Linley M. Sherin
McGill University, Department of Natural Resource Sciences, Macdonald Campus, 21,111 Lakeshore Road, Ste-Anne-de-Bellevue, Quebec, H9X 3V9, Canada. [email protected]
The Pachygastrinae are a large, globally distributed subfamily of soldier flies (Diptera, Stratiomyidae). Despite being widespread, they have poor representation in museum collections and are mostly known from larvae collected under the bark of fallen logs. All seven Canadian genera and nine of the eleven Canadian species are herein reviewed and keyed. An updated checklist to the Canadian species is presented. Twenty-one new provincial records are reported, and four species are recorded from Canada for the first time: Cosmariomyia pallidipennis (Williston), Gowdeyana punctifera (Malloch), Neopachygaster vitrea Hull and Pachygaster pulchra Loew. This increases the known fauna in Canada from seven to 11 species.
Cosmariomyia pallidipennis (Williston) (photo by Jeffrey Gruber)
The Stratiomyidae, or soldier flies, are a large, cosmopolitan family (~2850 spp.) of lower brachyceran flies that are diverse in both size and morphology (Hauser et al. 2017; James 1981). There are 114 species recorded from Canada. However, a recent review of Canadian Diptera predicted an additional 5-10 species remain to be recorded or described (Savage et al. 2019). The Canadian stratiomyid fauna includes species in seven of the twelve subfamilies: Antissinae, Stratiomyinae, Beridinae, Sarginae, Nemotelinae, Clitellariinae and Pachygastrinae (McFadden 1972, Woodley 2001). I here review the taxonomy of the 11 Canadian species of Pachygastrinae.
Pachygastrinae are a predominantly tropical subfamily with a global distribution. Possibly due in part to their small size, they have poor representation in collections when compared with larger stratiomyids. Pachygastrinae are morphologically distinguished from other Canadian Stratiomyidae in having five visible abdominal tergites (tergites 6 and 7 reduced), wing with m-cu crossvein connected to vein M3+4, and in having only three veins arising from the discal medial cell (Fig. 1; James 1981; Hauser et al. 2017). This subfamily was previously thought to be paraphyletic (Nagatomi and Iwata 1978; Woodley 2001), however the only molecular study of Stratiomyid phylogeny (Brammer and von Dohlen 2007) found evidence of a monophyletic Pachygastrinae. Monophyly was also supported in a 2010 morphological analysis (Brammer and von Dohlen 2010), though pachygastrine representation was limited.
Adults of the Nearctic Region are small (2.0-3.6 mm), predominantly black flies, often with silvery vestiture (Kraft and Cook 1961; Woodley 2001). Though sexual dimorphism is prevalent in soldier flies, Pachygastrinae often display only subtle variation between sexes (Curran 1927). Adults of many species form swarms, and the Nearctic species Gowdeyana punctifera Malloch has been recorded in swarms of up to 50 individuals (Woodley 2001). For unknown reasons, many species are collected from windows (Hull 1930) and few were collected prior to the common use of Malaise traps (Woodley 2001). Adults appear attracted to dead wood or fallen trees where females oviposit (Woodley 2001).
Relatively little is known about the habitat associations and feeding habits of larval Pachygastrinae (Woodley 2001). Larvae are universally brown to white, flattened dorso-ventrally and vary in length from 4.0-7.2 mm (Kraft and Cook 1961). Pachygastrine larvae are terrestrial and have been found under the bark and in tree holes of both deciduous and coniferous trees (Kraft and Cook 1961; McFadden 1967). They appear to feed on either the sap or microorganisms that occur on the moist wood of wounded and dying trees (McFadden 1967) and use cylindrical brush-like mouthparts to sweep food into their oral cavity (Cook 1953; Kraft and Cook 1961; McFadden 1967). It has been suggested previously that the larvae are predatory (Malloch 1917; Johannsen 1922; James 1981). However, McFadden (1967) and Woodley (2001) considered it doubtful given their common appearance in large numbers with McFadden (1967) recording over 100 larvae existing under the bark of a single log. The Canadian species Berkshiria albistylum Johnson has also been reared from a large number of larvae located under the bark of a fallen poplar tree (Marshall, S.A., pers. comm.).
