoid genus Pelecinus Latreille (Hymenoptera: Pelecinidae). Journal of Natural History 33: 1513–1543. https://doi.org/10.1080/002229399299879
  • Johnson NF, Musetti L (2004) Catalog of systematic literature of the superfamily Ceraphronoidea (Hymenoptera). Contributions of the American Entomological Institute 33(2): 1–149.
  • Kaltenpoth M, Corneli PS, Dunn DM, Weiss RB, Strohm E, Seger J (2012) Accelerated evolution of mitochondrial but not nuclear genomes of Hymenoptera: new evidence from crabronid wasps. PLoS ONE 7: e32826. https://doi.org/10.1371/journal.pone.0032826
  • Kimsey LS (2009) Taxonomic purgatory: Sorting out the wasp genus Myzinum Latreille in North America (Hymenoptera, Tiphiidae, Myzininae). Zootaxa 2224: 30–50.
  • Kimsey LS, Bohart RM (1991) [1990] The chrysidid wasps of the world. Oxford University Press, New York, 652 pp.
  • Kimsey LS, Wasbauer MS (2006) Phylogeny and checklist of the nocturnal tiphiids of the Western Hemisphere (Hymenoptera: Tiphiidae: Brachycistidinae). Journal of Hymenoptera Research 15: 9–25.
  • Kohl FF (1893) Ueber Ampulex Jur. (s.l.) und die damit enger verwandten Hymenopteren-Gattungen. Annalen des K.K. Naturhistorischen Hofmuseums 8: 455–516.
  • Krombein KV (1979a) Bethyloidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1203–1252.
  • Krombein KV (1979b) Scolioidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1253–1322.
  • Krombein KV (1979c) Pompiloidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1523–1572.
  • Krombein KV (1979d) Sphecoidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1573–1740.
  • Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) (1979) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 2735 pp.
  • Kurzenko NV (1996) A new Nearctic genus of Sapygidae with a key to the Nearctic and Palaearctic genera (Hymenoptera, Sapygidae). Memoirs of the Entomological Society of Washington 17: 89–94.
  • LaBerge WE (1956a) A revision of the bees of the genus Melissodes in North and Central America. Part I (Hymenoptera, Apidae). University of Kansas Science Bulletin 37: 911–1194. https://doi.org/10.5962/bhl.part.24549
  • LaBerge WE (1956b) A revision of the bees of the genus Melissodes in North and Central America. Part II (Hymenoptera, Apidae). University of Kansas Science Bulletin 38: 533–578.
  • LaBerge WE (1961) A revision of the bees of the genus Melissodes in North and Central America. Part III (Hymenoptera, Apidae). University of Kansas Science Bulletin 42: 283–663. https://doi.org/10.5962/bhl.part.9821
  • LaBerge WE (1969) A revision of the bees of the genus Andrena of the western hemisphere. Part II. Plastandrena, Aporandrena, Charitandrena. Transactions of the American Entomological Society 95: 1–47.
  • LaBerge WE (1973) A revision of the bees of the genus Andrena of the western hemisphere. Part VI. Subgenus Trachandrena. Transactions of the American Entomological Society 99: 235–371.
  • LaBerge WE (1977) A revision of the bees of the genus Andrena of the western hemisphere. Part VIII. Subgenera Thysandrena, Dasyandrena, Psammandrena, Rhacandrena, Euandrena, Oxyandrena. Transactions of the American Entomological Society 103: 1–143.
  • LaBerge WE (1980) A revision of the bees of the genus Andrena of the western hemisphere. Part X. Subgenus Andrena. Transactions of the American Entomological Society 106: 395–525.
  • LaBerge WE (1986) A revision of the bees of the genus Andrena of the western hemisphere. Part XI. Minor subgenera and subgeneric key. Transactions of the American Entomological Society 111: 441–567.
  • LaBerge WE (1987) A revision of the bees of the genus Andrena of the western hemisphere. Part XII. Subgenera Leucandrena, Ptilandrena, Scoliandrena, and Melandrena. Transactions of the American Entomological Society 112: 191–248.
  • LaBerge WE (1989) A revision of the bees of the genus Andrena of the western hemisphere. Part XIII. Subgenera Simandrena and Taeniandrena. Transactions of the American Entomological Society 115: 1–56.
  • LaBerge WE, Bouseman JK (1970) A revision of the bees of the genus Andrena of the western hemisphere. Part III. Tylandrena. Transactions of the American Entomological Society 96: 543–605.
  • LaBerge WE, Ribble DW (1972) A revision of the bees of the genus Andrena of the western hemisphere. Part V. Gonandrena, Geissandrena, Parandrena, Pelicandrena. Transactions of the American Entomological Society 98: 271–358.
  • LaBerge WE, Ribble DW (1975) A revision of the bees of the genus Andrena of the western hemisphere. Part VII. Subgenus Euandrena. Transactions of the American Entomological Society 101: 371–446.
