Checklist of Chalcidoidea and Mymarommatoidea (Hymenoptera) of Canada, Alaska and Greenland

A checklist of 1246 extant, described species, classified in 346 genera in 18 families of Chalcidoidea (Hymenoptera) are reported from Canada, Alaska (USA) and Greenland (Denmark) based on examined specimens and published records up to December 31, 2020. Of the reported species, 1214 (in 345 genera in 18 families) are listed from Canada, 113 (in 58 genera in 10 families) from Alaska, and 26 (in 22 genera in 4 families) from Greenland. The list includes 235 new species records and 53 new generic records for Canada (no new family records). Forty-one new species records, 22 new generic records and the families Chalcididae and Eurytomidae are newly reported for Alaska. No new records were found for Greenland. Two species (in one genus) of Mymarommatoidea are reported from Canada. For each species in Canada, distribution is tabulated by province or territory, except the province of Newfoundland and Labrador is divided into the island of Newfoundland and the region of Labrador. The inclusion of known species from Alaska and Greenland results in the first comprehensive distributional checklist for JHR 82: 69–138 (2021) doi: 10.3897/jhr.82.60058 https://jhr.pensoft.net Copyright Her Majesty the Queen in Right of Canada. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. CHECKLIST John T. Huber et al. / Journal of Hymenoptera Research 82: 69–138 (2021) 70 the entire northern part of the Nearctic region. A brief review of the history of cataloguing Chalcidoidea in North America and a comparison of this checklist with four published checklists from the Palaearctic


Introduction
The superfamily Chalcidoidea is one of the most diverse groups of organisms on the planet (Figs 2-13). More than 22,700 species are described (Huber 2017), but Heraty et al. (2013) estimated that there might be up to 500,000 species worldwide. Most chalcidoids, for which the biology is known, are parasitoids, having been reared from a wide variety (12 orders) of Insecta, and also 2 orders of Arachnida and the family Anguinidae (Nematoda) (Gibson 1993). A few are predators and some are phytophagous. For more comprehensive information on the biology of Chalcidoidea see, e.g., Clausen (1940), Askew (1971), Bendel-Janssen (1977), Gordh (1979a), Gauld and Bolton (1988), Hanson and Gauld (1995), Noyes (2019). In addition to Chalcidoidea, the small superfamily Mymarommatoidea is also included in this paper because it is generally considered to be the sister group to Chalcidoidea Huber et al. 2008;Heraty et al. 2013). The biology of Mymarommatoidea is unknown, except that one has been reared from a bracket fungus and most are collected in shady, moist areas such as deciduous forests (Huber et al. 2008).
The first published cataloguing efforts for Chalcidoidea of the Nearctic region began with Peck (1951), with supplements by Burks (1958Burks ( , 1967b. Peck (1963) catalogued the literature for each species up to and including 1958. The families comprising the Chalcidoidea section in Krombein et al. (1979) were catalogued by B. Burks, G. Gordh, and E. Grissell, former chalcidologists at United States National Museum of Natural History, Washington, DC (USNM), and included the relevant taxonomic literature to the end of 1972 or 1976 depending on the family. These catalogues also included species and records from Greenland. In his acclaimed Universal Chalcidoidea Database (UCD) for world Chalcidoidea, Noyes (2019) included the data from these previous catalogues. His database is now the only comprehensive compilation of taxonomic, biological, distributional and literature source information for world Chalcidoidea for the past 40-50 years, though it has not been updated since March 2019. Among other searches, it can be used to generate numbers and lists of Chalcidoidea names for any biogeographical region or country, and political subdivision within larger countries. Building on the information contained in the UCD, it is the purpose of this paper to provide the first checklist of the Chalcidoidea and Mymarommatoidea of Canada, Alaska and Greenland incorporating previously published, substantiated records as well as new records based on authoritatively identified specimens.

