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Integrative taxonomy of Nearctic and Palaearctic Aleocharinae: new species, synonymies, and records (Coleoptera, Staphylinidae)
expand article infoAdam J. Brunke, Mikko Pentinsaari§, Jan Klimaszewski|
‡ Canadian National Collection of Insects, Ottawa, Canada
§ University of Guelph, Guelph, Canada
| Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, Québec, Canada
Open Access

Abstract

A long tradition of separate Nearctic and Palaearctic taxonomic studies of the diverse aleocharine rove beetles (Coleoptera: Staphylinidae) has obscured the recognition of Holarctic species and detection of adventive species in both regions. Recently, integrated study of the two regions through detailed morphological comparisons and development of an authoritatively identified DNA barcode reference library has revealed the degree to which these two aleocharine faunas are interconnected, both naturally and through human activity. Here this approach is adopted to recognize new species, reveal Holarctic species, and recognize adventive species in both North America and Europe. The following new species are described: Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov. from British Columbia; Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov., from Ontario, Maryland and North Carolina; Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov., from Ontario and Virginia; and Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. from eastern Canada. Dasygnypeta velata and Philhygra angusticauda are revealed to be Holarctic species, resulting in the following synonymies: Dasygnypeta velata (Erichson, 1839) = Gnypeta minuta Klimaszewski & Webster, 2008, syn. nov. and Philhygra angusticauda (Bernhauer, 1909) = Atheta (Philhygra) pinegensis Muona, 1983, syn. nov. The Nearctic species Hylota ochracea (and genus Hylota), Thecturota tenuissima, and Trichiusa robustula are newly reported from the Palaearctic region as adventive, resulting in the following synonymies: Hylota ochracea Casey, 1906 = Stichoglossa (Dexiogyia) forticornis Strand, 1939, syn. nov.; Thecturota tenuissima Casey, 1893 = Atheta marchii Dodero, 1922, syn. nov.; and Trichiusa robustula Casey, 1893 = T. immigrata Lohse, 1984, syn. nov. The Palaearctic species Amarochara forticornis, Anomognathus cuspidatus, Oligota pumilio, and Parocyusa rubicunda are newly confirmed from the Nearctic region as adventive, resulting in the following synonymies: Parocyusa rubicunda (Erichson, 1837) = Chilopora americana Casey, 1906, syn. nov. and Anomognathus cuspidatus (Erichson, 1839) = Thectura americana Casey, 1893, syn. nov. The genus Dasygnypeta, sensu nov. is newly reported from North America, Paradilacra is newly reported from eastern North America, and Haploglossa is newly reported from Canada, resulting in the following synonymy: Paradilacra densissima (Bernhauer, 1909) = Gnypeta saccharina Klimaszewski & Webster, 2008, syn. nov. Native Cyphea wallisi is newly reported from across Canada and C. curtula is removed from the Nearctic fauna. The status of both Gyrophaena affinis and Homalota plana is uncertain but these species are no longer considered to be adventive in North America. Three new combinations are proposed: Dasygnypeta baranowskii (Klimaszewski, 2020) and D. nigrella (LeConte, 1863) (both from Gnypeta) and Mocyta scopula (Casey, 1893) (from Acrotona). Dolosota Casey, 1910, syn. nov. (type species Eurypronota scopula Casey), currently a subgenus of Acrotona, is therefore synonymized with Mocyta Mulsant & Rey, 1874. Additionally, four new Canadian records and 18 new provincial and state records are reported.

Keywords

Canada, DNA barcodes, faunistics, morphology, North America, rove beetles, United States

Introduction

Historically, taxonomic research on the hyperdiverse aleocharine rove beetle (Coleoptera: Staphylinidae) faunas of North America and better-known Europe has been conducted separately, with a few exceptions (e.g., Klimaszewski et al. 1979). More recently, a closer examination of Aleocharinae in these two regions has demonstrated that a number of species are shared between the Nearctic and Palaearctic, either naturally (Holarctic) or through human activity (adventive) (e.g., Muona 1984; Klimaszewski et al. 2007; Klimaszewski et al. in press). The interconnectedness of these assemblages, combined with the sheer diversity of the subfamily, have made it difficult to avoid describing synonyms of taxa from other regions, especially when those taxa have been described in entirely different genera (e.g., Gusarov 2003a). One strategy to broadly address this challenge is the publication of detailed illustrations of habitus and genitalia in comprehensive faunal treatments such as the recently available ‘Aleocharinae of Eastern Canada’ (Klimaszewski et al. 2018) and ‘the Danish Beetle Bank’ website (Hansen et al. 2017), the latter an online resource for the Danish beetle fauna. In the past few years, resources such as these have made it possible to efficiently cross-check Nearctic and Palaearctic aleocharines without consulting a comprehensive reference collection for each region.

In combination with careful morphological study, large-scale DNA barcoding (e.g., deWaard et al. 2019) has accelerated the discovery of Holarctic species, and the detection of new adventive species and potential synonyms in the Canadian beetle fauna (e.g., Pentinsaari et al. 2019) by algorithmically flagging potential taxonomic issues and novelties, and connecting authoritatively identified specimens to unidentifiable females, damaged specimens, or other life stages. This integrated taxonomic approach, as applied to Aleocharinae, has already resulted in the detection of adventive species of genera Amischa, Atheta, and Myllaena in North America (Pentinsaari et al. 2019), and has refined the classification of Holarctic species in Atheta (Klimaszewski et al. in press), Boreophilia (Klimaszewski et al. 2019), and Gnathusa (Klimaszewski et al. in press).

Here we broadly compare morphological and molecular data across the Nearctic and West Palaearctic Aleocharinae in order to better integrate the taxonomic knowledge of these two regions. We describe four new Nearctic species, propose revised generic concepts, report new distributional records, and propose a number of new synonyms that impact our understanding of Holarctic and adventive species.

Materials and methods

Almost all specimens used in this study were dissected and their genitalia were subsequently examined on microslides. The genital structures were dehydrated in absolute ethanol, mounted in Canada balsam on celluloid microslides, and pinned with the specimens from which they originated. The photographs of the entire body and the genital structures were taken using an image processing system (Nikon SMZ 1500 stereoscopic microscope; Nikon Digital Camera DXM 1200F) and processed in Adobe Photoshop. Terminology mainly follows that used by Lohse et al. (1990) and Klimaszewski et al. (2018). The ventral part of the median lobe of the aedeagus is considered to be the part of the bulbus containing the foramen mediale, the entrance of the ductus ejaculatorius, and the adjacent venter (ventral part of the tubus of the median lobe) of the tubus; the opposite side is referred to as the dorsal part.

Depository abbreviations:

CBG Centre for Biodiversity Genomics, University of Guelph, Guelph, Ontario, Canada;

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario, Canada;

cRW Personal collection of Reginald P. Webster, Charters Settlement, New Brunswick, Canada (also known as RWC);

LFC Laurentian Forestry Centre, Québec, Quebec, Canada;

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, United States (C. Maier);

NHMD Natural History Museum of Denmark, Copenhagen University, Copenhagen, Denmark (A. Solodovnikov);

NMNH National Museum of Natural History, Washington D.C., United States (F. Shockley);

UAM University of Alaska Museum Insect Collection, Fairbanks, Alaska, United States (D. Sikes);

ZFMK Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany;

ZMHB Museum für Naturkunde, Berlin, Germany (B. Jaeger);

ZMUO Zoology Museum, University of Oulu, Oulu, Finland (M. Mutanen);

ZSM Zoologische Staatssammlung, Munich, Germany (F. Koehler).

We have examined all DNA barcode data for Aleocharinae previously generated by a variety of projects in both Europe and North America (e.g., Rulik et al. 2017; Sikes et al. 2017; McClenaghan et al. 2019; and other studies summarized by Pentinsaari et al. 2019). Fifty-three barcode sequences, the majority of which are Canadian sequence records originating from various projects coordinated by CBG, are published here for the first time. All sequences were analyzed using the workbench tools of the BOLD platform (http://www.boldsystems.org) after applying filters to exclude those flagged as misidentifications, those with sequence lengths under 100 bp, those with stop codons, and those flagged as contaminated. Sequences were generally visualized as clusters in neighbour-joining trees (using the Taxon ID Tree tool). In addition, BIN Discordance Reports, which compare the taxonomy of the specimen records to their BIN assignments, were used to detect potential misidentifications and synonyms.

All COI barcode sequences in BOLD that fulfill quality criteria (minimum length 500 bp, less than 1% ambiguous bases) are automatically assigned into BIN clusters (Barcode Index Numbers; Ratnasingham and Hebert 2013). In addition, sequences between 300–500 bp can be assigned as members of an existing BIN, but they will not be accepted as founding members of a new BIN. BINs correspond to species at a high accuracy in northern and central European beetles (Hendrich et al. 2015; Pentinsaari et al. 2017), and we treat BINs here as provisional hypothetical species.

The DNA barcode sequences studied here, including both previously unpublished data and the sequences published in earlier studies, have been compiled into a publicly available dataset on BOLD (DS-ALEO2020, https://doi.org/10.5883/DS-ALEO2020) along with collecting data, images of the specimens (if available), and other metadata related to the specimens and sequences. The sequences are also available through GenBank (accessions provided in Suppl. material 1: Table S1).

Taxonomic accounts

Tribe Aleocharini Fleming, 1821

Amarochara forticornis (Lacordaire, 1835)

Fig. 1A–H

Material

(DNA barcoded specimens). Canada: Ontario: Fergus, Centre Wellington District High School, 43.704, -80.358, Malaise trap, 3.V.2013, M. Cottrill (1, CBG); Guelph, Biodiversity Institute of Ontario, 43.528, -80.229, Malaise trap, 25.VII.2013, BIO Collections Staff (1, CBG); Rouge National Urban Park, west of Glen Rouge campground, 43.804, -79.146, marsh scrub along riverside, pitfall trap, 9.VI.2013, BIObus 2013 (1, CBG); Cambridge, rare Charitable Research Reserve, Preston Flats, 43.3908, -80.3747, grassy wetland, pitfall trap, 31.V.2015, BIO Collections staff (2, CBG); Peterborough, 44.318, -78.372, farm, malaise trap, B. McClenaghan (1, CBG).

Distribution

Origin: Palaearctic (adventive in Nearctic). Canada: ON [new record].

Diagnosis

Amarochara forticornis may be easily recognized among the other Canadian species of the genus by the distal antennomeres, which are less than twice as wide as long. The species is also unique within the genus by having a distinct basal impression on abdominal tergite VI.

Bionomics

In its native range, A. forticornis occurs in a variety of open and forested habitats, including forests, edges of waterways, grasslands, agricultural fields, and gardens (Assing 2002). It has been mostly collected by pitfall traps in the spring and summer, and then from flood debris in the cooler months of the year (Assing 2002). Assing (2002) suggested that beetles in flood debris were washed from some cryptic, subterranean microhabitat. Canadian specimens were collected in similar ways as in Europe.

Comments

Newly reported as adventive in North America, from several localities in southern and eastern Ontario. It is native to the West Palaearctic and is known from most of Central Europe, Russian Central Territory, Armenia, and Georgia (Newton 2019).

Figure 1. 

Amarochara forticornis (Lacordaire) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H).

The key to Eastern Canadian Amarochara in Klimaszewski et al. (2018) should be modified as follows

1A Antennomere 10 only weakly transverse (Fig. 1A); abdominal tergite VI with distinct basal impression in addition to coarse punctures Amarochara forticornis (Lacordaire)
Antennomere 10 strongly transverse, at least twice as wide as long (native species); abdominal tergite VI with, at most, coarse punctures at base 1B
1B Pronotum with strong microsculpture and coarse, dense punctation, surface almost matte A. duryi (Casey)
Pronotum without microsculpture or with fine microsculpture, and with fine sparse to moderately dense punctation, surface moderately to highly glossy 2

Tribe Oxypodini C.G. Thomson, 1859

Subtribe Microglottina Fenyes, 1918

Haploglossa nebulosa (Casey, 1906)

Fig. 2A–H

Material

(DNA barcoded specimens). Canada: Ontario: Rouge National Urban Park, Toronto Zoo, 43.8223, -79.1897, forest, malaise trap, 21.V.2013, L. Attard and K. Greenham (1, CBG).

Distribution

Origin: Nearctic. Canada: ON [new record]. United States: OK, PA.

Diagnosis

Haploglossa nebulosa may be easily distinguished from the other Nearctic species of the genus, H. barberi (Fenyes), by the bicolored elytra and fusiform body (Klimaszewski and Ashe 1991). Based on the shape of the spermatheca with its narrow capsule and broadly rounded apex, H. nebulosa may be most closely related to Palaearctic H. marginalis (Gravenhorst) as is suggested by barcode clustering. However, H. nebulosa can be readily distinguished by the pronotum, which is dark and paler only along the margins, while H. marginalis has broad pale areas laterally. Haploglossa nebulosa was compared to Palaearctic H. villosula (Stephens) by Klimaszewski and Ashe (1991; as H. pulla (Gyllenhal)), but the species is quite different externally (much darker, finer pronotal punctation) and the spermatheca of the latter species is of the type with a large, rounded capsule.

Bionomics

All members of Haploglossa are nidicolous, mostly in bird nests but also in mammal and ant nests (summarized by Staniec et al. 2010). Some species with well-known life histories appear to specialize on particular types of host nests, such as birds of prey (H. picipennis (Gyllenhal)) or bank swallows (H. nidicola (Fairmaire)) (Staniec et al. 2010). The genus is very rarely collected in North America. The Nearctic species H. barberi (Fenyes) was collected in long series from bank swallow nests (Klimaszewski and Ashe 1991). One specimen of H. nebulosa has been found in a rodent nest within a hollow tree (Klimaszewski and Ashe 1991) but bird and mammal nests have been poorly sampled in the Nearctic and more collecting is needed to determine the biology of the Nearctic Haploglossa (Brunke and Buffam 2018).

