Integrative taxonomy of Nearctic and Palaearctic Aleocharinae: new species, synonymies, and records (Coleoptera, Staphylinidae).

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.

Introduction

Historically, taxonomic research on the hyperdiverse aleocharine rove beetle (: ) 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 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 ‘ 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 , has already resulted in the detection of adventive species of genera , , and in North America (Pentinsaari et al. 2019), and has refined the classification of Holarctic species in (Klimaszewski et al. in press), (Klimaszewski et al. 2019), and (Klimaszewski et al. in press).

Here we broadly compare morphological and molecular data across the Nearctic and West Palaearctic 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:

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

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

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

Laurentian Forestry Centre, Québec, Quebec, Canada;

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

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

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

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

Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany;

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

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

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

We have examined all DNA barcode data for 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 Fleming, 1821

(Lacordaire, 1835)

C1164C98-66CD-56B0-B0BE-3DF38E0703E4

BOLD:ACF6186

Fig. 1A–H

Figure 1.

(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).

Material

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

Distribution.

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

Diagnosis.

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, 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).

Tribe C.G. Thomson, 1859

Subtribe Fenyes, 1918

(Casey, 1906)

FAA43CAB-1D19-5EB0-8D93-E6F624AA22F8

BOLD:ACK6454

Fig. 2A–H

Figure 2.

(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).

Material

(DNA barcoded specimens). Canada: Ontario: Rouge National Urban Park, Toronto Zoo, , 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.

may be easily distinguished from the other Nearctic species of the genus, (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, may be most closely related to Palaearctic (Gravenhorst) as is suggested by barcode clustering. However, can be readily distinguished by the pronotum, which is dark and paler only along the margins, while has broad pale areas laterally. was compared to Palaearctic (Stephens) by Klimaszewski and Ashe (1991; as (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 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 ( (Gyllenhal)) or bank swallows ( (Fairmaire)) (Staniec et al. 2010). The genus is very rarely collected in North America. The Nearctic species (Fenyes) was collected in long series from bank swallow nests (Klimaszewski and Ashe 1991). One specimen of 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 (Brunke and Buffam 2018).

Comments.

The genus and 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).

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

Subtribe C.G. Thomson, 1859

Klimaszewski & Webster, 2016

05D18B83-114C-5324-8226-4A6086310487

BOLD:ACN2725

Fig. 3A–G

Figure 3.

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).

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

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

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

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.

Casey, 1906

5F0DA124-3E2B-55A8-801C-6C32F923749D

BOLD:ABW9176

Fig. 4A–H

Figure 4.

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.

Strand, 1939, syn. nov.

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

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

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.

, 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 . is also a new genus record for the Palaearctic region.Nearctic and Palaearctic 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 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). 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 (Smetana), which was originally described from Italy but later found to be a native Nearctic species (Smetana 1995).

With the above synonymy, the genus is now known only from externally similar sister species (Casey) (Nearctic) and (Erichson) (West Palaearctic), plus Afrotropical (Scheerpeltz). As in the former , is probably misplaced due to superficial similarity. is readily separated from 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 , the head is distinctly narrower than the pronotum.

Casey, 1893

AD374A1E-92D0-5E5E-8511-F4652EB0CEC0

Casey, 1906;

Casey, 1910 (in part);

Klimaszewski & Pelletier, 2004;

In Klimaszewski et al. (2020), was incorrectly attributed to Casey but was actually proposed by Klimaszewski and Pelletier (2004), apparently as an unnecessary replacement name for 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 Newman that preoccupied Casey’s name, so Casey stands as valid with as a synonym.

Klimaszewski, Brunke & Pentinsaari sp. nov.

0A48DBD4-50F3-56F5-813B-2D277A51D955

http://zoobank.org/A8A0402E-2950-4394-B9CF-25E3DD629804

BOLD:ACU5806

Fig. 5A–H

Figure 5.

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).

Type material.

(male): Canada, British Columbia, Prince George, Nukko Lake Elementary EQP-CLL-574, , 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). (3, CBG): Canada, British Columbia, Prince George, Nukko Lake Elementary EQP-CLL-574, , 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 group.

Origin: Nearctic. Canada: BC.

can be easily distinguished from all Nearctic species of the 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).

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).