Sources of data
All records are substantiated by evidence, either collection-or literature-based. The vast majority of records in this checklist are based on specimens in the Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa (CNC). Additional distributional records, for which specimens could not be examined, were obtained by mining previous literature. Because of the relatively poor knowledge of Chalcidoidea, regional collections were generally not consulted because of the immense amount of work required to identify and curate most specimens in these collections. However, a few records were based on examination of specimens (or their photographs) deposited in other collections, as follows: Royal Alberta Museum, Edmonton, Alberta, Canada (PMAE: M. Buck); Royal Ontario Museum, Toronto, Ontario, Canada (ROM: C. Darling); Royal Saskatchewan Museum, Regina, Saskatchewan, Canada (RSKM: C. Sheffield), University of Alaska, Fairbanks, Alaska, USA (UAM: D. Sikes). Whereas most records are Canadian, records from Alaska (USA), Greenland (Denmark) and the 242 km 2 French Overseas Collectivity of Saint Pierre and Miquelon islands located 25 km from the southern coast of Newfoundland are also included so as to provide complete coverage of the northern part of the Nearctic region. Most of the Alaska species records (74 of 113) were based on specimens in collections (CNC and UAM). The remainder were literature records, primarily from the chapters in Krombein et al. (1979), which mostly catalogued specimens in the USNM. The Greenland records were taken almost exclusively from the relevant chapters in Böcher et al. (2015) with some records substantiated by specimens in the CNC. The single species recorded from Saint Pierre and Miquelon was obtained from the TAXREF database (Gargominy et al. 2020) managed by the Muséum national d'Histoire naturelle, Paris, but specimens were not examined and this species is only discussed in the text, not included in Table 2. Because of relatively poor sampling of the chalcidoids of these last three regions, it is likely that the current survey is not as complete for them as it is for Canada. All records published up to December 31, 2020 were evaluated for the current checklist.
We exclude from the checklist the very few fossil species of Chalcidoidea described from Canadian Cretaceous amber; all are now classified in Mymaridae (Poinar and Huber 2011) and Rotoitidae (Gumovsky et al. 2018). Further, species introduced into Canada from other countries for biological control against introduced pests are included only if there is irrefutable evidence that they became established after release. Often, no follow up surveys were undertaken to determine if the species released had established and so their continued presence is unsubstantiated. Because there is no single compilation of intentionally released species, one must search for their names in the five volumes on biological control in Canada: McLeod et al. (1962), Kelleher et al. (1971), Kelleher and Hume (1984), Mason and Huber (2002) and Mason and Gillespie (2012). At least 18 species are or were commercially produced in Canada (Mason and Huber 2002), 14 of which are included in the checklist. The other four species, Aphytis melinus DeBach, Eretmocerus californicus Howard, Metaphycus helvolus (Compere) and Trichogrammatoidea bactrae Nagaraja, occur in the southern states of USA or outside the Nearctic region on pests of crops not, or not extensively, grown commercially in Canada so are most unlikely to be found there. Some of the commercially produced species may establish more or less permanent populations in areas where they are released, often in large numbers at intervals (usually in greenhouses) or may occur naturally outside the facilities that produce them. We do not include any records in Table 2 that are solely known from websites such as iNaturalist, BugGuide or online databases of specimens in museums because for Chalcidoidea, examination of specimens with reference to authoritatively identified material is generally required. Finally, we only include described species, not undescribed taxa or specimens identified only to genus.  Table 1 is a summary of the numbers of described, recorded species of Chalcidoidea and Mymarommatoidea in Canada, Alaska and Greenland (not including Saint Pierre and Miquelon) totalled for each family for all 17 regions. Table 2 is the species checklist arranged alphabetically by family for the same 17 regions. It contains three types of distributional records: 1) a published record for which we have examined a specimen; 2) a new (unpublished) record for which we have examined a specimen; and 3) a published record for which we have not examined a specimen, but is well-substantiated (see Assessing credibility of records section in Bennett 2021a). The different types of records are indicated by different fonts and colours in Table 2 (see Table heading). The absence of a provincial or territorial acronym for a species recorded from Canada indicates that the taxon was recorded from Canada but no province was specified. Literature references (shown in the far right column of Table 2) are only noted for previously published records for which no specimens were examined. Authors' names that have been spelled in different ways, such as with or without diacritic marks, are spelled in only one way for consistency, for example, Förster, not Foerster. Literature references for published records for which specimens were examined are not provided as this would dramatically increase the size of the study and make it practically impossible to present the distributional data in a table format. We do provide an extensive, but by no means comprehensive, list of references for higher taxa, e.g., revisions of genera and regional checklists, which are cited directly under the higher taxon names in Table 2. Our list is not a catalogue so synonyms and homonyms are generally excluded; these can be found in UCD. In addition to the published checklist, the data presented in Table 2 have been added to Canadensys (https://data.canadensys.net/ipt/resource?r=aafchymenoptera-canada-ak-gl) and are also registered on GBIF (Bennett 2021b).