  • Langor DW (2019) The diversity of terrestrial arthropods in Canada. In: Langor DW, Sheffield CS (Eds) The Biota of Canada – A Biodiversity Assessment. Part 1: The Terrestrial Arthropods. ZooKeys 819: 9–40. https://doi.org/10.3897/zookeys.819.31947
  • Leclercq J (2000) Hyménoptères Sphécides Crabroniens des Amériques du genre Crossocerus Lepeletier & Brullé, 1835. Notes fauniques de Gembloux 40: 3–75.
  • Leclercq J (2006) Hyménoptères Crabroniens des Amériques du genre Lestica Billberg, 1820 (Hymenoptera: Crabronidae: Crabronini). Notes fauniques de Gembloux 59: 215–218.
  • Leclercq J (2008) Hyménoptères Crabroniens d’Amérique du genre Crabro Fabricius 1775 (Hymenoptera: Crabronidae Crabroninae). Faunistic Entomology 61: 75–85.
  • Leclercq J (2012) Hyménoptères Crabroniens d’Amérique du Nord et d’Amérique centrale du genre Entomognathus Dahlbom 1844 (Hymenoptera: Crabronidae Crabronini). Faunistic Entomology 65: 119–126.
  • Lelej AS, Mokrousov MV (2015) Sierolomorpha sogdiana spec. nov. from Central Asia, and a world catalogue of Sierolomorphidae (Hymenoptera). Zootaxa 4018(4): 593–599.
  • Liu Z, Nordlander G (1992) Ibaliid parasitoids of siricid woodwasps in North America: Two new Ibalia species and a key to species (Hymenoptera: Cynipoidea). Proceedings of the Entomological Society of Washington 94: 500–507.
  • Liu Z, Nordlander G (1994) Review of the family Ibaliidae (Hymenoptera: Cynipoidea) with keys to genera and species of the world. Entomologica Scandinavica 25: 377–392. https://doi.org/10.1163/187631294X00153
  • Lohrmann V, Fox M, Solis M, Krogmann L (2012) Systematic revision of the New World Olixon Cameron with descriptions of O. melinsula sp. n. and the hitherto unknown female of O. bicolor (Hymenoptera, Rhopalosomatidae). Deutsche Entomologische Zeitschrift 59: 259–275.
  • MacKay WP (1987) The scoliid wasps of the southwestern United States (Hymenoptera: Scoliidae). The Southwestern Naturalist 32: 357–362. https://doi.org/10.2307/3671453
  • MacLachlan WB (1980) A key to and notes on Eumenes of America north of Mexico (Hymenoptera: Eumenidae). Journal of the Kansas Entomological Society 53: 617–621.
  • Marsh P (1979a) Family Braconidae. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press, Washington, DC, 144–294.
  • Marsh P (1979b) Family Aphidiidae. In: Krombein KV, Hurd Jr PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in American north of Mexico. Smithsonian Institution Press, Washington, DC, 295–312.
  • Marsh P (1991) Description of a phytophagous doryctine braconid from Brasil (Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington 93: 92–95.
  • Masner L (1976a) A Revision of the Ismarinae of the New World (Hymenoptera, Proctotrupoidea, Diapriidae). The Canadian Entomologist 108: 1243–1266. https://doi.org/10.4039/Ent1081243-11
  • Masner L (1976b) Revisionary notes and keys to world genera of Scelionidae (Hymenoptera: Proctotrupoidea). Memoirs of the Entomological Society of Canada 97: 1–87. https://doi.org/10.4039/entm10897fv
  • Masner L (1983a) A revision of Gryon Haliday in North America (Hymenoptera: Proctotrupoidea: Scelionidae). The Canadian Entomologist 115: 123–174. https://doi.org/10.4039/Ent115123-2
  • Masner L (1983b) Revision of the Nearctic species of Trichacis Foerster (Hymenoptera: Proctotrupoidea: Platygastridae). The Canadian Entomologist 115: 1071–1093. https://doi.org/10.4039/Ent1151071-9
  • Masner L (1991) Revision of Spilomicrus Westwood in America north of Mexico (Hymenoptera: Proctotrupoidea, Diapriidae). The Canadian Entomologist 123: 107–177. https://doi.org/10.4039/Ent123107-1
  • Masner L (1993a) Superfamily Proctotrupoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 537–557.
  • Masner L (1993b) Superfamily Platygastroidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 558–565.
  • Masner L (1993c) Superfamily Ceraphronoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 566–567.
  • Masner L, Barron JR, Danks HV, Finnamore AT, Francoeur A, Gibson GAP, Mason WRM, Yoshimoto CM (1979) Hymenoptera. In: Danks HV (Ed.) Canada and its Insect Fauna. Memoirs of the Entomological Society of Canada No. 108, 485–508. https://doi.org/10.4039/entm111108485-1
  • Masner L, Huggert L (1989) World review and keys to genera of the subfamily Inostemmatinae with reassignment of the taxa to the Platygastrinae and Sceliotrachelinae (Hymenoptera: Platygastridae). Memoirs of the Entomological Society of Canada No. 147, 214 pp. https://doi.org/10.4039/entm121147fv
  • Mason WRM (1993) Superfamilies Evanioidea, Stephanoidea, Megalyroidea and Trigonaloidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 510–520.