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

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, keys to genus Casey in Klimaszewski and Pelletier (2004). However, barcode sequences of this species do not cluster with but rather form a cluster with and the species of which bear a spermatheca with broad, circular loops, similar to those of . 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 is 11–12%. It is likely that belongs in a separate genus, but this is outside of the scope of this study. We here place tentatively in 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 group.

(Erichson, 1837)

9525D491-2F0D-5740-B065-B3C850EB878D

Fig. 6A–D

Figure 6.

(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).

Erichson, 1837

Casey, 1906, syn. nov.

:

:

(DNA barcoded specimens). Austria: Innervillgraten, Arntal, , 1580 m, 22.VIII.2010, F. Koehler and J. Koehler (2, ZSM). Canada: Ontario: Northumberland County, Peter’s Woods Protected Natural Area, , 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, , 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, 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).

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

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.

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 (Casey) (Klimaszewski et al. 2018). Assing (2021) recently reported this species from BC and treated 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 (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 as a genus separate from after the discovery that was not a monophyletic group. The type species of was found to be congeneric with that of , and the latter became a junior synonym of the former.

Tribe C.G. Thomson

Revised key to the Canadian genera of

Adapted from Klimaszewski et al. 2018.

(Bernhauer, 1909)

E571F2B1-45E2-5131-951F-93DE5877604D

BOLD:ACF7668

Fig. 7A–H

Figure 7.

(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.

Bernhauer, 1909

Klimaszewski & Webster, 2008, syn. nov.

(DNA barcoded specimens). Canada: Alberta: Waterton Lakes National Park, Highway 6 pulloff, , 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, , 702 m, malaise trap, 22.VIII.2013, C. Simon (1, CBG); New Brunswick: York Co., Fredericton at Saint John River, , margin of river in flood debris, 7.VII.2005, R.P. Webster [note: paratype of ] (1, LFC); Ontario: Guelph, University of Guelph Arboretum, , 12.VI.2019, M. Pentinsaari (1, CBG).

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

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

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

(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).

Lohse, 1974, sensu nov.

In his key to the genera of , Paśnik (2010) distinguished , with its single Palaearctic species (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 Klimaszewski & Webster is a synonym of (see below). A re-examination of other Nearctic species revealed two others that are closely related to : Klimaszewski, and (LeConte). Their morphological divergence from other was represented by an earlier placement in the ‘Nigrella species group’ of by Klimaszewski et al. (2008), together with (now a synonym of ). In corroboration with morphology, DNA barcodes of and form sister clusters (sequences of not available). Transfer of these two species to required a new concept for this genus as most of the distinguishing features were apomorphies of or found not to be of diagnostic value due to variability or overlap with other genera. Here we distinguish members of by their characteristic abdomen (Figs 10–12): 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 ); tergites III–V with very deep basal impressions, each creating strongly convex areas on the disc. Members of could be confused with , 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). was even originally described by LeConte (1863) in , likely based on this similarity.

Figure 10.

(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.

Figure 12.

(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.

Figure 9.

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.

(Klimaszewski, 2020) comb. nov.

07B2AE1C-D445-5110-AA12-4E67C8ECBA3B

Fig. 10A–H

Klimaszewski, 2020

Origin: Nearctic. Canada: BC.

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

We here transfer this species to sensu nov. based on morphology illustrated by Klimaszewski et al. (2020). This recently described western species is most similar to eastern 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 is superficially similar to but these taxa are externally quite different.

(LeConte, 1863) comb. nov.

E1AE07D9-4C82-5A3F-BDBF-05959CE93F6E

BOLD:ACS6831

Fig. 11A–H

Figure 11.

(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.

LeConte, 1863

:

(DNA barcoded specimens). Canada: New Brunswick: York Co., Fredericton at St. John River, , 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).

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

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

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

(Erichson, 1837)

1367A6AB-1432-5D19-BF2F-3ADF13AEBD27

BOLD:ACZ0581

Fig. 12A–H

Erichson, 1837

Klimaszewski & Webster, 2008, syn. nov.

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

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).

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

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).

is newly reported from North America and was previously known in the Nearctic region under the synonym (Klimaszewski et al. 2018). We here newly record this taxon from MB. 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 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.

Klimaszewski, Brunke & Pentinsaari sp. nov.