Classification
The family classification in Chalcidoidea has been extremely volatile, varying from 1 to 23 recognized families (see Grissell and Schauff 1997), with changes even in the last few years and more changes likely in the future. We mainly follow the family classification in Heraty et al. (2013) in which 22 families were recognized, except we also recognize the family Megastigmidae, which was raised from subfamily status within Torymidae by Janšta et al. (2018). Five extant families, the Agaonidae, Cynipencyrtidae, Eriaporidae, Rotoitidae, and Tanaostigmatidae do not occur in northern North America. Species of Agaonidae are associated exclusively with figs (Ficus spp.), which do not naturally occur in Canada, Cynipencyrtidae consists of one genus and species in Asia, Eriaporidae occur only in the Old World, the two described species of Rotoitidae occur only in Chile and New Zealand, and species of Tanaostigmatidae occur in the New World only as far north as the southern states of USA.

Results and discussion
A total of 1246 described, extant species of Chalcidoidea in 346 genera in 18 families are listed for Canada, Alaska and Greenland (Tables 1 and 2). Of these, 1214 species, classified in 345 genera in 18 families, are listed from Canada. To place the current number of species in perspective, it represents a 149% increase from the 500 species reported in Danks (1979). In terms of relative species richness within Hymenoptera, Chalcidoidea species comprise 13.5% of the 9250 species recorded in northern North America and 13.6% of the 8933 recorded in Canada (Bennett 2021a). Yet in comparison to other areas of the world the number of Chalcidoidea is relatively low. Gijswit (2003)  ) are only about 3.2% the size of Canada (9.985 million km 2 ), but despite this, the 1754 species recorded from there is almost 1.5 times greater than all the species we record from Canada. Canada and Russia have a much greater variety of ecozones and habitats than does Britain and Ireland, which have little or no tundra, temperate rain forest, grassland or semi-desert. While the colder climate over much of Canada contributes to the apparently depauperate fauna this is not the main reason. Lack of collecting as well as lack of study of what has been collected, in groups other than those of research interest to the few taxonomists who study Chalcidoidea in Canada, is probably the main factor contributing to poor knowledge of species and their distributions.

FAMILY TRICHOGRAMMATIDAE
Based on sequencing of the DNA barcode region of cytochrome oxidase I (COI) and using the Barcode Index Number (BIN) criterion of Ratnasingham and Hebert (2013) that 2% sequence divergence is indicative of species differences, Bennett et al. (2019) estimated that there are 3301 species of Chalcidoidea from Canada in the Barcode of Life Data System (BOLD) database (Ratnasingham and Hebert 2007). This represents 2.7 times the number of recorded named species of Chalcidoidea in Canada. These BINs have not yet been reconciled against the names in the checklist so the percent congruence is unknown, but it illus- trates that there are many unrecorded species. The comparison of COI sequences from unidentified specimens against those of named species in such databases as BOLD will certainly help to reveal yet more species to add to the checklist, e.g., Anastatus reduvii (Howard) (Eupelmidae), a recent, accidentally introduced parasi- Compared to Canada, the number of species of Chalcidoidea recorded from Alaska and Greenland is far lower (113 species in 58 genera in 10 families from Alaska and 26 described species in 22 genera in 4 families from Greenland) (Tables 1, 2). The summary of the entomofauna of Greenland (Böcher et al. 2015) included records from six chalcidoid families, but specimens of Aphelinidae and Trichogrammatidae were only identified to genus, and so are not included in our checklist. Relative to Canada, the smaller land masses and more northerly latitudes of Alaska and Greenland definitely contribute to lower species richness, but it is also probable that the numbers are lower than expected because of relatively poor sampling in Alaska and Greenland (as well as in the three Canadian territories, NT, NU and YT). In addition, one species of Chalcidoidea, Pteromalus elevatus (Walker, 1834) (Pteromalidae) has been recorded from the French Overseas Collectivity of Saint Pierre and Miquelon Islands (Gargominy et al. 2020). This record is derived from an online database and specimens have not been examined, but this species has been recorded previously from NB, NS and NL (Hoebeke and Wheeler 1996).