Comments

The genus Haploglossa and H. nebulosa are newly reported from Canada, from a single locality in southern Ontario. The species is also known from Oklahoma and Pennsylvania, United States (Klimaszewski and Ashe 1991).

Figure 2. 

Haploglossa nebulosa (Casey) A habitus B median lobe of aedeagus in lateral view (adapted from Klimaszewski and Ashe (1991)) C median lobe of aedeagus in dorsal view (adapted from Klimaszewski and Ashe (1991)) D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H).

The key to genera of Oxypodini in Eastern Canada in Klimaszewski et al. (2018) should be modified as follows

8A Pronotum strongly converging anteriad; posterolateral margin of elytra with strong sinuate emargination 8B
Pronotum not or, at most, weakly converging anteriad 9
8B Pronotum with fine punctures, not clearly visible at moderate magnification, shape strongly transverse, ~ 1.5 × wider than long Crataraea Thomson
Pronotum with coarse punctures, clearly visible with low magnification, shape weakly transverse, no more than 1.4 × wider than long Haploglossa Kraatz

Subtribe Oxypodina C.G. Thomson, 1859

Hylota cryptica Klimaszewski & Webster, 2016

Fig. 3A–G

Material

(DNA barcoded specimens). Canada: Ontario: Guelph, Hanlon Preservation Park, 43.506, -80.213, mixed forest, dead wood and beating, 11.VI.2017, M. Pentinsaari (1, CBG); Hartington, Eel Lake Cottage, 44.5628, -76.553, Lindgren funnel, 12.VII.2017, G. Blagoev (1, CBG); Kawartha Lakes, 44.28, -78.529, farm, malaise trap, 5.V.2016, B. McClenaghan (1, CBG); Murphy’s Point Provincial Park, 44.7812, -76.2336, forest, malaise trap, 19.VI.2014, CBG Collections staff (1, CBG); Newfoundland: Terra Nova National Park, Blue Hill Road, 48.598, -53.9702, malaise trap, old balsam fir forest, 2.VII.2013, E. Perry (1, CBG).

Distribution

Origin: Nearctic. Canada: AB, NB, NF [new record], ON [new record].

Bionomics

Little is known about the microhabitat preferences of this species, but it likely occurs in in nests or cavities within trees as does H. ochracea (Klimaszewski et al. 2018).

Comments

This recently described species, previously known from New Brunswick and Alberta (Klimaszewski et al. 2018) is newly recorded from Ontario and Newfoundland. It is likely to be widely distributed in North America east of the Rocky Mountains.

Figure 3. 

Hylota cryptica Klimaszewski & Webster A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Webster et al. (2016).

Hylota ochracea Casey, 1906

Fig. 4A–H

Stichoglossa (Dexiogyia) forticornis Strand, 1939, syn. nov.

Material

(DNA barcoded specimens). Canada: Ontario: Guelph, Dovercliffe Road, 43.51, -80.254, backyard, compost and mouldy hay piles, 6.VI.2018, M. Pentinsaari (3, CBG); Guelph, Hanlon Preservation Park, 43.51, -80.221, mixed forest, at UV light, 30.VI.2018, M. Pentinsaari (1, CBG); Whitby, Julie Payette Public School, Malaise trap, 43.886, -78.934, 22.IV.-03.V.2013, Z. Turner (1, CBG). Quebec: Montreal, Montreal Botanical Garden, 45.559, -73.566, Malaise trap, 24.VII-02.VIII.2014, M. Larrivee (1, CBG). Finland: Al: Lemland, Äspholm, 60.0675, 19.9583, 9.X.2011, M. Pentinsaari (1, ZMUO); Al: Lemland, Nåtö, 60.046, 19.981, 26.VI.2014, M. Pentinsaari (2, ZMUO); N: Sipoo, Sipoonkorpi, 60.304, 25.202, window trap, 2.VIII.2013, S. Karjalainen and P. Martikainen (1, ZMUO).

Distribution

Origin: Nearctic (adventive in Europe). Canada: NB, NS, NT, ON, QC, SK. United States: NY, VT.

Bionomics

Hylota ochracea is strongly associated with bird nests in forested habitats. It has also been collected from artificial analogs such as a pigeon coup, manmade nest boxes, and a plastic composter bin containing carrion and decaying vegetables (Klimaszewski et al. 2018). The specimens recently collected in Ontario, Canada were found in compost and at UV light.

Comments

Hylota ochracea, a widespread Nearctic species (Klimaszewski et al. 2018), is newly reported from the Palaearctic region and had been previously known from Finland, Denmark, Germany, Norway, Sweden, and Switzerland (Lundberg 2006; Schülke and Smetana 2015; Newton 2019) under the synonym Dexiogyia forticornis. Hylota is also a new genus record for the Palaearctic region.Nearctic Hylota ochracea and Palaearctic D. forticornis share a BIN and do not form separate clusters. One of the DNA barcode haplotypes is shared between Finnish and Canadian specimens. Nearctic and Palaearctic populations also have identical male and female genitalia. Based on its specialization on microhabitats in forests, we do not consider H. ochracea to be a naturally occurring Holarctic species. Holarctic beetles are generally those that occur north of the treeline and have crossed treeless Beringia in the last 2.8 Mya (reviewed in Brunke et al. 2020). Hylota ochracea may have been introduced to the Palaearctic region with the nest material of poultry or domestic pigeons, or with another form of decaying plant matter. A similar situation has occurred with the bird nest-associated staphylinid Bisnius palmi (Smetana), which was originally described from Italy but later found to be a native Nearctic species (Smetana 1995).

With the above synonymy, the genus Dexiogyia is now known only from externally similar sister species D. angustiventris (Casey) (Nearctic) and D. corticina (Erichson) (West Palaearctic), plus Afrotropical D. congoensis (Scheerpeltz). As in the former D. forticornis, D. congoensis is probably misplaced due to superficial similarity. Hylota is readily separated from Dexiogyia by the shape of the pronotum, which is strongly convergent anteriad, such that its apical width is subequal to the width of the head. In Dexiogyia, the head is distinctly narrower than the pronotum.

Figure 4. 

Hylota ochracea Casey A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), reproduced with permission.

Isoglossa Casey, 1893

Rheobioma Casey, 1906; Klimaszewski and Pelletier (2004), syn. of Neoisoglossa

Athetalia Casey, 1910 (in part); Klimaszewski and Pelletier (2004) syn. of Neoisoglossa

Neoisoglossa Klimaszewski & Pelletier, 2004; Gouix and Klimaszewski (2007), syn. of Isoglossa, unnecessary replacement name; Klimaszewski et al. (2020) as valid genus, incorrectly attributed to Casey (1893).

Comments

In Klimaszewski et al. (2020), Neoisoglossa was incorrectly attributed to Casey but was actually proposed by Klimaszewski and Pelletier (2004), apparently as an unnecessary replacement name for Isoglossa Casey 1893. The previous treatment of these generic names and two other synonyms in the catalog of Gouix and Klimaszewski (2007) is correct and followed here. Blackwelder (1952) was wrong and there is no Isoglossa Newman that preoccupied Casey’s name, so Isoglossa Casey stands as valid with Neoisoglossa as a synonym.

Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov.

Fig. 5A–H

Type material

Holotype. (male): Canada, British Columbia, Prince George, Nukko Lake Elementary EQP-CLL-574, 54.0831°N, 122.988°W, 764 m asl, Holly Sapun 04/20/2015 to 05/08/2015, Barcode of life, DNA voucher specimen, Sample ID: BIOUG22036-B02, Process ID: SMTPM2682-15 (CNC). Paratypes (3, CBG): Canada, British Columbia, Prince George, Nukko Lake Elementary EQP-CLL-574, 54.0831°N, 122.988°W, 764 m asl, Holly Sapun 04/20/2015 to 05/08/2015, Barcode of life, DNA voucher specimen, Sample ID: BIOUG22036-B07, Process ID: SMTPM2682-15 (1 male, CBG); same label data except: Sample ID: BIOUG22035-H08, Process ID: SMTPM2665-15 (1 female, CBG); Sample ID: BIOUG22036-A04, Process ID: SMTPM2672-15 (1 female, CBG).

Etymology

The species epithet refers to the remarkably separated triangular apex of the median lobe of the aedeagus, distinguishing it from all other members of the Ocalea group.

Distribution

Origin: Nearctic. Canada: BC.

Diagnosis

Isoglossa triangularis can be easily distinguished from all Nearctic species of the Ocalea group of genera by a combination of the strongly transverse and sparsely punctate pronotum, transverse antennomere 4, distinct triangular apex of the median lobe in lateral view (Fig. 5B), and distinct and simple ‘walking cane’ shape of the spermatheca (Fig. 5D).

Figure 5. 

Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H).

Description

Body length 3.0–3.3 mm, dark brown with elytra, antennomeres 1–2 or 1–3, legs and apical part of abdomen yellow-brown, forebody moderately glossy and abdomen strongly so (Fig. 5A); antenna moderately stout, antennomere 4 slightly transverse, antennomeres 5–10 strongly transverse, terminal antennomere ca. as long as two preceding ones combined; pronotum transverse (width/length ratio = 1.6), impressed medially at base, lateral edges evenly arcuate, length ratio of base to apex 1.2 ×, punctures fine and sparse, distance between punctures ~ 3 × diameter of a puncture, space between punctures with faint isodiametric microsculpture, pubescence directed laterad from midline of disc forming arcuate lines on both sides; elytra transverse (width/length ratio = 1.3), 1.5 × as long as pronotum; abdomen arcuate laterally and gradually narrowing toward apex. MALE. Tergite VIII broadly arcuate apically (Fig. 5E); sternite VIII with apical part broadly triangularly produced (Fig. 5F); median lobe of aedeagus in lateral view with narrowly elongate crista apicalis at base of bulbus, tubus moderately long, strongly produced ventrally, apex narrowly triangular constricted baso-dorsally in lateral view (Fig. 5B), internal sac structures not pronounced (Fig. 5B, C). FEMALE. Tergite VIII truncate apico-medially (Fig. 5G); sternite VIII arcuate apically (Fig. 5H); spermatheca with capsule approximately spherical with short neck, stem narrow, long and straight (Fig. 5D).

Bionomics

The specimens were collected in a Malaise trap on an open field surrounded by mixed forest.

Comments

Based on a combination of small size (< 4.5 mm), superficial, meshed microsculpture, sparse pronotal punctation, with punctures separated by more than two puncture diameters, pronotum transverse, shorter and narrower than elytra, and the transverse antennomeres 5–10, I. triangularis keys to genus Isoglossa Casey in Klimaszewski and Pelletier (2004). However, barcode sequences of this species do not cluster with Isoglossa agnita but rather form a cluster with Gennadota canadensis and the species of Neothetalia which bear a spermatheca with broad, circular loops, similar to those of Gennadota. Isoglossa triangularis has a simple spermatheca with a long straight stem and is not externally similar to these taxa (see above), and the barcode divergence between these species and I. triangularis is 11–12%. It is likely that I. triangularis belongs in a separate genus, but this is outside of the scope of this study. We here place I. triangularis tentatively in Isoglossa as not to disturb the existing morphological diagnoses of the genera and identification keys (e.g., Klimaszewski and Pelletier 2004; Klimaszewski et al. 2020), pending generic revision of the Ocalea group.

Parocyusa rubicunda (Erichson, 1837)

Fig. 6A–D

Tachyusa rubicunda Erichson, 1837

Chilopora americana Casey, 1906, syn. nov.

Tetralaucopora americana: Klimaszewski et al. 2018 (as valid species)

Parocyusa americana: Assing 2021 (possible synonym of P. rubicunda)

Material

(DNA barcoded specimens). Austria: Innervillgraten, Arntal, 46.8362, 12.3348, 1580 m, 22.VIII.2010, F. Koehler and J. Koehler (2, ZSM). Canada: Ontario: Northumberland County, Peter’s Woods Protected Natural Area, 44.124, -78.039, under rock in streambed,12.IX.2011, A. Brunke and S. Paiero (1, DEBU); Crieff Bog, 3 km W Puslinch, sedge meadow, 26.VI.1987, D. Blades (1, DEBU). United States: Connecticut: East Hartford, Two Rivers Magnet Middle School, 41.757, -72.655, 4.VI.2005, J. DeWaard (1, CBG).

Additional non-barcoded material

Canada: Ontario: Ancaster, 21.X.1967 (1, CNC); Rondeau Prov. Pk., Tulip Tree Trail, Carex and moss on logs in pond, 5.VI.1985, A. Davies and J.M. Campbell (1, CNC); Quebec: Montreal, 20.IX.1969, E.J. Kiteley, 1 (CNC); Mt. Orford Park, 20.IX.-11.X.1972, Dondale and Redner, 1 (CNC).

Distribution

Origin: West Palaearctic (adventive in North America). Canada: BC, ON, QC, NB, NF. United States: CT, NY, PA.

Bionomics

In North America, most specimens of this species have been collected from near water, including a sandy creek bank, in a dried streambed and in moss near the splash zone of a waterfall (Klimaszewski et al. 2018). Nearctic populations of this species are only known from female specimens and the species may be parthenogenetic in North America. In its native distribution, the northern and northwestern populations are also parthenogenetic (Assing 2021) and most likely represent the source population for the Nearctic introduction.