  • McGinley RJ (1986) Studies of Halictinae (Apoidea: Halictidae), I: revision of New World Lasioglossum Curtis. Smithsonian Contributions to Zoology 429: 1–294. https://doi.org/10.5479/si.00810282.429
  • Melika G, Abrahamson WG (2002) Review of the world genera of oak cynipid wasps (Hymenoptera: Cynipidae, Cynipini). In: Melika G, Thuróczy C (Eds) Parasitic Wasps: Evolution, Systematics, Biodiversity and Biological Control. Agroinform, Budapest, 150–190.
  • Menke ASE (1965) A revision of the North American Ammophila (Hymenoptera, Sphecidae). PhD Thesis, University of California, Davis, 247 pp.
  • Menke ASE, Evenhuis HH (1991) North American Charipidae: key to genera, nomenclature, species checklists, and a new species of Dilyta Förster (Hymenoptera: Cynipoidea). Proceedings of the Entomological Society of Washington 93: 136–158.
  • Michener CD (1938b) Bees of the genera Chelostomopsis, Formicapis, Robertsonella, and Prochelostoma (Hymenoptera: Megachilidae). Entomological News 69: 127–132.
  • Michener CD (1938c) American bees of the genus Chelostoma. Pan-Pacific Entomologist 14: 36–45.
  • Michener CD (1947) A revision of the American species of Hoplitis (Hymenoptera, Megachilidae). Bulletin of the American Museum of Natural History 89: 257–318.
  • Michener CD (2000) The bees of the world. Johns Hopkins University Press, Baltimore, 913 pp.
  • Michener CD (2007) The bees of the world. Second Edition. Johns Hopkins University Press, Baltimore, 953 pp.
  • Michez D, Eardley C (2007) Monographic revision of the bee genus Melitta Kirby 1802 (Hymenoptera: Apoidea: Melittidae). Annales de la Société entomologique de France (n. s. ) 43: 379–440.
  • Michez D, Patiny S (2005) World revision of the oil-collecting bee genus Macropis Panzer 1809 (Hymenoptera: Apoidea: Melittidae) with a description of a new species from Laos. Annales de la Société entomologique de France (n. s. ) 41: 15–25.
  • Mickel CE (1967) A review of the mutillid genus Chyphotes Blake (Hymenoptera; Mutillidae; Apterogyninae). Transactions of the American Entomological Society 93: 125–234.
  • Middlekauff WW (1958) The North American sawflies of the genera Acantholyda, Cephalicia, and Neurotoma (Hymenoptera: Pamphiliidae). University of California Publications in Entomology 14(2): 51–174.
  • Middlekauff WW (1964) North American sawflies of the genus Pamphilus (Hymenoptera: Pamphiliidae). University of California Publications in Entomology 38: 1–84.
  • Middlekauff WW (1983) Revision of the sawfly family Orussidae of North and Central America (Hymenoptera: Symphyta, Orussidae). University of California Publications Entomology 101: 1–46.
  • Milliron HE (1949) Taxonomic and biological investigations in the genus Megastigmus. American Midland Naturalist 41: 257–420. https://doi.org/10.2307/2421720
  • Mitchell TB (1960) Bees of the eastern United States. Vol. I. North Carolina Agricultural Experiment Station Technical Bulletin 141: 1–538.
  • Mitchell TB (1962) Bees of the eastern United States. Vol. II. North Carolina Agricultural Experiment Station Technical Bulletin 152: 1–557.
  • Muesebeck CFW (1979) Proctotrupoidea. In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 1121–1186
  • Noyes JS, Valentine EW (1989) Chalcidoidea (Insecta: Hymenoptera) – introduction and review of genera in smaller families. Fauna of New Zealand No. 18: 1–91.
  • Olmi M (1984) A revision of the Dryinidae (Hymenoptera), Parts 1 and 2. Memoirs of the American Entomological Institute 37: 1–1913.
  • Olmi M (1991) Supplement to the revision of the world Dryinidae (Hymenoptera, Chrysidoidea). Frustula Entomologica 12(1989): 371–456.
  • Olmi M (1995) A revision of the world Embolemidae (Hymenoptera Chrysidoidea). Frustula Entomologica 18(31): 85–146.
  • Osten T (2005) Checkliste der dolchwespen der welt (Insecta: Hymenoptera, Scoliidae). Bericht der Naturforschenden Gesellschaft Augsburg 62: 1–62.