Previous work addressing the Canadian pachygastrine fauna has been limited and largely attributable to a small number of dedicated dipterists. Kraft and Cook (1961) provided a revision of the Pachygastrinae of America north of Mexico, including keys to 21 species, 13 of which were newly described in that paper. Following Kraft and Cook, James (1965) provided a supplementary revision and key to the genera of the southwestern United States and parts of Mexico, adding two species to the Nearctic fauna (one new species, Zabrachia yuccae James, and one new Nearctic record, Cosmariomyia pallidipennis (Williston)). James (1981) provided illustrated adult and larval keys to the subfamilies and genera of Nearctic Stratiomyidae within the Manual of Nearctic Diptera, referencing 24 pachygastrine species in its generic treatment. Most recently, Woodley (2001, 2011) provided a catalogue of the global Stratiomyidae. This catalogue included 25 Nearctic pachygastrine species in nine genera, adding the species Artemita nana (Bellardi) to the Nearctic fauna. Of these 25 species, Woodley reported seven species from Canada (Berkshiria albistylum Johnson, Eidalimus fuscus (Kraft & Cook), Neopachygaster maculicornis (Hine), N. occidentalis Kraft & Cook, N. reniformis Hull, Zabrachia plicata (Kraft & Cook) and Z. polita (Coquillett)) and another five species were recorded from the adjacent USA.
Despite past attempts to generate a larval key to species of Nearctic Pachygastrinae, no complete key currently exists. Kraft and Cook (1961) provided a larval key to genera and larval keys to species for three genera (Eidalimus Kertesz, Gowdeyana Curran (as Eupachygaster Kertesz) and Zabrachia Coquillett). Most recently, McFadden (1967) provided a larval key to the genera of Stratiomyidae in America north of Mexico and incomplete larval keys to species.
Current pachygastrine adult keys are also inadequately illustrated and none present an up-to-date review of the Canadian fauna; the last Nearctic revision (Kraft and Cook 1961) included only five of the 11 Canadian species. I here key the seven Canadian genera including nine of the 11 Canadian species, illustrating adult diagnostic characters with photographs, adding new characters and highlighting taxa that need formal revision. Four of the species included here are new Canadian records (Table 1). By providing an updated and interactive key to the Canadian fauna, I hope to encourage the future study of the Pachygastrinae in Canada and bring attention to the need for a global revision of the subfamily. I also hope that this key serves as a tool for amateur naturalists to discover these beautiful and elusive flies.
Figure 1. Wing of Pachygaster pulchra Loew, edited to highlight characteristic wing venation of the subfamily Pachygastrinae in red; m-cu crossvein connected to M3+4, and only three veins arising from cell dm (M1, M2 apical and M4). Terminology follows Cumming and Wood (2017). Abbreviations: dm – discal medial cell; M – medial vein or media; M1+2 – fused first and second branch of medial vein; M1 – first branch of medial vein; M2 – second branch of medial vein; M3+4 – fused third and fourth branch of medial vein; M3 – third branch of medial vein; M4 – fourth branch of medial vein; m-cu – medial-cubital crossvein, m-m – medial crossvein.
Species included in this key are those previously known from Canada (Woodley 2001) as well as new Canadian records reported from specimen data for the first time (Table 1).
Specimen data were obtained from the University of Guelph Insect Collection (DEBU, Guelph, Ontario), the Biodiversity Institute of Ontario (BOLD, Guelph, Ontario), the Canadian National Collection of Insects, Arachnids and Nematodes (CNC, Ottawa, Ontario), the Lyman Entomological Museum (LEMQ, Montréal, Quebéc), the Ouellet-Robert Entomological Collection (QMOR, Montréal, Quebéc), the Spencer Entomological Collection (UBCZ, Vancouver, British Columbia) and the online platform iNaturalist [accessed 2020 October 17].
The following collections were also examined but held no Canadian Pachygastrinae: the Royal Ontario Museum (ROM, Toronto, Ontario), the Royal British Columbia Museum (RBCM, Victoria, British Columbia), the Cornell University Insect Collection (CUIC, Ithaca, New York, USA), as well as the online platform BugGuide (https://bugguide.net/) [accessed 2020 October 17].