Comments

Parocyusa rubicunda is a widespread West Palaearctic species (Europe, European Russia, Turkey, Georgia, Iran, Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan) (Assing 2021). It is confirmed as established in the Nearctic region and had been previously known from North America under the synonym Tetralaucopora americana (Casey) (Klimaszewski et al. 2018). Assing (2021) recently reported this species from BC and treated T. americana as a tentative synonym based on the results presented in this paper.

Although all available sequences of this species are partial (382–407 bp) and a BIN has not been established as that would require at least one founding member with a minimum sequence length of 500 bp, Nearctic and Palaearctic sequences form a distinct cluster with only a single variable nucleotide site. External morphology and that of the spermatheca are identical. As spermathecae are of generally poor diagnostic value (especially the distal part) in Parocyusa (Assing 2021), the barcode evidence was quite critical for the resolution of this issue. Based on this evidence and a distribution centered around populated areas in northeastern and western North America, we here consider this species to be adventive in the Nearctic region. At the moment, it is not yet possible to determine whether the population in BC is a separate introduction from the northeastern population.

Recently, Assing (2021) revalidated Parocyusa as a genus separate from Tectusa after the discovery that Tectusa was not a monophyletic group. The type species of Parocyusa was found to be congeneric with that of Tetralaucopora, and the latter became a junior synonym of the former.

Figure 6. 

Parocyusa rubicunda (Erichson) A habitus B spermatheca C female tergite VIII D female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–D). Illustrations after Klimaszewski et al. (2016c).

Tribe Tachyusini C.G. Thomson

Revised key to the Canadian genera of Tachyusini

Adapted from Klimaszewski et al. 2018.

1 Elytra at humerus only slightly broader than pronotum at base (Figs 7, 8); impressions of abdominal tergites shallow, with punctation similar to that of disc (Figs 7, 8) 2
Elytra at humerus distinctly broader than pronotum at base (Figs 913); impressions of abdominal tergites with at least a few coarse punctures and glossy areas, punctation distinctly different from that of disc (Figs 913) 3
2 Pronotum with pubescence directed straight posteriad; hind tarsus subequal in length to hind tibia or longer (Fig. 8) Brachyusa Mulsant & Rey
Pronotum with pubescence directly posteriolaterad from midline; hind tarsus shorter, slightly longer than half the length of hind tibia or shorter (Fig. 7) Paradilacra Bernhauer
3 Abdomen clavate, at base distinctly narrower than head (Fig. 9); tergite III ca. as long as wide or longer; tergal impressions with median carina (Fig. 9) Tachyusa Erichson
Abdomen at most slightly constricted at base, subequal to or wider than head (Figs 1013); tergite III ca. twice as wide as long or wider; tergal impressions never with median carina (Figs 1013) 4
4 Abdomen at base elongate and moderately constricted, ca. as wide as head (Figs 1012); tergite III, at most, twice as wide as long; tergal impressions deep and sharply delineated from strongly convex disc (Figs 1012) Dasygnypeta Lohse, sensu nov.
Abdomen at base at most slightly constricted, wider than head (Fig. 13); tergite III strongly transverse, ~ 2.5 × wider than long or wider; tergal impressions shallower, gradually sloping to disc at base (Fig. 13) Gnypeta Thomson

Paradilacra densissima (Bernhauer, 1909)

Fig. 7A–H

Atheta (Paradilacra) densissima Bernhauer, 1909

Gnypeta saccharina Klimaszewski & Webster, 2008, syn. nov.

Material

(DNA barcoded specimens). Canada: Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane forest, 27.VI.2012, BIOBus 2012 (1, CBG); British Columbia: 10 km W Kamloops, New Afton Mine, Wetland Protected Area, 50.662, -120.504, 702 m, malaise trap, 22.VIII.2013, C. Simon (1, CBG); New Brunswick: York Co., Fredericton at Saint John River, 45.959, -66.625, margin of river in flood debris, 7.VII.2005, R.P. Webster [note: paratype of G. saccharina] (1, LFC); Ontario: Guelph, University of Guelph Arboretum, 43.53, -80.21, 12.VI.2019, M. Pentinsaari (1, CBG).

Distribution

Origin: Nearctic. Canada: AB, BC, NB, ON [new record], SK [new record]. United States: CA, MT, NV, ND, OR, UT.

Bionomics

This species has been collected from various wetland microhabitats including the edges of lakes, rivers, and a beaver pond (Gusarov 2003a).

Figure 7. 

Paradilacra densissima (Bernhauer) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission.

Comments

Paradilacra densissima and the genus Paradilacra, widespread in western and central North America (Gusarov 2003a), are newly reported from SK (records in Klimaszewski et al. 2016a, as G. saccharina) and eastern North America based on records from NB (Klimaszewski et al. 2008) and ON (this study), including one sequenced paratype of synonym Gnypeta saccharina. Under the present concept, only one widespread species of this genus is known.

Figure 8. 

Brachyusa helenae (Casey) A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2016a).

Dasygnypeta Lohse, 1974, sensu nov

In his key to the genera of Tachyusini, Paśnik (2010) distinguished Dasygnypeta, with its single Palaearctic species Dasygnypeta velata (Erichson), from the Nearctic genera by the following features: the narrow and slender abdomen with base approximately as wide as head, the ‘very deep’ tergal impressions, abdominal pubescence of tergites III–V directed posteriad, basal segment of metatarsus shorter than following two segments. Through an analysis of barcode data, we have discovered that the recently described Gnypeta minuta Klimaszewski & Webster is a synonym of D. velata (see below). A re-examination of other Nearctic Gnypeta species revealed two others that are closely related to D. velata: G. baranowskii Klimaszewski, and G. nigrella (LeConte). Their morphological divergence from other Gnypeta was represented by an earlier placement in the ‘Nigrella species group’ of Gnypeta by Klimaszewski et al. (2008), together with G. saccharina (now a synonym of Paradilacra densissima). In corroboration with morphology, DNA barcodes of D. velata and G. nigrella form sister clusters (sequences of G. baranowskii not available). Transfer of these two Gnypeta species to Dasygnypeta required a new concept for this genus as most of the distinguishing features were apomorphies of D. velata or found not to be of diagnostic value due to variability or overlap with other genera. Here we distinguish members of Dasygnypeta by their characteristic abdomen (Figs 1012): base of abdomen ca. as wide as head; basal half of abdomen elongate, tergite III (first visible) only moderately transverse, ~ 2 × as wide as long (at least 2.5 × in Gnypeta); tergites III–V with very deep basal impressions, each creating strongly convex areas on the disc. Members of Dasygnypeta could be confused with Tachyusa, which also bears an elongate basal abdomen, but in the latter genus tergites III–V are far more elongate and the abdominal base is narrower than the head (Fig. 9). Dasygnypeta nigrella was even originally described by LeConte (1863) in Tachyusa, likely based on this similarity.

Figure 9. 

Tachyusa obsoleta Casey A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view (adapted from Paśnik 2006) D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission.

Dasygnypeta baranowskii (Klimaszewski, 2020), comb. nov.

Fig. 10A–H

Gnypeta baranowskii Klimaszewski, 2020

Distribution

Origin: Nearctic. Canada: BC.

Bionomics

The type series was collected by sifting litter (Klimaszewski et al. 2020).

Comments

We here transfer this species to Dasygnypeta sensu nov. based on morphology illustrated by Klimaszewski et al. (2020). This recently described western species is most similar to eastern D. nigrella based on the moderately elongate antennae, more robust body and the distinctive deep emargination of female sternite VIII. However, it is easily distinguished by the coarser pronotal punctation, and male and female genitalia (Fig. 10). The aedeagus of D. baranowskii is superficially similar to P. densissima but these taxa are externally quite different.

Figure 10. 

Dasygnypeta baranowskii (Klimaszewski) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2020), used with permission.

Dasygnypeta nigrella (LeConte, 1863), comb. nov.

Fig. 11A–H

Tachyusa nigrella LeConte, 1863

Gnypeta nigrella: Klimaszewski et al. 2008

Material

(DNA barcoded specimens). Canada: New Brunswick: York Co., Fredericton at St. John River, 45.959, -66.625, margin of river in drift (mostly maple seeds), 4.VII.2004, R.P. Webster (1, LFC).

Additional non-barcoded material

Canada: Manitoba: 5 miles SW of Shilo, 5.VI.1958, J.F. McAlpine (1, CNC); Quebec: Montreal, 30.VIII.1968, E.J. Kiteley (1, CNC); Montreal 14.VI.1972, E.J. Kiteley (1, CNC); Berthierville, 5.VI.1976, E.J. Kiteley (3, CNC); Kazabazua, 15.VIII.1968, R.C. Lawrence (3, CNC); Wakefield, 4.VI.1930, W.J. Brown (1, CNC); Drummondville, 18.VII.1977, river mudflat, L. LeSage (5, CNC).

Distribution

Origin: Nearctic. Canada: MB [new record], NB, NF, ON, QC [new record]. United States: IL, MA, MD, NJ, NY, PA, VT, WV.

Bionomics

Collected along the edge of a variety of running and standing water-based habitats.

Comments

We here transfer this species to Dasygnypeta sensu nov. based on morphology and close clustering of DNA barcode sequences with D. velata. Dasygnypeta nigrella is a widespread species in eastern North America and is here newly reported from Manitoba and Quebec.

Figure 11. 

Dasygnypeta nigrella (LeConte) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission.

Dasygnypeta velata (Erichson, 1837)

Fig. 12A–H

Homalota velata Erichson, 1837

Gnypeta minuta Klimaszewski & Webster, 2008, syn. nov.

Material

(DNA barcoded specimens). Germany: Thuringia, Ufergehoelze am Speicher Loessau, 50.5665, 11.894, 460 m, 1.I.2013, GBOL-Team ZFMK (1, ZFMK); Thuringia, NE, Freibad, Werraufer, 50.9768, 10.0963, 20.X.2014, GBOL-Team ZFMK (1, ZFMK). United States: Alaska: Selawik NWR, Kugarak River, 66.561, -158.996, mud bank, shore washing, 23.VI.2010, D.S. Sikes (3, UAM).

Additional non-barcoded material

Canada: Manitoba: 5 miles SW of Shilo, 5.VI.1958, J.F. McAlpine (2, CNC); Northwest Territories: Inuvik, 24.VI.1972, A. Smetana (1, CNC). United States: Alaska: Hess Creek, mi. 24 Wales Hwy, 1.VII.1978, J.M. Campbell and A. Smetana (1, CNC); Kenai Peninsula, Anchor River at Hwy 1, 450’, 4.VI.1978, A. Smetana and E. Becker (1, CNC); mi. 1259 Alaska Hwy, 7.VII.1968, J.M. Campbell and A. Smetana (1, CNC).

Distribution

Origin: Holarctic. Canada: MB [new record], NB, NF, NT, SK. United States: AK.

Bionomics

Nearctic specimens have been collected most frequently along the margins of running water but also along the margins of a forest pool (Klimaszewski et al. 2018).

Comments

Dasygnypeta velata is newly reported from North America and was previously known in the Nearctic region under the synonym Gnypeta minuta (Klimaszewski et al. 2018). We here newly record this taxon from MB. Dasygnypeta velata is here considered a Holarctic species as it has a broad, transpalaearctic distribution (Europe to Siberia; Newton 2019), occurs along rivers and has been collected north of the treeline in Alaska.

This species has been collected together with D. nigrella in southern Manitoba (see above). The barcode sequences of the specimens from Alaska are all partial (386 to 407 bp), but the overlapping parts of the sequences are identical to the two German sequences.

Figure 12. 

Dasygnypeta velata (Erichson) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission.

Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov.

Fig. 13A–H

Type material

Holotype. (male, CNC): Canada, Ontario, Hartington, Eel Lake, South Frontenac, Paul Hebert’s cottage property, 44.563°N, 76.549°W, 6.13.2017, Mikko Pentinsaari, Barcode of Life DNA voucher specimen, Sample ID: BIOUG34206-H01, Process ID: MPCAN465-17. Paratypes (3 CBG, 4 CNC): Canada, labelled as the holotype except: Sample ID: BIOUG34206-H02, Process ID: MPCAN466-17 (1 male, CBG); Sample ID: BIOUG34206-G12, Process ID: MPCAN464-17 (1 male, CBG); Sample ID: BIOUG34206-G11, Process ID: MPCAN463-17 (1 female, CBG). United States: North Carolina: Haywood Co., 3 mi N Dellwood, 19.VIII.1972, A. Smetana (3, CNC); Maryland: Patuxent Wildl. Res. Ctr., 5 km E Montpelier, treading pond vegetation, 16.VI.1982, Bousquet & Davies (1, CNC).

Etymology

The species epithet refers to the longitudinal impression on the pronotum, most strongly developed in males.

Diagnosis

Gnypeta impressicollis can be easily distinguished from all Nearctic species of the genus (except eastern G. baltifera (LeConte)) by the hexagonal pronotum with a longitudinal impression in the basal half (females) to nearly entire pronotal length (males). Males also have an impression on the vertex of the head. We have examined the female type of G. baltifera and it is externally similar but differs by the shorter, less angulate hexagonal pronotum, reddish and longer elytra and spermatheca with an elongate stem (C-shaped in G. impressicollis).