  • Onuferko TM (2017) Cleptoparasitic bees of the genus Epeolus Latreille (Hymenoptera: Apidae) in Canada. Canadian Journal of Arthropod Identification 30: 1–62. https://doi.org/10.3752/cjai.2017.30
  • Paiero SM, Marshall SA, Pratt PD, Buck M (2010) Insects of Ojibway prairie, a southern Ontario tallgrass prairie. Arthropods of Canadian Grasslands 1: 199–225. https://doi.org/10.3752/9780968932148.ch9
  • Parker FD (1962) On the subfamily Astatinae, with a systematic study of the genus Astata of America north of Mexico (Hymenoptera: Sphecidae). Annals of the Entomological Society of America 55(6): 643–659. https://doi.org/10.1093/aesa/55.6.643
  • Parker FD (1969) On the subfamily Astatinae. Part VI. The American species in the genus Dryudella Spinola (Hymenoptera: Sphecidae). Annals of the Entomological Society of America 62(5): 963–976. https://doi.org/10.1093/aesa/62.5.963
  • Parker FD (1972) On the subfamily Astatinae. Part VII. The genus Diploplectron Fox (Hymenoptera: Sphecidae). Annals of the Entomological Society of America 65(5): 1192–1203. https://doi.org/10.1093/aesa/65.5.1192
  • Pate VSL (1947) A conspectus of the Tiphiidae with particular reference to the Nearctic forms. Journal of the New York Entomological Society 55: 115–145.
  • Payette A (2013) First record of the bee Melitta americana (Smith) (Hymenoptera: Melittidae) for Quebec and Canada. The Canadian Field Naturalist 127: 60–63. https://doi.org/10.22621/cfn.v127i1.1409
  • Peck O (1985) The taxonomy of the Nearctic species of Pediobius (Hymenoptera: Eulophidae), especially Canadian and Alaskan forms. The Canadian Entomologist 117: 647–704. https://doi.org/10.4039/Ent117647-6
  • Peters RS, Krogmann L, Mayer C, Donath A, Gunkel S, Meusemann K, Kozlov A, Podsiadlowski L, Petersen M, Lanfear R, Diez PA, Heraty J, Kjer KM, Klopfstein S, Meier R, Polidori C, Schmitt T, Liu S, Zhou X, Wappler T, Rust J, Misof B, Niehuis O (2017) Evolutionary history of the Hymenoptera. Current Biology 27(7): 1013–1018. https://doi.org/10.1016/j.cub.2017.01.027
  • Pilgrim EM, von Dohlen CD, Pitts JP (2008) Molecular phylogenetics of Vespoidea indicate paraphyly of the superfamily and novel relationships of its component families and subfamilies. Zoologica Scripta 37: 539–560. https://doi.org/10.1111/j.1463-6409.2008.00340.x
  • Pinto JD (1999) Systematics of the North American species of Trichogramma Westwood (Hymenoptera: Trichogrammatidae). Memoirs of the Entomological Society of Washington 22: 1–287.
  • Pinto JD (2004) A review of the genus Doirania Waterston (Hymenoptera: Trichogrammatidae), with a description of a new species from North America. Proceedings of the Entomological Society of Washington 106: 352–360.
  • Pitts JP (2007) Revision of Odontophotopsis Viereck (Hymenoptera: Mutillidae) Part 1, with a description of a new genus Laminatilla. Zootaxa 1619: 1–43.
  • Prous M, Kramp K, Vikberg V, Liston A (2017) North-western Palaearctic species of Pristiphora (Hymenoptera, Tenthredinidae). Journal of Hymenoptera Research 59: 1–190.
  • Raff JW (1934) Observations on saw-flies of the genus Perga, with notes on some reared primary parasites of the families Trigonalidae, Ichneumonidae and Tachinidae. Proceedings of the Royal Society of Victoria 47: 54–77.
  • Rankin R, Austin M, Rice J (2011) Ecological classification system for the ecosystem status and trends report. Canadian Biodiversity: Ecosystem Status and Trends 2010, Technical Thematic Report No. 1. Canadian Councils of Resource Ministers. Ottawa, Ontario, ii + 14 pp. http://www.biodivcanada.ca/default.asp?lang=En&n=137E1147-1
  • Rasnitsyn AP (1988) An outline of evolution of the hymenopterous insects (Order Vespida). Oriental Insects 22: 115–145.
  • Rehan SM, Sheffield CS (2011) Morphological and molecular delineation of a new species in the Ceratina dupla species-group (Hymenoptera: Apidae: Xylocopinae) of eastern North America. Zootaxa 2873: 35–50.
  • Ribble DW (1968) Revisions of two subgenera of Andrena: Micrandrena Ashmead and Derandrena, new subgenus (Hymenoptera: Apoidea). Bulletin of the University of Nebraska State Museum 8: 237–394.
  • Ribble DW (1974) A revision of the bees of the genus Andrena of the western hemisphere, subgenus Scaphandrena. Transactions of the American Entomological Society 100: 101–189.
  • Ries DT (1937) A revision of the Nearctic Cephidae (Hymenoptera: Tenthredinoidea). Transactions of the American Entomological Society 63: 259–324.
  • Rightmyer M (2008) A review of the cleptoparasitic bee genus Triepeolus (Hymenoptera: Apidae), Part I. Zootaxa 1710: 1–170.