Terminology used follows Cumming and Wood (2017). Data included within species pages were taken from specimen label data unless otherwise cited. Labels without GPS coordinates for locality data had coordinates generated retroactively using Google Maps GPS Coordinates (https://www.gps-coordinates.net). Distribution maps were created using the online software SimpleMappr (Shorthouse 2010). Previous Canadian distribution data were taken from literature (Woodley 2001) and mapped using whole provinces to indicate the presence of a species. This does not imply distribution throughout the entire highlighted regions. Canadian collection localities for the examined specimens were included in the distribution map as point data. For brevity, the term “provincial” is used when describing both provincial and territorial records throughout this manuscript and the provided key.
Specimen photographs were taken using a Canon DSLR mounted on a Stackshot rail and operated using Helicon Remote. Photographs were then stacked using Helicon Focus and edited using Adobe Photoshop. All specimen photographs were taken and edited by the author unless otherwise indicated. Photographs of living flies are from multiple sources; credit is given below each photograph.
Table 1. Checklist of Canadian Pachygastrinae arranged from west to east, by province and then territory: BC, British Columbia; AB, Alberta; SK, Saskatchewan; MB, Manitoba; ON, Ontario; QC, Quebec; NB, New Brunswick; NS, Nova Scotia; PE, Prince Edward Island; NL, Newfoundland and Labrador; YT, Yukon; NT, the Northwest Territories; NU, Nunavut. *New provincial record. **New national record.
|Berkshiria albistylum Johnson||7*||15||1*||23|
|Cosmariomyia pallidipennis (Williston)||1*||1**|
|Eidalimus fuscus (Kraft & Cook)||8||2*||10|
|Gowdeyana punctifera (Malloch)||1*||1*||6*||1*||9**|
|Neopachygaster maculicornis (Hine)||6*||6||15*||13*||40|
|Neopachygaster occidentalis Kraft & Cook||1||1|
|Neopachygaster reniformis Hull||5*||7*||5*||9*||25||38||89|
|Neopachygaster vitrea Hull||35*||18*||53**|
|Pachygaster pulchra Loew||8*||8**|
|Zabrachia spp. Coquillett||83||22*||16||75||5*||1*||202|
In total, 439 specimens were examined (52 DEBU, 20 BOLD, 262 CNC, 1 LEMQ, 101 QMOR, 2 UBCZ, 1 iNaturalist) that correspond to nine species and seven genera (Table 1). Within this material, three specimens were damaged and not identifiable to species. See Appendix 1 for a complete list of examined specimens. Notably, no pachygastrine species have been recorded from provinces east of Quebec. This is potentially due to sampling bias, as the most widespread species appear to exist wherever there are forested areas.
The genus Zabrachia needs revision, which is beyond the scope of this review. Therefore, the key identifies Zabrachia specimens only to generic level. Neopachygaster Austen is also in need of revision. All Neopachygaster species are included within the key although only a single character separates the three most widespread species (Neopachygaster reniformis Hull, N. vitrea Hull and N. maculicornis (Hine)). As Pachygastrinae are a predominantly tropical subfamily, it is recommended that revisions of any Canadian genera take place within larger revisions, ideally of the global fauna. It has been previously recommended that the entire subfamily is revised (Woodley 2001; Hauser, M., pers. comm) and the findings presented here support this recommendation for a global revision of the Pachygastrinae.
In total, I present 21 new provincial distribution records and four new national distribution records. This includes records from four provinces from which the subfamily had not previously been known. The following species are recorded from Canada for the first time; Cosmariomyia pallidipennis (Williston), Gowdeyana punctifera (Malloch), Neopachygaster vitrea Hull and Pachygaster pulchra Loew. This increases the known pachygastrine fauna in Canada from seven to 11 species, and the known stratiomyid fauna from 114 to 118 species (Savage et al. 2019).
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I would like to thank Steve Marshall for advising me, this review could not have been completed without his support and guidance, and without access to the Guelph Insect Collection (DEBU) where the lab work for this study was conducted. I am grateful to all the collections which loaned me material for study and/or allowed me to identify material at their institutions (listed in full under Methods). I thank Steve Paiero, Morgan Jackson and Jessica Gillung for editing the text and key multiple times and for their continued mentorship in all things dipterology. I also thank Martin Hauser for his edits and for sharing his expertise of Stratiomyidae, which helped improve the manuscript immensely. Finally, I would like to thank Brad Sinclair and Joel Gibson for their helpful and thorough reviews.