Description

Body length 3.2–3.4 mm; colour dark brown, elytra brown with irregular rust-brown patches, first two or three basal tergites rust-brown with posterior edge yellow, apex of abdomen rust-brown, legs and antennae rust-brown; integument highly glossy (Fig. 13A); pubescence yellowish grey, moderately long and moderately sparse; all antennomeres distinctly elongate; head round with short neck (visible only when head is distended from thorax), vertex in males with broad central impression, vertex of females with much smaller and narrower median impression, maximum width of head slightly less than maximum width of pronotum; pronotum hexagonal in shape, ca. as long as head, with a longitudinal impression in the basal half (females) to nearly entire pronotal length (males), pubescence on disc directed anteriad along midline and obliquely laterad elsewhere; elytra wider than either head or pronotum, at suture shorter than pronotum along midline, pubescence directed obliquely posteriad forming wavy pattern medially on each side; abdomen arcuate laterally, broadest in apical third, at base distinctly narrower than elytra; legs very long, hind tarsus with basal tarsomere ca. as long as the two following ones combined. MALE. Tergite VIII with apical margin truncate medially and arcuate laterally (Fig. 13E); sternite VIII elongate, narrowed apically, apex truncate medially and oblique laterally (Fig. 13F); median lobe of aedeagus in lateral view with tubus very short, triangular and gradually tapering to narrowly rounded apex, ventral margin broadly curved ventrad in apical half (Fig. 13B); in dorsal view bulbus moderately large and tubus swelled basally and triangular apically (Fig. 13C); internal sac with complex membranous structures (Fig. 13B,C). FEMALE. Tergite VIII broadly arcuate apically (Fig. 13G); sternite VIII rounded apically with very shallow median emargination (Fig. 13H); spermatheca C-shaped, capsule subspherical with broad apical invagination, stem tubular and C-shaped (Fig. 13D).

Figure 13. 

Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H).

Distribution

Origin: Nearctic. Canada: ON. United States: MD, NC. Gnypeta impressicollis is probably broadly distributed in eastern North America.

Bionomics

Specimens were collected by sifting leaf litter along a lake margin and by treading pond vegetation.

Comments

It was challenging to place this species in either Gnypeta or Ischnopoda Stephens based on the concepts of Paśnik (2010). The extremely long legs, pronotal shape, C-shaped spermatheca and superficial punctation of the pronotum and abdomen are consistent with at least some Neotropical members of Ischnopoda but the ligula of G. impressicollis is divided to the base, which is considered to be a feature of Gnypeta (Pašnik 2010). The C-shaped spermatheca of Gnypeta impressicollis also bears some similarity to the G. crebrepunctata group of Klimaszewski et al. (2008) but it is rather different in external morphology. We place this species in Gnypeta pending future systematic research.

Tribe Hypocyphtini Laporte, 1835

Oligota parva Kraatz, 1862

Fig. 14A–G

Material

(DNA barcoded specimens). Germany: Bornheim-Hemmerich, Ortslage, 50.7596, 6.93151, 30.VII.2010, F. Koehler (2, ZSM); Bornheim-Hemmerich, Ortslage, 50.7596, 6.93151, 25.VIII.2013, GBOL-Team ZFMK (1, ZFMK); Wutha-Farnroda, Wartburgkreis, 50.947, 10.4214, 25.VIII.2012, GBOL-Team ZFMK (2, ZFMK). Canada: Ontario: Kawartha Lakes, 44.296, -78.452, farm, malaise trap, 24.VII.2015, B. McClenaghan (1, CBG); same except 19.VII.2016 (1, CBG); Guelph, Arboretum, Urban Organic Farm, 43.5381, -80.222, compost heaps and mouldy hay pile, 17.IX.2017, M. Pentinsaari (4, CBG).

Distribution

Origin: West Palaearctic (adventive in North America). Canada: NB, ON [new record], PE. United States: CA, MA, MO, NV, TX.

Bionomics

This species is generally found in anthropogenic habitats, including compost, dung, and old hay and grass (Klimaszewski et al. 2018). In Canada, it has been collected in compost and in ocean coastline drift at the top of the littoral zone (Klimaszewski et al. 2018).

Comments

Oligota parva is a cosmopolitan species that is adventive in Canada. Here we newly report it from Ontario.

Figure 14. 

Oligota parva Kraatz A habitus B median lobe of aedeagus in lateral view C male tergite VIII D male sternite VIII E female tergite VIII F female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–F). Illustrations after Webster et al. (2016).

Oligota pumilio Kiesenwetter, 1858

Fig. 15A–C

Material

(DNA barcoded specimens). Belgium: Blanden, BR Meerdaalboos, 50.7976, 4.71622, 8.V.2010, F. Koehler (1, ZSM). Germany: Neuburg, Altrheine, 48.9943, 8.24412, 29.IX.2011, F. Koehler (2, ZSM); Erftstadt-Bliesheim, NWZ Altwald Ville, 50.792, 6.844, 4.X.2010, F. Koehler (1, ZSM); Jockgrim, Sandmagerrasen, 49.0805, 8.26568, 14.XI.2010, F. Koehler (1, ZSM); Ochtendung, Michelsberg, 50.3631, 7.3889, 17.III.2012, F. Koehler (1, ZSM); Edenkoben-Rhodt, Villa Ludwigshoehe, 49.2767, 8.08991, 20.V.2012, F. Koehler (1, ZSM); Bad Muenster-Traisen, Rotenfels, 49.822, 7.832, 20.V.2012, F. Koehler & J. Koehler (1, ZSM); Osterholz bei Blankenburg, 51.9519, 11.0526, 18.III.2015, GBOL-Team ZFMK (2, ZFMK). Canada: Alberta: Waterton Lakes National Park, Red Rock Parkway, 49.088, -113.883, Moraine grassland, intercept trap, 1328 m, 11.VIII.2012, BIOBus 2012 (1, CBG). United States: Montana: Missoula County, Florence, MPG Ranch, 46.702, -114.034, grassland, pitfall trap, 05.VI.2019, M. Seidensticker (1, CBG).

Distribution

Origin: West Palaearctic (adventive in North America). Canada: AB [new record]. United States (all except MT need verification): DC, IL, OH, MT [new record].

Diagnosis

Among Canadian species of Oligota, O. pumilio is extremely similar to O. pusillima in the narrow, parallel body (Fig. 15A) and in male and female genitalia (Klimaszewski et al. 2018). However, it can be distinguished by the more abruptly truncate apex of the median lobe in lateral view (Fig. 15B), differently shaped sclerites of the internal sac (Fig. 15B), medially projected apex of male sternite VIII, and the transverse capsule of the spermatheca (Fig. 15C) (Kapp 2019).

Figure 15. 

Oligota pumilio Kiesenwetter A habitus (image by A. Bogri – www.BilleBank.dk) B median lobe of aedeagus in lateral view (drawn from Kapp 2019) C spermatheca (drawn from Kapp 2019). Scale bars: 1 mm (A); 0.2 mm (B, C).

Bionomics

This species occurs in a wide variety of habitats across a broad elevational range, including hollow trees, plant debris, old hay in cattle barns, moldy substrates and in mushrooms (Kapp 2019). The barcoded Nearctic specimens were collected from grassland habitats by an intercept trap (Alberta) and a pitfall trap (Montana).

Comments

Oligota pumilio is a West Palaearctic species that is adventive in Canada. Although it has been previously reported from the United States (OH, IL, DC) (Newton 2019), these records need confirmation as they are in the east, some distance away from the present records. This species’ presence in North America is thus verified here for the first time, from both Canada (AB) and United States (MT). It has also been reported as adventive from Argentina, Chile, and New Zealand (Newton 2019). The barcoded specimens of O. pumilio from Canada and the United States share the same DNA barcode haplotype, which is also shared by some of the specimens from Germany.

Oligota pusillima (Gravenhorst, 1806)

Fig. 16A–G

Material

(DNA barcoded specimens). Finland: Oba: Oulu, Hietasaari, 65.0225, 25.4247, 22.IV.2011, M. Pentinsaari (1, ZMUO); Germany: Edenkoben-Rhodt, Villa Ludwigshoehe, 49.277, 8.09, 20.V.2012, F. Koehler (2, ZSM); Edenkoben-Rhodt, Villa Ludwigshoehe, 49.277, 8.09, 23.VI.2012, F. Koehler (2, ZSM); Zweibruecken-Mauschbach, Monbijou-Wald, 49.2038, 7.39891, 16.X.2011, F. Koehler & W. Koehler (1, ZSM). Canada: Alberta: Two Hills, Two Hills School EQP-CLL-553, 53.7104N, 111.7437W, 613 m, Malaise trap, 21.IX.–2.X.2015, K. Warawa (2, CBG).

Additional non-barcoded material

Ontario: Ottawa, Ottawa River, Deschênes Lookout, Berlese flood debris, 1.V.1985, A. Davies (1, CNC).

Distribution

Origin: West Palaearctic (adventive in North America). Canada: AB [new record], NB, ON [new record]. United States: MA, NY.

Bionomics

This species occurs in a variety of moist to dry, decaying organic matter including rotting hay, compost, hollow trees, and ant nests (Kapp 2019). Canadian specimens were collected in compost (Webster et al. 2016), and in malaise traps and flood debris in a suburban setting (present study).

Comments

Oligota pusillima is a Palaearctic species that has been introduced to North America, South America, Australia, Africa, and southeast Asia (Kapp 2019, Newton 2019). It is here reported from Ontario and Alberta for the first time, the latter representing the westernmost record in North America. Specimens from Alberta were sequenced and their barcodes match those of Palaearctic specimens with no divergence. These specimens were also morphologically consistent with O. pusillima.

Figure 16. 

Oligota pusillima (Gravenhorst) A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G).Illustrations after Webster et al. (2016).

Homalotini Heer, 1839

Anomognathus athabascensis Klimaszewski, Hammond & Langor, 2016

Fig. 17A–G

Material

Non-sequenced specimens. Canada: Manitoba: Winnipeg, under bark of rotten ‘N. aceroides’ [= Acer negundo], 27.VIII.1918, J.B. Wallis (2, CNC).

Distribution

Origin: Nearctic. Canada: AB, MB [new record].

Bionomics

The MB specimens were collected under bark, confirming that this species lives in a way similar to other members of the genus.

Comments

Anomognathus athabascensis, recently described from Alberta (Klimaszewski et al. 2016b), is newly reported from Manitoba. This native Nearctic species is likely transcontinental but rarely reported due to its small size and elusive habits.

Figure 17. 

Anomognathus athabascensis Klimaszewski, Hammond & Langor A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2016b).

Anomognathus cuspidatus (Erichson, 1839)

Figs 18A–D, 19A–D, 20A–D.

Homalota cuspidata Erichson, 1839

Thectura americana Casey, 1893, syn. nov.

Anomognathus americanus: Seevers (1978) (as valid species)

Type material

Homolota cuspidata Erichson, 1839. Lectotype, male, here designated (ZMHB): cuspidata Er: [handwritten label] / 5387 [typed label] / Hist.-Coll. (Coleoptera), Nr. 5387, Homalota cuspidata, Erichs., Europa, Zool. Mus. Berlin [typed white label] / Lectotype Homalota cuspidata des. J. Klimaszewski 2019 [white printed label]. Paralectotypes (3, ZMHB, without original labels): Hist.-Coll., (Coleoptera), Nr. 5387, Homolota cuspidata Erichs., Europa, Zool. Mus. Berlin; Syntype Homolota cuspidata Erichson, 1837, labelled by MNHUB 2010; Paralectotype Homalota cuspidata des. J. Klimaszewski 2019 [white printed label] [1 female, spermatheca and terminalia dissected in Canada balsam on microslide attached to specimen]; same labels except: SYNTYPUS, Homalota cuspidata Erichson, 1837 [typed red label, added by MNHUB 2010] [1 female, spermatheca and terminalia dissected in Canada balsam on microslide attached to specimen]; same labels as before [1 damaged specimen, sex undetermined].

Males and females of the syntype series were morphologically consistent with the specimens forming molecular cluster BOLD:AAO0339, including those sequenced from Ontario, Canada. As the most obvious difference between A. cuspidatus and the potential new Central European species (see Diagnosis) was the shape of the median process on male tergite VIII (in lateral view) (Fig. 18B, D), a male syntype (see above) was designated as the lectotype of this species to fix its identity. Morphology of the aedeagus itself was difficult to study due to its small size and obvious differences between molecular clusters (see below) were not observed (Fig. 19).

Figure 18. 

A, B Anomognathus cuspidatus (Erichson), and C, D Anomognathus sp., putative undescribed species (Europe) A, C habitus and B, D male tergite VIII in lateral view. Scale bars: 1 mm (A, C); 0.2 mm (B, D).

Thectura americana Casey, 1893, syn. nov. Holotype (male) (NMNH): NY/ TYPE USNM 39614/ Thectura americana Casey (handwritten by Casey).

Figure 19. 

Aedeagi of A–C Anomognathus cuspidatus (Erichson) and D potential undescribed species, in lateral view (top row) and dorsal view (bottom row) A sequenced non-type (Ontario, Canada) B lectotype of A. cuspidatus (‘Europe’) C, D sequenced non-types (Finland). Scale bar: 0.2 mm.

Casey (1893) gave numerous characters to distinguish A. americanus from A. cuspidatus but all of these were observed to be highly variable within populations in the material studied, including the shape of apical antennomeres, shape of the pronotum, position of the abdominal tubercles in the male, and the type of dorsal expansion of the median process of male tergite VIII. We could not find the depression at the base of tergite VIII on the holotype of A. americanus mentioned by Casey (1893). Although the aedeagus of the holotype was not studied (not extracted from partly damaged and fragile pygidium), male tergite VIII was intact and its median process in lateral view bears an apical hook, matching the present concept for A. cuspidatus (Fig. 20). Therefore, in corroboration with Fenyes (1918), we treat A. americanus as a synonym of A. cuspidatus.

Figure 20. 