  • Rightmyer M, Griswold T, Arduser M (2010) A review of the non-metallic Osmia (Melanosmia) found in North America, with additional notes on Palearctic Melanosmia (Hymenoptera, Megachilidae). ZooKeys 60: 37–77. https://doi.org/10.3897/zookeys.60.484
  • Ritchie AJ, Masner L (1983) Revision of the Nearctic species of Baryconus (Hymenoptera: Scelionidae, Scelioninae). Canadian Journal of Zoology 61: 704–720. https://doi.org/10.1139/z83-094
  • Ritchie AJ (1993) Superfamily Cynipoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 521–536.
  • Roberts RB (1972) Revision of the bee genus Agapostemon (Hymenoptera: Halictidae). University of Kansas Science Bulletin 49: 437–590.
  • Roberts RB (1973) Bees of northwestern America: Halictus (Hymenoptera: Halictidae). Agricultural Experiment Station, Oregon State University Technical Bulletin 126: 1–23.
  • Rodriguez JJ, Fernández-Triana J, Smith MA, Janzen DH, Hallwachs W, Erwin T, Whitfield JB (2013) Extrapolations from field studies and known faunas converge on dramatically increased estimates of global microgastrine parasitoid wasp species richness (Hymenoptera: Braconidae). Insect Conservation and Diversity 6: 530–536. https://doi.org/10.1111/icad.12003
  • Rohwer SA (1917) A Nearctic species of Dolichurus. Proceedings of the Entomological Society of Washington 18: 212–213.
  • Ronquist F, Nieves-Aldrey J-L, Buffington ML, Liu Z, Liljeblad J, Nylander JAA (2015) Phylogeny, Evolution and Classification of Gall Wasps: The Plot Thickens. PLoS ONE 10(5): e0123301. https://doi.org/10.1371/journal.pone.0123301
  • Rosen D, DeBach P (1979) Species of Aphytis of the world (Hymenoptera: Aphelinidae). Series Entomologica 17: 1–801.
  • Ros-Farré P, Pujade-Villar J (2009) Revision of the genus Callaspidia Dahlbom, 1842 (Hymenoptera: Figitidae: Aspicerinae). Zootaxa 2105: 1–31.
  • Ros-Farré P, Pujade-Villar J (2011) Revision of the genus Omalaspis Giraud, 1860 (Hym.: Figitidae: Aspicerinae). Zootaxa 2917: 1–28.
  • Ross HH (1955) The taxonomy and evolution of the sawfly genus Neodiprion. Forest Science 1: 196–209.
  • Sandhouse GA (1939) North American bees of the genus Osmia (Hymenoptera: Apoidea). Memoirs of the Entomological Society of Washington 1: 1–167.
  • Sann M, Niehuis O, Peters RS, Mayer C, Kozlov A, Podsiadlowski L, Bank S, Meusemann K, Misof B, Bleidorn C, Ohl M (2018) Phylogenomic analysis of Apoidea sheds new light on the sister group of bees. BMC Evolutionary Biology 18: 72. https://doi.org/10.1186/s12862-018-1155-8
  • Schauff ME (1991) The Holarctic genera of Entedoninae (Hymenoptera: Eulophidae). Contributions of the American Entomological Institute 26(4): 1–109.
  • Schauff ME, Evans GA, Heraty JM (1996) A pictorial guide to the species of Encarsia (Hymenoptera: Aphelinidae) parasitic on whiteflies (Homoptera: Aleyrodidae) in North America. Proceedings of the Entomological Society of Washington 98: 1–35.
  • Schiff NM, Goulet H, Smith DR, Boudreault C, Wilson AD, Scheffler BE (2012) Siricidae (Hymenoptera: Symphyta: Siricoidea) of the Western Hemisphere. Canadian Journal of Arthropod Identification 21: 1–305. https://doi.org/10.3752/cjai.2012.21
  • Schmidt S, Schmid-Egger C, Morinière J, Haszprunar G, Hebert PDN (2015) DNA barcoding largely supports 250 years of classical taxonomy: identifications for Central European bees (Hymenoptera, Apoidea partim). Molecular Ecology Resources 15: 985–1000. https://doi.org/10.1111/1755-0998.12363
  • Schmidt S, Taeger A, Morinière J, Liston A, Blank S, Kramp K, Kraus M, Schmidt O, Heibo E, Prous M, Nyman T, Malm T, Stahlhut J (2017) Identification of sawflies and horntails (Hymenoptera, ‘Symphyta’) through DNA barcodes: successes and caveats. Molecular Ecology Resources 17: 670–685. https://doi.org/10.1111/1755-0998.12614
  • Schwarzfeld MD (2014) Ichneumonidae (Hymenoptera) of the Canadian Prairies Ecozone: A Review. In: Giberson DJ, Cárcamo HA (Eds) Arthropods of Canadian Grasslands (Volume 4): Biodiversity and Systematics Part 2. Biological Survey of Canada, Ottawa, Ontario, 317–397.
  • Sharkey MJ (2007) Phylogeny and classification of Hymenoptera. Zootaxa 1668: 521–548.