Two species of Mymarommatoidea are recorded, one known only from Ontario, and one from Ontario, Quebec and New Brunswick (see bottom of Table 2). The latter species (Mymaromella pala Huber and Gibson) has also been collected from Montana, USA (Hatten et al. 2011), so it is likely that future collecting in central and western Canada will reveal its presence there as well. This further illustrates the work still needed to obtain reliable records of the species and their distributions in Canada and the rest of North America.
In terms of species richness by distributional area, the political region with the highest recorded number of species of Chalcidoidea is Ontario (852, 68.4% of 1246 species), followed by Quebec (566, 45.4%) and British Columbia (440, 35.3%) ( Fig. 1 and Table 1). The greater relative species richness in these areas is certainly strongly influenced by higher sampling effort compared to more northern or central regions (Langor 2019). Despite this bias, higher species diversity is expected in these three provinces relative to most other regions because of the higher number of ecozones and habitats, compared to more northern areas (Scudder 1979).
In total, 235 new species records of Chalcidoidea are reported for Canada, which represents 19.4% of the total number of described species recorded. The number of new Canadian species records by family is shown in Table 1 (in parentheses following the Canada totals). The checklist includes 53 new generic records for Canada (those for which the only Canadian records are shown in boldface in Table 2). All families in our checklist were previously recorded from Canada, although some, like Azotidae (see Heraty et al. 2013) and Megastigmidae (see Janšta et al. 2018), were not recognized as families in previous catalogues (e.g., chapters in Krombein et al. 1979), whereas some other previously recognized families have been subsumed within others, e.g., Elasmidae is now classified as part of Eulophidae (Gauthier et al. 2000). For Alaska, there were 41 new species records (36.3% of the total), 22 new generic records and the families Chalcididae and Eurytomidae are also newly recorded. There were no new records for Greenland.
The distributions given by province and territory for Canada, the state of Alaska for USA, and Greenland must be accepted with caution. Records taken from the literature, particularly pre-1980 records, could be based on misidentifications because of the difficulty in identifying all taxa reliably to species when fewer workable species keys existed. Although Yoshimoto (1984) provided a key to the families and some subfamilies of Chalcidoidea from Canada, a comprehensive key to the genera of the Nearctic region did not exist prior to 1997 . The superfamily is so diverse and speciose, even in the relatively cold, mid-to-high latitude region encompassed by our checklist, that the relatively few authorities available, in the past or now, simply could not correctly identify every specimen encountered. In preparing our checklist, there was no time or sufficient expertise for exhaustive study of all specimens from various localities in Canada and comparison with previously identified CNC specimens, which may or may not have been authoritatively and correctly identified in the first place. As often occurs, cataloguing efforts greatly outpace production of taxonomic revisions, including comparative descriptions and comprehensive identification keys to genera or species. Yet, once published, past identifications are the basis of the names and distributions presented in our checklist and had to be included even if some are wrong. Nevertheless, this checklist is our best summary of the current state of knowledge. It provides baseline data for future studies on the taxonomy, natural history and distribution of chalcidoids and will be useful to more applied fields such as the biological control of insect pests.