Male tergite VIII of A–C Anomognathus cuspidatus (Erichson) and D potential undescribed species, in dorsal (top row) and lateral (middle and bottom rows) A lectotype of A. cuspidatus (‘Europe’) B holotype of A. americanus (Casey) (= A. cuspidatus) C, D sequenced, non-types (Finland). Scale bar: 0.2 mm.

Non-type material

(sequenced specimens indicated in square brackets). Canada: Alberta: Peace River, 25 km NW Peace River, 17–23.VIII.1993, J. Hammond (2, CNC); Ontario: Wellington County, Guelph, Eramosa River Trail, 43.539, -80.236, deciduous forest, 14.IV.2017, M. Pentinsaari (4, CBG [4 barcoded]).

A photo record of this species from Ontario is available on bugguide.net (/view/1816108): Toronto, 19.V.2020, under bark, O. Strickland.

Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 28.IV.2010, F. Koehler (1, ZSM [1 sequenced]). Czech Republic (all CNC): Bohemia, Poděbrady 50 km, Smetana, 1959, car net trap (1); Bohemia, Chvojno, Smetana (1); Moravia, Drnholec, Smetana (1). Denmark (all NHMD): Staksrode, EJ, 24.IX.1983 (1); Æbelø F, 18.V.1997 (1); Faested Mose, SJ, 12.IV.1986 (1); Dyrehaven, 14.4.1934 (1); same except 21.3.1923 (1); same except 21.10.1932 (1); 30.4.1922 (1); same except 19.5.1911 (1); Lyng Huse, 29.3.1997 (1).

Germany: Nationalpark Mueritz, Babke-Zartwitz-Speck-Schwarzenhof, 53.4125, 12.8463, car net, 20.VI.2015, GBOL-Team ZFMK (2, ZFMK [2 sequenced]); Hoenningen bis Insul, Ahrtal, 50.45, 6.942, 24.IV.2010, F. Koehler (1, ZSM [1 sequenced]); Oberheimbach, Franzosenkopf, 50.004, 7.805, 27.V.2012, W. Koehler (1, ZSM [1 sequenced]). Finland (all ZMUO): N: Espoo, Saunalahti, 60.1643, 24.6263, 17.IX.2012, fungusy aspen logs, E. Helve (1) [barcoded]; Al: Bjoerkoe, 59.9769, 20.1879, sifting, 24.IX.2014, M. Pentinsaari (1) [barcoded]; Ta: Lammi, R. Linnavuori leg. (1); Ab: Naantali, R. Linnavuori leg. (1); Kb: Lieksa, R. Linnavuori (1); Rynmattyla, 24.VI.1945, Karvonen (2); same except 14.VIII.1945 (2). Slovakia (all CNC): Cenkov, Smetana, 1963 (11); Nová Sedlica, Smetana, 1961 (2); Ruská Poruba, Smetana, 1956 (2). United Kingdom (all CNC): Essex (6).

Putative undescribed Anomognathus (corresponding to BIN BOLD:ACA9191):

Finland (all ZMUO): N: Espoo, E. Helve, 1978 (1); same except 1976 (1); same except 1977 (1); same except 1979 (1); same except 1981 (1); same except 1982 (1); Ks: Taivalkoski, 728.53 Window trap, 2003, E. Hurme (2); same except Polyporus trap (1); Kb, Kitee, 23.05.2016, M. Pentinsaari leg., [1 sequenced]; Obb: Rovaniemi, Rovajärvi, 16.6–8.7.2010, M. Pentinsaari and E. Kuusela [1 sequenced]. Germany: Schleiden-Wolfgarten, Dachsloecher, 50.6098, 6.42237, 26.VII.2012, F. Koehler (1, ZSM [1 sequenced]).

Diagnosis

Anomognathus cuspidatus is distinctive for its trident-shaped apex of male and female tergite VIII (Fig. 20A–C) and can be distinguished from all described species by this feature alone. However, in the course of this study, specimens representing a remarkably divergent barcode cluster (BOLD:ACA9191; 9.63% uncorrected p-distance to A. cuspidatus) were investigated and found to likely represent an undescribed species of Anomognathus in Europe (confirmed specimens from Finland and Germany). Although most morphological characters of A. cuspidatus and the putative new species are highly variable, including the median lobe of the aedeagus, males can be dependably separated based on the shape of their median process of tergite VIII in lateral view: A. cuspidatus bears a minute to distinct hook at the apex (Fig. 20A–C), while in the undescribed species, the median process converges evenly to a single point, creating an elongate, turnip-shape (Fig. 20D). The shape of tergite VIII in females was observed to be extremely variable and no features were deemed to be diagnostic. Externally, most specimens can be recognized as either species (especially males) by the relative proportions of the head versus the pronotum, with A. cuspidatus generally bearing a small pronotum, narrower than the head (Fig. 18A) and the undescribed species bearing a wider, longer pronotum, wider than the head (Fig. 18C). The limits of this taxon need further investigation and should include morphological study of a much wider range of sequenced material.

Distribution

Origin: West Palaearctic (adventive in North America). Canada: AB, NB, ON. United States: NY.

Bionomics

This species occurs under the bark of dead trees. One specimen (NB) was collected from a Lindgren funnel.

Comments

Anomognathus cuspidatus is a widespread West Palaearctic species that is known from Europe, European Russia and Algeria (Newton 2019) and has been previously known in North America under the synonym A. americanus. The record from Beijing, China should be verified. The species has become introduced in North America (before 1893) and it is unclear whether the population in Alberta represents a separate introduction from Europe, a secondary introduction from eastern North America or a broad adventive distribution across Canada.

After the results of the present study, two species of Anomognathus are known to occur in North America: native A. athabascensis Klimaszewski, Hammond & Langor and the adventive A. cuspidatus. These are easily separated by the drastically different shapes of male and female tergites VIII (Figs 17D, 20A–C). Previously, only females of A. cuspidatus (as A. americanus) were available from Canada (Klimaszewski et al. 2016b; Webster et al. 2016). Here we demonstrate that all available Nearctic Anomognathus specimens with a trident-shaped tergite VIII correspond to Palaearctic A. cuspidatus.

Cyphea wallisi Fenyes, 1921

Figs 21A–G, 22A–D

Cyphea wallisi Fenyes, 1921

Agaricomorpha vincenti Klimaszewski & Webster, 2016, syn. nov.

Agaricomorpha vincenti: Klimaszewski et al. 2018 (as synonym of C. curtula)

Type material

Cyphea wallisi Fenyes, 1921. Paratype, male (MCZ). Winnipeg, Man. [handwritten label] / Wallis [handwritten label] / 25490. / Cyphea, Wallisi, Feny [handwritten label] / Type., 9989, 9983 [typed red label].

Agaricomorpha vincenti Klimaszewski & Webster, 2016, syn. nov. Holotype, male (LFC). Canada, New Brunswick, Carleton Co., Jackson Falls, “Bell Forest”, 46.2200°N, 67.7231°W, 7–21.VI.2012, C. Alderson & V. Webster, coll. [white typed label] / Rich Appalachian hardwood forest, Lindgren funnel trap in canopy of Fagus grandifolia [white typed label] / Holotype Agaricomorpha vincenti Klimaszewski & Webster, 2016 [red typed label] / Cyphea curtula (Erichson) det. Klimaszewski 2017 [white typed label] / Cyphea wallisi Fenyes det. A. Brunke 2020.

Figure 21. 

Cyphea wallisi Fenyes A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2018), used with permission.

The aedeagi of the male paratype (holotype in collection of the California Academy of Sciences) of C. wallisi and holotype of A. vincenti are identical and both differ from that of Palaearctic Cyphea curtula (image by V. Assing) by the broader distal lobe in lateral view, which only slightly extends beyond the distal plate (Fig. 22A–C). Therefore, we transfer Agaricomorpha vincenti from synonymy with Cyphea curtula to synonymy with Cyphea wallisi.

Figure 22. 

Aedeagi of Cyphea wallisi Fenyes (A, B, D) and C. curtula (Erichson) (C), in lateral (A–C) and dorsal (D) view. Paratype of C. wallisi (A, D); holotype of Agaricomorpha vincenti Klimaszewski and Webster (= C. wallisi) (B); non-type, C. curtula (image by V. Assing) (C). Scale bar: 0.2 mm.

Other material

(DNA barcoded specimens). Canada: Ontario: Rouge National Urban Park, Toronto Zoo, 43.8223, -79.1897, forest, malaise trap, 25.VI.2013, L. Attard and K. Greenham (2, CBG); Hartington, Eel Lake Cottage, Lindgren funnel trap, 44.5628, -76.553, 25.VII.2017, G. Blagoev (1, CBG); Nova Scotia: Clyburn Valley Road, near golf course, Cape Breton National Highlands NP, forest, Malaise trap, 46.6553, -60.4285, 28.VI.2013, CBH staff (1, CBG).

Non-barcoded specimens

Canada: Quebec: Mont St. Bruno Prov Park, 45.541, -73.319, Lindgren funnel, trap 5, tree 2, beech-maple canopy, 21.VII-3.VIII.2005 (1, CNC); Oka Prov Park, Lindgren funnel, trap 3, tree 1, beech-maple canopy, 27.VII.30.VIII.2005 (1, CNC).

Distribution

Origin: Nearctic. Canada: AB, MB, NB, NS [new record], ON [new record], QC.

Bionomics

Specimens have been collected in Malaise traps, window traps and Lindgren funnels placed in forests. Both the closely related West Palaearctic C. curtula and C. latiuscula Sjöberg have been consistently collected under bark, where they occur in the larval burrows of various longhorn beetles (Cerambycidae), bark beetles (Curculionidae: Scolytinae) and the carpenter moth (Cossus L.) (Palm 1968).

Comments

Cyphea wallisi is a broadly distributed native Nearctic species, reported from AB east to NS. Here we treat Nearctic records of Cyphea as C. wallisi (previously treated as Palaearctic C. curtula, e.g., Klimaszewski et al. 2018) and newly report the genus from ON and NS. Cyphea wallisi is probably far more broadly distributed in North America than currently known and has been underreported due to its small size.

Sequenced Nearctic specimens of Cyphea from ON and NS formed a barcode cluster that was nearly 5% divergent from those of Palaearctic specimens of C. curtula (BOLD:AAO1175, one published sequence record from Belgium and three unpublished records from the Netherlands). Northern European C. latiuscula, the only other species of the genus, has a broader body outline, different male genitalia and is quite differently colored (bicolored pronotum and pale elytra). No barcode sequence data are currently available for C. latiuscula. Based on the study of one paratype of C. wallisi, described from Manitoba and not reported since, it was discovered that Nearctic specimens of Cyphea correspond to this species and differ from Palaearctic C. curtula by the broader distal lobe of the median lobe of the aedeagus in lateral view, which only slightly extends beyond the distal plate (Fig. 22A–C). The shape of the median lobe of the aedeagus in dorsal view may also be diagnostic (Fig. 22D) but a preparation in this view was unavailable for C. curtula. The illustration in Palm (1968) of the aedeagus of C. curtula in dorsal view appears to be less angulate than that of C. wallisi but this needs verification. Based on these differences in male genitalia (Fig. 22A–C) and the COI barcodes, Cyphea wallisi is morphologically and genetically distinct from Palaearctic C. curtula, and the latter species does not occur in North America as far as known. Both of these species have a median tubercle on male tergite VII, mentioned earlier by Fenyes (1921) but this structure was omitted from the illustrations in Klimaszewski et al. (2018), though it was present in the original description of synonym Agaricomorpha vincenti (Webster et al. 2016). Previous differences between the two species given by Klimaszewski et al. (2018) (e.g., projecting pronotal angles, lighter/darker body) proved to be highly variable.

Gyrophaena affinis Mannerheim, 1830

Fig. 23A–G

Material

(DNA barcoded specimens). Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 135 m, 16.VI.2010, F. Koehler (1, ZSM). Finland: Oba: Oulu, Linnanmaa, 65.0633, 25.4712, 7.VI.2011, M. Pentinsaari (1, ZMUO); Obb: Tornio, Kalkkimaa, 65.9014, 24.4711, 10.VII.2012, M. Pentinsaari (1, ZMUO); Al: Lemland, Herrövägen, 59.9796, 20.1954, car net, 5.VII.2012, M. Pentinsaari (1, ZMUO). Germany: Brohl-Luetzing, Brohltal, 50.4727, 7.31272, 22.V.2010, F. Koehler (1, ZSM), Riedlhuette, Diensthuettenstrasse, 48.937, 13.412, 09.VII.2011, F. Koehler & M. Koehler (1, ZSM), Waldhaeuser, Lusen- und Boehmstrasse, 48.93, 13.492, 09.VII.2011, F. Koehler & M. Koehler (1, ZSM). Canada: Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane forest, 27.VI.2012, BIOBus 2012 (2, CBG).

Distribution

Origin. Uncertain. Canada: AB [new record], BC, MB, NB, NF, NS, ON, QC, SK. United States: AZ, DC, IL, IN, IA, KY, MA, ME, MI, MN, MO, NC, NH, NJ, NM, NY, OH, PA, TN, WA, WI, WV.

Comments

Gyrophaena affinis is newly reported from AB based on barcoded material.

Sequenced Nearctic specimens from ON, AB, NB, and QC form a distinct barcode cluster, separate from all sequenced Palearctic specimens and divergent by 4.65%. This pattern is inconsistent with a species that is adventive in North America and we remove G. affinis from the list of adventive species in Canada. In comparing images between those of Nearctic specimens (Fig. 23B) and those of Enushchenko and Semenov (2016) for Palaearctic specimens, there appear to be slight differences in the median lobe of the aedeagus in lateral view. In the Palaearctic illustration, the apex of the median lobe is more acute and its secondary lobe is evenly rounded at apex, while the Nearctic illustration shows a more rounded apex of the median lobe and knob-like apex of the secondary lobe (Fig. 23B). More research is needed to determine the status of the Nearctic and Palaearctic populations, though the level of genetic divergence between discrete Nearctic and Palaearctic populations suggests that two sister species are involved.