  • Sharkey MJ, Carpenter JM, Vilhelmsen L, Heraty J, Liljeblad J, Dowling APG, Schulmeister S, Murray D, Deans AR, Ronquist F, Krogmann L, Wheeler WC (2012) Phylogenetic relationships among superfamilies of Hymenoptera. Cladistics 28: 80–112. https://doi.org/10.1111/j.1096-0031.2011.00366.x
  • Sharkey MJ, Wahl DB (1992) Cladistics of the Ichneumonoidea (Hymenoptera). Journal of Hymenoptera Research 1: 15–24.
  • Sheffield CS (2017) Diploplectron ferrugineum Ashmead, 1899 from Canada, with comments on the type locality (Hymenoptera: Crabronidae: Astatinae). The Pan-Pacific Entomologist 93: 159–162. https://doi.org/10.3956/2017-93.2.159
  • Sheffield CS, Frier SD, Dumesh S (2014) The bees (Hymenoptera: Apoidea, Apiformes) of the Prairies Ecozone, with comparisons to other grasslands of Canada. In Giberson DJ, Cárcamo HA (Eds) Arthropods of Canadian Grasslands (Volume 4): Biodiversity and Systematics Part 2. Biological Survey of Canada, Ottawa, Ontario, 427–467.
  • Sheffield CS, Heron J (2018) A new western Canadian record of Epeoloides pilosulus (Cresson), with discussion of ecological associations, distribution, and conservation status in Canada. Biodiversity Data Journal 5: e22837. https://doi.org/10.3897/BDJ.6.e22837
  • Sheffield CS, Heron J, Gibbs J, Onuferko TM, Oram R, Best L, deSilva N, Dumesh S, Pindar A, Rowe G (2017) Contribution of DNA barcoding to the study of the bees (Hymenoptera: Apoidea) of Canada: progress to date. The Canadian Entomologist 149: 736–754. https://doi.org/10.4039/tce.2017.49
  • Sheffield CS, Ratti C, Packer L, Griswold T (2011) Leafcutter and mason bees of the genus Megachile Latreille (Hymenoptera: Megachilidae) in Canada and Alaska. Canadian Journal of Arthropod Identification 18: 1–107. https://doi.org/10.3752/cjai.2011.18
  • Shirley X, Woolley JB, Hopper KR (2017) Revision of the asychis species group of Aphelinus Hymenoptera: Aphelinidae). Journal of Hymenoptera Research 54: 1–32. https://doi.org/10.3897/jhr.54.10457
  • Shorthouse JD, Ritchie AJ (1984) Description and biology of a new species of Diplolepis Fourcroy (Hymenoptera: Cynipidae) inducing galls on the stems of Rosa acicularis. The Canadian Entomologist 116: 1623–1636. https://doi.org/10.4039/Ent1161623-12
  • Skvarla MJ, Tripodi A, Szalanski A, Dowling A (2015) New records of Orussus minutus Middlekauf, 1983 (Hymenoptera: Orussidae) represent a significant western range expansion. Biodiversity Data Journal 3: e5793. https://doi.org/10.3897/BDJ.3.e5793
  • Smith DR (1969a) Key to genera of Nearctic Argidae (Hymenoptera) with revisions of the genera Atomacera Say and Sterictiphora Billberg. Transactions of the American Entomological Society 95: 439–457.
  • Smith DR (1969b) Nearctic Sawflies II. Selandriinae: Adults (Hymenoptera: Tenthredinidae). United States Agricultural Research Technical Bulletin 1398: 1–48.
  • Smith DR (1969c) Nearctic Sawflies I. Blennocampinae: Adults (Hymenoptera: Tenthredinidae). United States Agricultural Research Technical Bulletin 1397: 1–177.
  • Smith DR (1971a) Nearctic sawflies of the genera Neoptilia Ashmead, Schizocerella Forsius, Aprosthema Konow, and Sphacophilus Provancher (Hymenoptera: Argidae). Proceedings of the Entomological Society of Washington 76: 409–418.
  • Smith DR (1971b) Nearctic sawflies. III. Heterarthrinae: Adults and larvae (Hymenoptera: Tenthredinidae). Agricultural Research Service, United States Department of Agriculture Technical Bulletin 1420: 1–84.
  • Smith DR (1974) Conifer sawflies, Diprionidae – key to North American genera, checklist of world species, and new species from Mexico. Transactions of the American Entomological Society 97: 537–594.
  • Smith DR (1976) The xiphydriid woodwasps of North America (Hymenoptera: Xiphydriidae). Transactions of the American Entomological Society 102: 101–131.
  • Smith DR (1977) The fir shoot-boring sawflies of the genus Pleuroneura in North America (Hymenoptera: Xyelidae). Annals of the Entomological Society of America 70: 761–767. https://doi.org/10.1093/aesa/70.5.761
  • Smith DR (1979a) Symphyta In: Krombein KV, Hurd PD, Smith DR, Burks BD (Eds) Catalog of Hymenoptera in America North of Mexico. Smithsonian Institution Press, Washington, DC, 3–136.