Figure 23. 

Gyrophaena affinis Mannerheim A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2018), used with permission.

Gyrophaena gracilis Seevers, 1951

Fig. 24A–H

Material

(non-sequenced material). Canada: Quebec: Gatineau Park, wolf trail, near trail start, 45.541, -75.912, hardwood forest, Polyporus squamosus on large beech log, 8.VI.2019, A. Brunke & J. Smith (1, CNC).

Distribution

Origin. Nearctic. Canada: NB, QC [new record]. United States: WI.

Bionomics

Specimens have been collected from a partly dried Pleurotus mushroom, from within the pores of a Trametes polypore, and from the nest contents of a Barred owl (Strix varia Barton) (Klimaszewski et al. 2018). The specimen from QC was collected from Polyporus squamosus on a beech tree.

Comments

The new record from QC, near the ON border, bridges the wide gap between previous records in NB and WI.

Figure 24. 

Gyrophaena gracilis Seevers A habitus B, C median lobe of aedeagus in lateral view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission.

Gyrophaena simulans Seevers, 1951

Fig. 25A–G

Material

(DNA barcoded specimens). Canada: Ontario: Hartington, Eel Lake Cottage, 44.563, -76.549, deciduous forest, mushrooms, 4.X.2017, M. Pentinsaari (2, CBG).

Distribution

Origin. Nearctic. Canada: ON [new record]. United States: IL, MD, PA.

Diagnosis

Gyrophaena simulans is extremely similar to G. criddlei and G. pseudocriddlei but has a slightly more transverse and flatter pronotum, with straighter apical and basal margins, and differently shaped upper process of the median lobe in lateral view (Fig. 25B): longer than that of G. pseudocriddlei but shorter and broader than that of G. criddlei. The emargination of male tergite VIII in G. simulans appears to be shallower and broader than that of G. criddlei but more specimens are needed to confirm this.

Figure 25. 

Gyrophaena simulans Seevers A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G).

Bionomics

The Canadian specimens were collected by sifting mushrooms in a deciduous forest. No detailed data on the host fungus were recorded.

Comments

Gyrophaena simulans is a native Nearctic species distributed in eastern North America and is newly reported from Canada. The barcode cluster BOLD:ACY8004 also contains specimens identified as related species G. criddlei (female) and G. pseudocriddlei but more research, with broader sampling of sequenced, identified males, is needed to determine whether these species share a BIN or these specimens are misidentified. As we were unable to verify the identifications at this time, these records are not published here.

Homalota plana (Gyllenhal, 1810)

Fig. 26A–G

Material

(DNA barcoded specimens). Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 28.IV.2010, F. Koehler (1, ZSM). Germany: Arnsberg-Breitenbruch, NWZ Hellerberg, 51.446, 8.135, 30.V.2011, F. Koehler (2, ZSM); Heimbach-Blens, Linkheld, 50.648, 6.468, 29.VIII.2012, F. Koehler (2, ZSM); Erftstadt-Bliesheim, NWZ Altwald Ville, 50.7917, 6.84384, 03.VI.2011, F. Koehler (1, ZSM); westl. Klein-Quenstedt, 51.9239, 11.0478, 20.III.2015, GBOL-Team ZFMK (1, ZFMK). Finland: Al: Finström, Norrö, 60.2458, 19.822, 5.VII.2012, M. Pentinsaari (1, ZMUO); Ka: Joutseno, Kuurmanpohja, 61.071, 28.75, 3.VIII.2012, M. Pentinsaari (1, ZMUO). Canada: Ontario: Guelph, Eramosa River Trail, 43.539, -80.236, deciduous forest, 14.IV.2017, M. Pentinsaari (2, CBG).

Distribution

Origin. Uncertain. Canada: AB, MB, NB, NF, NS, ON. United States: AZ, CA, CO, IA, ID, IN, MT, NY, OH, PA, TX.

Bionomics

Specimens occur under bark of dead trees.

Comments

Sequenced Nearctic specimens from ON form a distinct barcode cluster, separate from all sequenced Palearctic specimens and divergent by 7.58%. This pattern is inconsistent with a species adventive in North America and we remove H. plana from the list of adventive species in Canada. Preliminary comparisons between images of Palaearctic and Nearctic specimens revealed that there may be some slight differences in the shape of the spermatheca. More research is needed to determine the status of the Nearctic and Palaearctic populations, though the level of genetic divergence between discrete Nearctic and Palaearctic populations suggests that two sister species are involved.

Figure 26. 

Homalota plana (Gyllenhal) A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2018), used with permission.

Thecturota tenuissima Casey, 1893

Fig. 27A–G

Thecturota tenuissima Casey, 1893

Atheta marchii Dodero, 1922, syn. nov.

Pragensiella magnifica Machulka, 1941, syn. nov.

Thecturota marchii: Muona 1984 (as valid species)

Thecturota magnifica: Schülke and Smetana 2015 (as syn. of T. marchii)

Material

(DNA-barcoded specimens). Germany: Kobern-Gondorf, Ortslage/Weinberge, 50.308, 7.460, 21.V.2010, F. Koehler (1, ZSM); Edenkoben-Rhodt, Villastrasse, 49.279, 8.092, 20.X.2012, F. Koehler (1, ZSM). Finland: Oba: Oulu, Linnanmaa, 65.0633, 25.4712, botanical garden, compost heap, flight-intercept trap, 7.VI.2011, M. Pentinsaari (1, ZMUO).

Non-sequenced material

Several males and females of T. tenuissima from Denmark (NMHD) were compared with illustrations from Klimaszewski et al. (2017).

Distribution

Origin. Nearctic (adventive in West Palaearctic). Canada: ON, QC. United States: RI.

Bionomics

Canadian specimens were collected by car-netting in mixedwood forests, while Palaearctic specimens are known from compost and other plant-based debris (Horion 1967).

Comments

Thecturota tenuissima is native to the Nearctic region and has become accidentally introduced to the West Palaearctic, including the Canary Islands, where it was previously known under the synonym T. marchii (Newton 2019). We expect this species to be broadly distributed in eastern North America and has been overlooked over much of its range because car-netting, an effective method for collecting small, obscure staphylinids, is rarely used in the Nearctic region.

Nearctic and Palaearctic populations do not differ in male and female genitalia or in external morphology. Molecular data were unavailable for the Nearctic population, which was recently reported from Canada (Klimaszewski et al. 2017) but described from Rhode Island, USA in 1893 (Casey 1893). However, we are confident that these species are synonyms. Muona (1984) stated that T. marchii is a ‘recent’ introduction to Europe but from an unknown source. Thecturota is primarily a New World genus, with ten described species in North and South America (Newton 2019). Thecturota magnifica (Machulka) is currently treated as a synonym of T. marchii (Newton 2019) and we simply transfer this name to synonymy with T. tenuissima. The only Palaearctic species remaining is poorly known T. williamsi (Bernhauer, 1936), known only from the type collected in Great Britain and probably a synonym of T. tenuissima. The characters Bernhauer (1936) gave to separate his species from T. tenuissima (as T. marchii) are slight differences in coloration and body proportions, which are both highly variable in the Palaearctic specimens of T. tenuissima studied. Therefore, we consider T. tenuissima to be a native Nearctic species that has become adventive in the West Palaearctic and suggest that genus Thecturota is naturally restricted to the Nearctic and Neotropical regions.

Figure 27. 

Thecturota tenuissima Casey A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2017).

Geostibini Seevers, 1978

Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov.

Fig. 28A–G

Type material

Holotype (male) (CNC): Canada: ON: Waterloo County, Cambridge, Rare Charitable Research Reserve, 43.390, -80.374, soybean field, pitfall trap, 29.VI.2010, A. Brunke [white printed label] / HOLOTYPE Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari sp. nov., des A. Brunke 2020 [red printed label]. Paratypes (1 NMNH, 6 DEBU, 1 CNC): Canada: ON: Wellington County, Eramosa, 43.616, -80.215, soybean field, pitfall trap, 13.VII.2010, A. Brunke [white printed label] / PARATYPE Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari sp. nov., des A. Brunke 2020 [yellow printed label] [CNC, DEBU, 7 specimens]. United States: VA: Arlington County, Marcey Creek, 38.9087, -77.1083, 70 m, suburban backyard, Malaise trap, 14–21.VI.2015, S. Miller [white printed label] / Barcode of life, DNA voucher specimen, Sample ID: BIOUG42376-E12, Process ID: GMUAF1698-18 [yellow printed label] / PARATYPE Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov., des A. Brunke 2020 [yellow printed label] [NMNH, 1 specimen].

Non-type material

(barcoded specimens). Canada: Ontario: Guelph, 25 Division St., 43.554, -80.264, Malaise trap, 14.VII.2010, A. Smith (1, CBG); Guelph, John F. Ross CVI, 43.5621, -80.247, Malaise trap, 22.IV-03.V.2013, G. Staines (1, CBG); Milverton, Milverton Public School, 43.568, -80.928, Malaise trap, 22.IV-03.V.2013, J. Van Bakel (1, CBG); Collingwood, Collingwood Collegiate Institute, 44.489, -80.215, 188 m, Malaise trap, 22.IV-05.V.2014, A. Breton (1, CBG); Cambridge, rare Charitable Research Reserve, 43.3736, -80.3652, 304 m, 04–11.VI.2015, BIO Collections Staff (1, CBG).

Etymology

The species epithet refers to the similarity to related species A. gregaria (Erichson), which was originally treated separately from other Aloconota under subgenus Glossola Fowler (e.g., Benick 1954) because it lacks obvious male secondary sexual characters.

Diagnosis

Aloconota pseudogregaria can be easily distinguished from all other species of the genus occurring in eastern North America by the distinctly bicolored abdomen (Fig. 28A). Among Central European species, the spermatheca of A. pseudogregaria is most similar to that of Palaearctic A. gregaria but in the latter the apex is distally truncate, median lobe is distinctly sinuate and only weakly projected ventrad, the abdomen is darker and not distinctly bicolored, and the microsculpture of the forebody is much stronger, creating a dull reflection.

Figure 28. 

Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII (structure accidentally over-cleared in balsam preparation) F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G).

Description

Body length 2.4–2.7 mm, moderately flattened (stronger so on elytra), narrowly subparallel, colour of head, pronotum, scutellar region of elytra, apical part of abdomen and antennomeres 5–11 dark brown to dark reddish brown, elytra and antennomeres 1–3 paler, red-brown and legs yellow; forebody finely and densely punctate, microsculpture shallow, consisting of meshes; head slightly elongate and with small, shallow impression medially, head slightly narrower than pronotum, postocular region elongate, ca. as long as maximum diameter of eye, tempora with carinae dorsally only; antennae slender, as long as pronotum and elytra combined, basal three antennomeres strongly elongate, 4 subquadrate, 5–10 subquadrate to slightly transverse, and terminal one strongly elongate and ca. as long as two preceding antennomeres combined; pronotum slightly transverse (width/length ratio 1.3), trapezoidal in shape, flattened, pubescence directed straight posteriad in central part of disc and obliquely posteriad laterally; elytra at suture ca. as long as pronotum along midline, flat, distinctly transverse (width/length ratio 1.5), ~ 1/3 broader than pronotum, humeri angular, posterior margins slightly sinuate laterally, pubescence directed straight posteriad forming slightly arcuate lines in sutural region of disc; abdomen subparallel, tergites III–VI distinctly impressed at base; basal metatarsomere ~ 1/3 longer than the following one. MALE. Tergite VIII rounded apically with minute median emargination, lacking apical teeth (Fig. 28D); sternite VIII rounded apically (Fig. 28E); tubus of median lobe of aedeagus long, ventrally ca. straight in basal two-thirds and moderately projecting ventrad apically in lateral view (Fig. 28B). FEMALE. Tergite VIII rounded apically and slightly pointed medially (Fig. 28F); sternite VIII rounded apically (Fig. 28G); spermatheca S-shaped, capsule pitcher-shaped with short neck, stem strongly sinuate and swollen apically (Fig. 28C).

Distribution

Origin: Nearctic. Canada: ON. United States: VA.

Bionomics

This species has only been collected by passive traps, including malaise and pitfall traps. All specimens have been collected from at least partly disturbed habitats, such as forest edges, agricultural fields, and suburban environments. This species corresponds to ‘Aleocharinae sp. 5’ in Brunke et al. (2014), which was collected in both soybean fields and adjacent forest edges by pitfall traps.

Comments

Aloconota pseudogregaria is probably broadly distributed in northeastern North America. We have compared the male and female genitalia of A. pseudogregaria with all Central European and Nearctic species of Aloconota, and are confident that this taxon has not been previously described from Europe or North America, despite its occurrence in disturbed habitats in North America, which is typical for introduced species. Although Aloconota pseudogregaria clustered most closely with A. gregaria (BOLD:ABU6164) in our barcode dataset, its BIN is ~ 8% different from that of the latter. Based on morphology of the aedeagus and spermatheca, Aloconota pseudogregaria is probably even more closely related to East Palaearctic Aloconota described from Japan and Korea (e.g., Sawada 1970 [as Tomoglossa], Lee and Ahn 2017) rather than to A. gregaria. However, the described species all differ markedly in external morphology.

Athetini Casey, 1910

Atheta (Datomicra) nigra (Kraatz, 1856)

Fig. 29A–G

Material

(DNA-barcoded specimens). Canada: Ontario: Peterborough, 44.253N, 78.415W, farm, malaise trap, 24–30.V.2015, B. McClenaghan (1, CBG). Germany: Koeln-Worringen, Worringer Bruch, 51.044, 6.87427, 01.VII.2010, F. Koehler & J. Koehler (1, ZSM).

Distribution

Origin. Palaearctic (adventive in North America). Canada: ON [new record], SK.

Bionomics

Canadian specimens have been collected on farmland and directly from horse manure.

Comments

Atheta nigra is a Palaearctic species reported from across Europe, European Russia, Kazakhstan, North Korea and southern China (Newton 2019). It is adventive in North America and New Zealand (Newton 2019) and is here newly reported from Ontario. The new record from Ontario indicates that this species is far more widely distributed in North America than previously known.

Figure 29. 

Atheta (Datomicra) nigra (Kraatz) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in ventral view D spermatheca E apical part of dorsal male abdomen showing 4 dents on tergite VIII F female tergite VIII G female sternite VIII; A, D, F, G after Klimaszewski et al. (2016a), based on female from Saskatchewan, Canada B, C after Benick and Lohse (1974) E after Klimaszewski et al. (2016a), based on a male from Germany. Scale bars: 1 mm (A); 0.2 mm (B–G).

Mocyta scopula (Casey, 1893), comb. nov.

Fig. 30A–F

Eurypronota scopula Casey, 1893

Pancota laetabilis Casey, 1906

Dolosota abundans Casey, 1910

Dolosota flaccida Casey, 1910

Dolosota redundans tergina Casey, 1910

Dolosota scopula: Casey (1910) (as type species of Dolosota)

Dolosota secunda Casey, 1910

Dolosota sequax Casey, 1910

Acrotona (Dolosota) abundans: Seevers (1978) (as valid species)

Acrotona (Dolosota) flaccida: Seevers (1978) (as valid species)

Acrotona (Dolosota) scopula: Seevers (1978) (as valid species) (Dolosota syn. of Acrotona, in part; some species moved to Pancota)

Acrotona (Dolosota) secunda: Seevers (1978) (as valid species)

Acrotona (Dolosota) sequax: Seevers (1978) (as valid species)

Pancota laetabilis: Seevers (1978) (as valid species)

Pancota redundans tegrina: Seevers (1978) (implied, subspecies not directly mentioned)

Acrotona abundans: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b))

Acrotona flaccida: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b))

Acrotona laetabilis: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b))

Acrotona redundans tergina: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b))

Acrotona scopula: Majka and Sikes (2009) (valid species following Gusarov (2003b))

Acrotona secunda: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b))

Acrotona sequax: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b))

Material

(DNA-barcoded specimens). Canada: Ontario: Georgian Bay Islands National Park, Fairy Lake, 44.8929, -79.8514, mostly conifer forest with moss, Berlese funnel, 5.VIII.2015, BIObus 2015 (1, CBG).

Distribution

Origin. Nearctic. Canada: ON [new record]. United States: IA, MO, MS, NY, PA, RI.

Diagnosis

Mocyta scopula can be distinguished from bicolored Canadian species and paler specimens of M. fungi by its finely punctate pronotum that is almost as wide as the elytra and ca. as long, and the distinctly transverse antennomeres 6–10 (Fig. 30A). The barcode sequences of M. scopula forms a sister cluster with M. luteola (BOLD:ABW2813), with a sequence divergence of ~ 7.5%. These species can be easily separated using the above diagnosis.

Figure 30. 

Mocyta scopula (Casey) A habitus (hind legs missing on specimen, taken from related species) B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D male tergite VIII E male sternite VIII F spermatheca (female syntype). Scale bars: 1 mm (A); 0.2 mm (B–F).

Bionomics

The Canadian specimen was collected from forest litter with a Berlese funnel but nothing specific is known about this species’ microhabitat preferences.

Comments

Mocyta scopula is a native Nearctic species distributed in eastern North America. Here we newly report it from Canada based on one male specimen collected in southern Ontario. Its distribution in the United States is based on type material, including its putative synonyms, which should be verified.

Mocyta scopula is the type species of Dolosota Casey, which has been treated as a subgenus of Acrotona since Seevers (1978). However, using the generic concepts of Klimaszewski et al. (2015), this species best fits in genus Mocyta based on the following character states: dorsal surface without fine white pubescence; broad tergite VIII with basal line not touching base of tergite; spermatheca with pear-shaped capsule and distinct but small invagination, and thin and irregularly shaped stem ending in a tightly deflexed apex (Fig. 30F) (based on images taken of female syntypes (NMNH). Further evidence comes from barcode sequences of this species, which cluster with the other species of Mocyta. Therefore, we synonymize Dolosota Casey syn. nov. with Mocyta Mulsant and Rey. The other species included in Dolosota by Seevers (1978) were treated as synonyms of M. scopula by Majka and Sikes (2009), in addition to two other Casey names (see above synonymy), following the unpublished results of a type revision by V. Gusarov (Gusarov 2003b). These synonyms and M. scopula are here comb. nov. in Mocyta.

The aedeagus, coloration and punctation of the Canadian specimen are consistent with type material of M. scopula, previously examined and imaged by JK. The two other members of the BIN BOLD:ACH8720 originate from a study by Elven et al. (2010), and were mined into BOLD from GenBank. They were collected in the USA and identified verbatim as Mocyta scopula by V. Gusarov.

The key to Canadian Mocyta in Klimaszewski et al. (2015) can be modified as follows (bicolored species)

2a Pronotum much broader than elytra; antennal articles 5–10 in specimens slightly elongate; spermatheca forming concentric circles posteriorly M. discreta (Casey)
Pronotum ca. as broad as elytra or slightly narrower (Fig. 30A); antennal articles 5–10 subquadrate to transverse (Fig. 30A); spermatheca forming irregular coils posteriorly 2b
2b Pronotum coarsely punctate and extremely transverse with weakly rounded base and apex; antennal articles 5–10 subquadrate; median lobe in lateral view strongly produced ventrad M. luteola (Erichson)
Pronotum finely punctate and transverse, but more rounded at base and apex (Fig. 30A); antennal articles 5–10 distinctly transverse (Fig. 30A); median lobe in lateral view only weakly produced ventrad (Fig. 30B) M. scopula (Casey)

Philhygra angusticauda (Bernhauer, 1909)

Fig. 31A–G

Atheta (Metaxya) angusticauda Bernhauer, 1909

Atheta (Philhygra) pinegensis Muona, 1983, syn. nov.

Material

(DNA barcoded specimens). Canada: Alberta: Jasper National Park, Miette Hotsprings, 53.124, -117.7755, Malaise trap placed in valley with creek bed, sides rocky and mossy, 1439 m, 21.VII.2012, BIObus 2012 (1, CBG). Finland: Lkoc: Muonio, Sarvijärvi, 68.0909, 24.103, 11.VII.2012, M. Pentinsaari (2, ZMUO).

Distribution

Origin. Holarctic. Canada: AB[new record], BC, NB. United States: AK, NH.

Bionomics

As with other species of the genus, P. angusticauda is associated with riparian habitats.

Comments

Philhygra angusticauda is a Holarctic species that was previously recognized in the Palaearctic (Finland, Norway, European Russia, Russian Far East) (Schülke and Smetana 2015; Newton 2019) under the synonym P. pinegensis (Muona). We newly report this species from Alberta and suggest that it broadly occurs across northern Canada. Specimens from the Nearctic and Palaearctic were found to have identical genitalia and their DNA barcodes form a cluster with only 0.3% divergence between Finnish and Canadian specimens.

Figure 31. 

Philhygra angusticauda (Bernhauer) A habitus B median lobe of aedeagus in lateral view C female pygidium D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2018), reproduced with permission.

Philhygra finitima (Casey, 1910)

Fig. 32A–H

Material

(non-sequenced specimens). Canada: Ontario: Algonquin Park, ~45.87, -77.33, car net, 20.VII.2016, T. Struyve (10, CNC, LFC [4 males, 6 females])

Distribution

Origin. Nearctic. Canada: ON [new record]. United States: MA, RI.

Diagnosis

This species can be readily recognized by a combination of its small size, large eyes and relatively simple, ventrally projecting median lobe of the aedeagus in lateral view (Fig. 32B).

Figure 32. 

Philhygra finitima (Casey) A habitus B, C median lobe of aedeagus in lateral view C with internal sac everted (unpublished lectotype) D female pygidium E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H).

Bionomics

Nothing specific is known about this species’ microhabitat preferences but it probably occurs near water as do other species of the genus. The series of Ontario specimens was collected using a car net, which is typically effective for collecting small staphylinids.

Comments

Philhygra finitima is a native Nearctic species distributed in northeastern North America. Here, we newly report it from Canada. Canadian specimens were identified based on comparison with images (Fig. 32C) of the unpublished lectotype of P. finitima in the Casey collection at NMNH.

Philhygra laevicollis (Mäklin, 1852), sensu nov.

Fig. 33A–I

Material

(DNA-barcoded specimens). Canada. Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane forest, 21–27.VI.2012, BIOBus 2012 (1, CBG); same data except 06–11.VIII.2012 (1, CBG). British Columbia: Prince George, Nukko Lake Elementary School, EPQ-CLL-574, 54.083, -122.988, 764 m, 8.V.2015, H. Sapun (1, CBG). United States. Alaska: Dall Island, 54.998, -133.016, 15.VII.2011, D. S. Sikes (1, UAM); Prince of Wales Island, Luck Point, 55.98, -132.772, clear cut, berlese, 9.VIII.2011, J. Stockbridge and B. Wong (1, UAM).

Distribution

Origin. Nearctic. Canada: AB [new record], BC. United States: AK, WA.

Diagnosis

Philhygra laevicollis can be distinguished from most species of the genus by the general shape of the median lobe in lateral view. It is most similar to P. pseudolaevicollis but has a sinuate ventral face of the median lobe in lateral view and large spines in the internal sac (Fig. 33B).

Figure 33. 

Philhygra laevicollis (Mäklin) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in ventral view D, E female pygidium F male tergite VIII G male sternite VIII H female tergite VIII I female sternite VIII. A, B, E–H after Klimaszewski et al. (2020), used with permission C, D after Klimaszewski and Winchester (2002).Scale bars: 1 mm (A); 0.2 mm (B–I).

Bionomics

Specimens have been collected from clear cut areas, transitional zone of a coniferous forest, seepages, and river and creek edges, from moss, leaf litter, gravel, dung, carrion and pitfall traps (Klimaszewski et al. 2020).

Comments

Philhygra laevicollis is a western Nearctic species that was previously considered to include eastern populations that we here treat as Philhygra pseudolaevicollis sp. nov. that differs in male genitalia but also by the divergent DNA barcode sequence.

Neither this species nor P. laevicollis are known from MB, this error was corrected by Klimaszewski et al. (2020). We have observed some variation in the shape of the sclerotized structure present on the female pygidium between specimens collected in BC, but it is not yet clear whether additional species are overlooked within the present concept of P. laevicollis.

Philhygra palustris (Kiesenwetter, 1844)

Fig. 34A–H

Material

(DNA-barcoded specimens). Canada: Ontario: Puslinch, Hanner property, 43.4464, -80.2512, Malaise trap in hardwood forest, 21.VIII.2008, T. Terzin (1, CBG); Puslinch, concession 11/Hume Rd., 43.537, -80.134, Malaise trap in temperate mixed forest, 18–24.IV.2010, P. Hebert (1, CBG); Milverton, Milverton Public School, 43.568, -80.928, Malaise trap, 3.V.2013, J. Van Bakel (1, CBG); Cambridge, rare Charitable Research Reserve, Hogsback forest, 43.3729, -80.354, edge of hardwood forest, intercept trap, 31.V.2015, BIO collections staff (2, CBG); same except pan traps (1, CBG); Kawartha Lakes, 44.366, -78.478, farm, Malaise trap, 13.VI.2015, B. McClenaghan (2, vouchers not preserved); Guelph, Arkell Research Station, 43.5187, -80.1709, between corn and soy fields, w/ nearby pasture, Malaise trap, 8.V.2015, BIO collections staff (1, CBG); same except soy field, 43.5264, -80.1796, 4-headed SLAM trap, 17.V.2017 (1, CBG); Hamilton, Royal Botanical Gardens, Cootes Paradise, 43.281, -79.904, forest, deadwood and UV lights at night, 21.VII.2017, M. Pentinsaari (1, CBG); Markham, 43.9371, -79.2285, mixed habitat, Berlese funnel, 25.VI.2017, Rouge NUP BioBlitz Volunteers (1, CBG). Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 16.VI.2010, F. Koehler (1, ZSM). Estonia: Piusa, 57.844, 27.466, 05.VII.2010, J. Salokannel (2, ZMUO). Finland: Ab: Nauvo, Sandö, 60.1747, 22.1338, 18.VI.2011, M. Pentinsaari (1, ZMUO); Ok: Vaala, Manamansalo, 64.3365, 27.0879, 21.VIII.2011, M. Pentinsaari (1, ZMUO); Ks: Kuusamo, Oulanka, 66.3686, 29.3188, 07.VIII.2011, M. Pentinsaari (1, ZMUO); Ka: Virolahti, Hailiniemi, 60.5259, 27.7366, 20.VII.2012, M. Pentinsaari (1, ZMUO). Germany: Riedlhuette, Diensthuettenstrasse, 48.937, 13.412, 09.VII.2011, F. Koehler and M. Koehler (2, ZSM); Spiegelau, Schwarzachstrasse, 48.9456, 13.3619, 09.VII.2011, F. Koehler and M. Koehler (2, ZSM); Waldhaeuser, Lusen- und Boehmstrasse, 48.93, 13.492, 09.VII.2011, F. Koehler and M. Koehler (2, ZSM); Arnsberg-Breitenbruch, NWZ Hellerberg, 51.4461, 8.13539, 30.V.2011, F. Koehler (2, ZSM); Bornheim-Hemmerich, Hellenmaar, 50.7402, 6.91803, 14.VIII.2012, F. Koehler (1, ZSM); Erftstadt-Bliesheim, NWZ Altwald Ville, 50.7917, 6.84384, 03.VI.2011, F. Koehler (1, ZSM); Kandel, Bienwald, 49.01, 8.103, 05.VI.2010, F. Koehler (1, ZSM); Eisenach, E, Rothenhof, Hoerselufer, 50.9643, 10.3644, 06.VII.2013, GBOL-Team ZFMK (2, ZFMK).