  • Smith DR (1979b) Nearctic sawflies. IV. Allantinae: adults and larvae (Hymenoptera: Tenthredinidae). United States Department of Agriculture Technical Bulletin 1595: 1–172.
  • Smith DR (1986) The berry and rose stem-borers of the genus Hartigia in North America (Hymenoptera: Cephidae). Transactions of the American Entomological Society 112: 129–145.
  • Smith DR (1989) The sawfly genus Arge (Hymenoptera: Argidae) in the Western Hemisphere. Transactions of the American Entomological Society 115: 83–205.
  • Smith DR (1996) Review of the Gasteruptiidae (Hymenoptera) of eastern North America. Proceedings of the Entomological Society of Washington 98(3): 491–499.
  • Smith DR (2001) World catalog of the family Aulacidae (Hymenoptera). Contributions on Entomology, International 4(3): 263–319.
  • Smith DR (2006) Checklist of the Pergidae (Hymenoptera: Symphyta) of the Western Hemisphere, with a new genus and two new species. In: Blank SM, Schmidt S, Taeger A (Eds) Recent Sawfly Research: Synthesis and Prospects. Goecke & Evers, Keltern, 613–626.
  • Smith DR, Schiff NM (1998) The genera Macroxyela Kirby and Megaxyela Ashmead (Hymenoptera: Xyelidae) in North America. Proceedings of the Entomological Society of Washington 100: 636–657.
  • Smith DR, Schiff NM (2005) A new western Nearctic species of Calameuta Konow (Hymenoptera : Cephidae). Proceedings of the Entomological Society of Washington 107: 864–868.
  • Smith DR, Solomon NM (1989) A new Janus (Hymenoptera: Cephidae) from Quercus, and a key to North American species. Entomological News 100: 1–5.
  • Smith NJ (2008) A review of Nearctic Ammoplanus Giraud 1869 (Hymenoptera: Crabronidae). Pan-Pacific Entomologist 84(4): 301–333. https://doi.org/10.3956/2007-56.1
  • Smith NJ (2009) A review of the Nearctic species of Parammoplanus (Pate) with descriptions of new species (Hymenoptera: Crabronidae). Pan-Pacific Entomologist 85(3): 107–149. https://doi.org/10.3956/2009-02.1
  • Snelling RR (1966a) Studies on North American bees of the genus Hylaeus. 1. Distribution of the western species of the subgenus Prosopis with descriptions of new forms (Hymenoptera: Colletidae). Los Angeles County Museum, Contributions in Science 98: 1–18.
  • Snelling RR (1966b) Studies on North American bees of the genus Hylaeus. 3. The Nearctic subgenera (Hymenoptera: Colletidae). Bulletin of the Southern California Academy of Science 65: 164–175.
  • Snelling RR (1970) Studies on North American bees of the genus Hylaeus. 5. The subgenera Hylaeus, s. str. and Paraprosopis (Hymenoptera: Colletidae). Los Angeles County Museum, Contributions in Science 180: 1–59.
  • Steiner A (1973) Solitary wasps from subarctic North America – II. Sphecidae from the Yukon and Northwest Territories, Canada: distribution and ecology. Quaestiones Entomologicae 9: 13–34.
  • Stephen WP (1954) A revision of the genus Colletes in America north of Mexico. University of Kansas Science Bulletin 36: 149–527.
  • Sugar A, Finnamore A, Goulet H, Cumming J, Kerr JT, De Giusti M, Packer L (1999) A preliminary survey of symphytan and aculeate Hymenoptera from oak savannahs in southern Ontario. Proceedings of the Entomological Society of Ontario 129: 9–18.
  • Taeger A, Blank SM, Liston AD (2010) World catalog of Symphyta (Hymenoptera). Zootaxa 2580: 1–1064.
  • Timberlake PH (1943) Racial differentiation in Nearctic species of Dianthidium (Hymenoptera, Apoidea). Journal of the New York Entomological Society 51: 71–110.
  • Timms LL, Schwarzfeld M, Sääksjärvi IE (2016) Extending understanding of latitudinal patterns in parasitoid wasp diversity. Insect Conservation and Diversity 9: 74–86. https://doi.org/10.1111/icad.12144
  • Townes HK (1949a) The Nearctic species of Stephanidae (Hymenoptera). Proceedings of the United States National Museum 99: 361–370.
  • Townes HK (1949b) The Nearctic species of Evaniidae (Hymenoptera). Proceedings of the United States National Museum 99: 525–537.
  • Townes HK (1977a) A revision of the Rhopalosomatidae (Hymenoptera). Contributions of the American Entomological Institute 15(1): 1–34.
  • Townes HK (1977b) A revision of the Heloridae (Hymenoptera). Contributions of the American Entomological Institute 15(2): 1–12.
  • Townes HK (1983) Revisions of twenty genera of Gelini (Hymenoptera). Memoirs of the American Entomological Institute 35: 1–281.