Distribution

Origin. Palaearctic (adventive in North America). Canada: MB, ON [new record]. USA: CT, MA, ME, NH, NY, PA, RI, SC, VT, WI.

Diagnosis

Males of this species are easily recognized among other Canadian Philhygra by the simple, non-projecting median lobe in lateral view (Fig. 34B).

Figure 34. 

Philhygra palustris (Kiesenwetter) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in ventral view D female pygidium E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H).

Bionomics

Most specimens of this species were collected by passive traps in a variety of habitats. In Sweden, P. palustris is considered a eurytopic species that occurs in various types of decaying plant matter, including compost, seaweed and hay piles, and along muddy shores of water bodies (Palm 1970). It can be collected in very large numbers using a car net (V. Assing, pers. comm.).

Comments

Philhygra palustris is a Palaearctic species that has become adventive and widespread in eastern North America. In the Palaearctic, it is very broadly distributed and reported from Europe, North Africa (Morocco), Russia (European and Siberia), Mongolia, North and South Korea (Lee and Ahn 2012), Japan, and northern China (Newton 2019). It is also known from the Azores and the Canaries (Newton 2019), though it is likely introduced there as well.

This species was reported from Canada (Manitoba) for the first time in the checklist by Bousquet et al. (2013) but no specimens could be found in the CNC to support this record. It is likely that the species P. tenuicula (Casey, 1911) described from Manitoba and treated as a synonym of P. palustris (Newton 2019), is the basis of this record. Philhygra palustris was first recorded from North America by Muona (1984) from New York, Maine and Pennsylvania but detailed specimen level data were not provided. Several specimens from various localities in southern Ontario have been sequenced, and their barcodes cluster with European specimens, with multiple haplotypes shared between Canada and Europe. This common European species is here confirmed to be adventive in Canada and is probably broadly distributed in at least eastern North America.

Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari, sp. nov.

Fig. 35A–G

Type material

Holotype (male) (CNC): NEW BRUNSWICK. YORK CO: New Maryland, Charters Settlement, 45.8341°N, 66.7445°W, 22 April 2005, R.P. Webster coll. / mature spruce and cedar forest, seepage area, in saturated sphagnum and leaf litter / HOLOTYPE Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari sp. nov., des. Klimaszewski 2021 [red printed label]. Paratypes (12: LFC, CNC): same data as holotype (1 male, CNC). Canada, New Brunswick, York Co., New Maryland, Charters Settlement, 45.8331°N, 66.7410°W, 14.04.2005, mixed forest in litter and sphagnum, R.P. Webster (1 male, LFC); York Co., New Maryland, Charters Settlement, 45.8390°N, 66.7308°W, 18.04.2005, mixed forest under bark, R.P. Webster (1 male, LFC); York Co., New Maryland, Charters Settlement, 45.8428°N, 66.7279°W, 20.04.2005, mixed forest small sedge marsh in moist grass litter and sphagnum, R.P. Webster (2 females, LFC, 1 female CNC); York Co., New Maryland, Charters Settlement, mixed forest, near small shaded brook, in leaf litter and moss, 9.05.2005, R.P. Webster (1 female, CNC); York Co., Canterbury Trail to Browns Mtn. Fen, 45.8978°N, 67.6273°W, mature cedar forest near stream, sifting leaf litter, 02.05.2005, M. Giguere and R. Webster (1 male, CNC); Northumberland Co., Goodfellow Brook Protected Area, 46.8943°N, 65.3796°W, old growth, wet eastern cedar swamp, in litter and moss on hummocks, near water, 23.05.2007, R.P. Webster (1 female, CNC). Quebec, Scotstown, 28.04.2008, C. Levesque, Barcode sample, BCO1 vial #X16, 26.05.2010, R. Civade (1 male, LFC). Ontario, Nipissing Co., Algonquin Prov. Park near Brent, 19.08.1980, R. Baranowski (1 female, LFC); same except: 21.08.1980 (1 male, LFC).

Non-types

(DNA-barcoded specimens). Canada: New Brunswick: Restigouche Co., 9 km S of Saint Arthur, 47.818, -66.756, eastern white cedar swamp, in moss and litter near small ponds, 14.VI.2006, R.P. Webster (1, cRW).

Etymology

Prefix -pseudo meaning false/not genuine, added to the sibling species name P. laevicollis (Mäklin).

Diagnosis

This species is similar externally and genitally to P. laevicollis but may be distinguished from it by the following combination of characters: body on average narrower, antennomeres 6–7 more elongate (Fig. 35A), ventral margin of tubus of the median lobe of aedeagus straight apically (Fig. 35B) (sinuate in P. laevicollis, Fig. 33B), apical sclerites of internal sac without large spike-like projections (Fig. 35B).

Figure 35. 

Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C female pygidium D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2018), reproduced with permission.

Description

Body narrowly subparallel, moderately flattened, length 3.0–4.2 mm; colour dark brown, elytra dark brownish to brownish yellow, except for darker scutellar area and paler legs, basal antennomeres rust-brown (Fig. 35A); integument moderately glossy, forebody sparsely punctate and pubescent, with pubescence long, punctation fine, microsculpture distinct and consisting of round and slightly convex meshes; head slightly elongate, round, ca. as wide as pronotum, eyes as long as genae in dorsal view, postocular carina strong basally, diffuse apically; antennae slender, at least as long as pronotum and elytra combined, antennomeres 1–3 strongly elongate, 6 and 7 slightly elongate, 8 and 9 slightly elongate or subquadrate, and terminal one as long as two preceding antennomeres combined; pronotum slightly transverse, impressed medially, arcuate laterally and basally, pubescence sparse, hypomeron visible almost for entire length of pronotum; elytra transverse, broader than pronotum; abdomen subparallel. MALE. Tergite VIII slightly transverse, arcuate apically (Fig. 35D); sternite VIII highly elongate, rounded apically and with wide distance between antecostal suture and base of disc (Fig. 35E); median lobe of aedeagus with moderate-sized bulbus, tubus short, ventral margin arcuate basally and straight apically, tubus narrowly triangular at apex in lateral view (Fig. 35B); internal sac sclerites without spike-like projections, complex as illustrated (Fig. 35B); in dorsal view bulbus roughly oval, tubus short, triangular apically. FEMALE. Tergite VIII transverse and truncate apically (Fig. 35F); sternite VIII rounded apically, apex slightly produced, distance between antecostal suture and base of disc wide (Fig. 35G); pygidium as illustrated, with weakly sclerotized central plate, slightly broader than in P. laevicollis (Fig. 35C); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination and with short narrow stem.

Distribution

Origin. Nearctic. Canada: NB, NS, ON, QC.

Bionomics

This species has been recorded from various wetland and riparian habitats in NB: in moss and leaf litter near brook and in litter, grasses, and moss on hummocks in old-growth eastern white cedar swamps and a wet alder swamp, in moist leaves along vernal pond margins in various mixed forests, and a red oak/red maple forest; also from pitfall traps in regenerating red spruce forests (NB) and from vernal pool litter in ON (summarized by Klimaszewski et al. 2018). Collecting period: IV-V, VIII. Collecting method: sifting leaf litter, grasses, and moss, under bark (probably overwintering).

Comments

Although they were not re-examined here, the specimens reported by Majka and Klimaszewski (2008) as P. laevicollis, certainly belong to P. pseudolaevicollis. This species is very similar externally and genitally to P. laevicollis occurring in western North America (AK, AB, BC, WA). Previously, it was tentatively identified as P. laevicollis pending additional study (e.g., Klimaszewski et al. 2005; Klimaszewski et al. 2020). The present evidence from DNA barcodes (8.5% divergence between the eastern and western specimens) and morphology of the aedeagus revealed that eastern and western populations represent two distinct, cryptic species. The single barcoded specimen of P. pseudolaevicollis produced a 407 bp sequence and therefore, no BIN has been generated.

Trichiusa robustula Casey, 1893

Fig. 36A–I

Trichiusa robustula Casey, 1893

Trichiusa immigrata Lohse, 1984, syn. nov.

Material

(DNA-barcoded specimens). Austria: Innervillgraten, Arntal, 46.8362, 12.3348, mountain forest and alpine pastures, car net, 25.VIII.2013, GBOL-Team ZFMK (1, ZFMK). Germany: Nuernberg, N Flughafen, 49.5006, 11.0789, sifting compost, date not provided, GBOL-Team ZFMK (1, ZFMK); Schoenau/Hoersel, W, Gewerbegebiet, 50.947, 10.4214, sifting compost, 25.VIII.2012, GBOL-Team ZFMK (2, ZFMK); Kahlenberg/Eisenach, Pferdekoppel, 50.9469, 10.4287, in horse dung, 16.IX.2013, GBOL-Team ZFMK (1, ZFMK). Finland: Ka: Hamina, Meltti, 60.5798, 27.2016, 12.X.2011, M. Pentinsaari (1, ZMUO); Al: Mariehamn, Dalen, 60.0703, 19.9595, 9.X.2011, M. Pentinsaari (2, ZMUO). Canada: Ontario: Guelph, Division Street, 43.5544, -80.2644, malaise trap, 14.VII.2010, A. Smith (1, CBG); Chelsey, Chelsey District Community School, EQP-CLL-581, 44.3028, -81.0967, 281 m, malaise trap, 22.IX–3.X.2014, A. Grieve (1, CBG); Guelph, Arboretum, Urban Organic Farm, 43.5381, -80.222, compost heaps/mouldy hay pile, 17.IX.2017, M. Pentinsaari (6, CBG).

Additional material

(non-barcoded). Numerous dissected specimens from Denmark were examined in the collection of NHMD.

Distribution

Origin. Nearctic (adventive in Europe). Canada: ON, NB. United States: IA.

Bionomics

In its native range, this species has been collected in a variety of decaying plant matter, especially near water. This species was also common in compost in NB. In Europe, this species has been collected from similar microhabitats including grass clippings and compost (Denton 1998; Anderson and Bryan 2012).

Comments

Trichiusa robustula is a Nearctic species that is broadly distributed in eastern North America but not well collected. It was previously recognized under the synonym T. immigrata Lohse in the West Palaearctic (Europe, Canary Islands, Madeira; Newton 2019), where it is adventive.

When describing his new species, Lohse (1984) noted that it must have originated from North America, since Trichiusa is otherwise endemic to that region. Lohse (1984) stated that T. immigrata was compared with types of North American species described by Casey (1893), but this taxon is a morphological and molecular match to T. robustula. Although most of the specimens collected in North America are bicolored (reddish/dark), study of extensive material from Denmark (NMHD) revealed a grade between fully dark brown to reddish/dark bicolored. Trichiusa robustula is distinctive for the shape of its spermatheca, which bears a rectangular capsule, and stem that has a single 180-degree bend followed by a twisted apical portion (Fig. 36C). The figure of the spermatheca in Klimaszewski et al. (2018) is atypical (Fig. 36C) and the original illustrations in Brunke et al. (2012), reproduced here (Fig. 36D, E), better show these features.

Figure 36. 

Trichiusa robustula Casey A habitus B median lobe of aedeagus in lateral view C–E spermatheca F male tergite VIII G male sternite VIII H female tergite VIII I female sternite VIII. A–C, F–I after Webster et al. (2016) D, E after Brunke et al. 2012. Scale bars: 1 mm (A); 0.2 mm (B–I).

Acknowledgements

We would like to thank the curators listed under Materials and methods for access to the material under their care. Thanks to J. Pedersen (NMHD) and V. Assing (Hanover, Germany) for their valuable input regarding the synonymy of Nearctic and Palaearctic species, and to V. Assing, A. Hansen (NHMD), and A. Bogri (NHMD) for generously sharing habitus and genitalia images for several species. We thank A. Newton (FMNH) for bringing the Neoisoglossa/Isoglossa issue to our attention. We also thank L. Hendrich, S. Schmidt (ZSM), S. Miller (NMNH), D. Sikes (UAM) and M. Seidensticker (MPG Ranch, Montana) for allowing us to publish their unpublished sequences in BOLD. We are grateful to T. Struyve (Belgium) for providing specimens collected by car-netting in eastern Canada. Thanks to C. Bourdon and S. Roberge (LFC) for preparing the vast majority of images included here. Financial support for this study was provided by A-base funding to AJB (Agriculture and Agri-Food Canada: Systematics of Beneficial Arthropods – J-002276), and funding from Natural Resources Canada to JK. All of the barcode sequence analysis and processing of Canadian voucher specimens at CBG, as well as CBG’s collection program, were supported by grants from the Ontario Ministry of Research and Innovation, and from Genome Canada through Ontario Genomics in support of the International Barcode of Life (iBOL) project. Subsequent work to identify the barcoded specimens and analyze data was enabled by the Canada First Research Excellence Fund through its support for the Food From Thought project at the University of Guelph. Two reviewers are thanked for their input, which significantly improved the manuscript.

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