  • Townes HK, Gupta VK, Townes MJ (1992) The ichneumon-flies of America north of Mexico: Part 11. Tribes Oedemopsini, Tryphonini and Idiogrammatini (Hymenoptera: Ichneumonidae: Tryphoninae). Memoirs of the American Entomological Institute 50: 1–296.
  • Townes HK, Townes MJ (1981) A revision of the Serphidae (Hymenoptera). Memoirs of the American Entomological Institute 32: 1–541.
  • Triapitsyn SV (2017) Revision of Alaptus (Hymenoptera: Mymaridae) in the Holarctic Region, with taxonomic notes on some extralimital species. Zootaxa 4279: 1–92. https://doi.org/10.11646/zootaxa.4279.1.1
  • Triapitsyn SV, Berezovskiy VV, Hoddle MS, Morse JG (2007) A review of the Nearctic species of Erythmelus (Hymenoptera: Mymaridae), with a key and new additions to the New World fauna. Zootaxa 1641: 1–64.
  • Triapitsyn S, Headrick DH (1995) A review of the Nearctic species of the thrips-attacking genus Ceranisus Walker (Hymenoptera: Eulophidae). Transactions of the American Entomological Society 121(4): 227–248.
  • Trjapitzin VA, Gordh G (1978a) Review of genera of Nearctic Encyrtidae (Hymenoptera: Chalcidoidea). I Entomologischeskoe Obozrenie 57(2): 364–385. [In Russian; English translation: Entomological Review 57: 257–270].
  • Trjapitzin VA, Gordh G (1978b) Review of genera of Nearctic Encyrtidae (Hymenoptera: Chalcidoidea). II Entomologischeskoe Obozrenie 57(3): 636–653. [In Russian; English translation: Entomological Review 57(3): 437–448].
  • van Achterberg C (2002) A revision of the Old World species of Megischus Brullé, Stephanus Jurine and Pseudomegischus gen. nov., with a key to the genera of the family Stephanidae (Hymenoptera: Stephanoidea). Zoologische Verhandelingen Leiden 339: 1–206.
  • various contributors (2018) Hymenoptera Online (HOL). https://hol.osu.edu [accessed 24.I.2018].
  • Veijalainen A, Wahlberg N, Broad GR, Erwin TL, Longino JT, Sääksjärvi IE (2012) Unprecedented ichneumonid parasitoid wasp diversity in tropical forests. Proceedings of the Royal Society B: Biological Sciences 279: 4694–4698. https://doi.org/10.1098/rspb.2012.1664
  • Wahis R (1986) Catalogue systématique et codage des Hyménoptères Pompilides de la région ouest-européenne. Notes fauniques de Gembloux 12: 1–91.
  • Wahis R (2006) Mise à jour du Catalogue systématique des Hyménoptères Pompilides de la région ouest-européenne. Additions et Corrections. Notes fauniques de Gembloux 59: 31–36.
  • Wahl DB, Sharkey MJ (1993) Ichneumonoidea. In: Goulet H, Huber JT (Eds) Hymenoptera of the world: an identification guide to families. Agriculture Canada, Publication 1894/E, Ottawa, Ontario, 358–509.
  • Wasbauer MS, Kimsey LS (1985) California spider wasps of the subfamily Pompilinae (Hymenoptera: Pompilidae). Bulletin of the California Insect Survey 26: 1–130.
  • Wharton RA, Marsh PM, Sharkey MJ (1997) Manual of the New World genera of the family Braconidae. Special publication of the International Society of Hymenopterists 1: 1–439.
  • Williams KA, Manley DG, Deyrup M, von Dohlen CD, Pitts JP (2012) Systematic review of the Dasymutilla monticola species-group (Hymenoptera: Mutillidae): using phylogenetics to address species-group placement and sex associations. Zootaxa 3554: 1–29.
  • Williams PH., Thorp RW, Richardson L, Colla SR (2014) A guide to the bumble bees of North America. Princeton University Press, Princeton, 208 pp.
  • Woolley JB (1988) Phylogeny and classification of the Signiphoridae (Hymenoptera: Chalcidoidea). Systematic Entomology 13: 465–501.
  • Yoshimoto CM (1978) Two new species of Epiclerus from the New World (Hymenoptera: Chalcidoidea, Tetracampidae). The Canadian Entomologist 110: 1207–1211. https://doi.org/10.4039/Ent1101207-11
  • Yu DSK, Achterberg van C, Horstmann K (2016) Taxapad 2016, Ichneumonoidea 2015. Database on flash-drive. www.taxapad.com, Nepean, Ontario, Canada.
  • Zhang YM, Gates MW, Shorthouse JD (2017) Revision of Canadian Eurytomidae (Hymenoptera, Chalcidoidea) associated with galls induced by cynipid wasps of the genus Diplolepis Geoffrey (Hymenoptera, Cynipidae) and description of a new species. Journal of Hymenoptera Research 61: 1–29. https://doi.org/10.3897/jhr.61.13466