CDA6F43E-AA37-5C10-9DEA-6ABBD17D3B65

http://zoobank.org/C6DC72D8-A182-43D3-9691-A352F0AF5D0E

BOLD:ADH7347

Fig. 13A–H

Figure 13.

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).

Type material.

(male, CNC): Canada, Ontario, Hartington, Eel Lake, South Frontenac, Paul Hebert’s cottage property, , 6.13.2017, Mikko Pentinsaari, Barcode of Life DNA voucher specimen, Sample ID: BIOUG34206-H01, Process ID: MPCAN465-17. (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.

can be easily distinguished from all Nearctic species of the genus (except eastern (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 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 ).

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).

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

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

It was challenging to place this species in either or 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 but the ligula of is divided to the base, which is considered to be a feature of (Pašnik 2010). The C-shaped spermatheca of also bears some similarity to the group of Klimaszewski et al. (2008) but it is rather different in external morphology. We place this species in pending future systematic research.

Tribe Laporte, 1835

Kraatz, 1862

17544666-193D-5AF3-AE5F-B1095FE817A0

BOLD:AAP9955

Fig. 14A–G

Figure 14.

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).

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

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

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).

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

Kiesenwetter, 1858

F4C78CF8-59A3-5149-AABF-7C1B68144429

BOLD:AAN4271

Fig. 15A–C

Figure 15.

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).

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

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

Among Canadian species of , is extremely similar to 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).

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).

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 from Canada and the United States share the same DNA barcode haplotype, which is also shared by some of the specimens from Germany.

(Gravenhorst, 1806)

EC04853E-57A2-5C81-B6BC-92C243700B56

BOLD:ABW7320

Fig. 16A–G

Figure 16.

(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).

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

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

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

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).

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 .

Heer, 1839

Klimaszewski, Hammond & Langor, 2016

7735FC0D-73F4-51C0-9AA5-7B2FBE84AFE7

Fig. 17A–G

Figure 17.

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).

Material.

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

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

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

, 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.

(Erichson, 1839)

A7476206-A27C-52E7-8349-BDD471A204A8

BOLD:AAO0339

Figs 18A–D , 19A–D , 20A–D.

Figure 18.

A, B (Erichson), and C, D 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).

Figure 19.

Aedeagi of A–C (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 (‘Europe’) C, D sequenced non-types (Finland). Scale bar: 0.2 mm.

Figure 20.

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

Erichson, 1839

Casey, 1893, syn. nov.

:

Erichson, 1839. , male, here designated (ZMHB): Er: [handwritten label] / 5387 [typed label] / Hist.-Coll. (), Nr. 5387, , Erichs., Europa, Zool. Mus. Berlin [typed white label] / Lectotype des. J. Klimaszewski 2019 [white printed label]. (3, ZMHB, without original labels): Hist.-Coll., (), Nr. 5387, Erichs., Europa, Zool. Mus. Berlin; Syntype Erichson, 1837, labelled by MNHUB 2010; Paralectotype 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, 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 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).

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

Casey (1893) gave numerous characters to distinguish from 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 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 (Fig. 20). Therefore, in corroboration with Fenyes (1918), we treat as a synonym of .

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, , 28.IV.2010, F. Koehler (1, ZSM [1 sequenced]). Czech Republic (all : Bohemia, Poděbrady 50 km, Smetana, 1959, car net trap (1); Bohemia, Chvojno, Smetana (1); Moravia, Drnholec, Smetana (1). Denmark (all : 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, , car net, 20.VI.2015, GBOL-Team ZFMK (2, ZFMK [2 sequenced]); Hoenningen bis Insul, Ahrtal, , 24.IV.2010, F. Koehler (1, ZSM [1 sequenced]); Oberheimbach, Franzosenkopf, , 27.V.2012, W. Koehler (1, ZSM [1 sequenced]). Finland (all : N: Espoo, Saunalahti, , 17.IX.2012, fungusy aspen logs, E. Helve (1) [barcoded]; Al: Bjoerkoe, , 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 : Cenkov, Smetana, 1963 (11); Nová Sedlica, Smetana, 1961 (2); Ruská Poruba, Smetana, 1956 (2). United Kingdom (all : Essex (6).

Putative undescribed (corresponding to BIN BOLD:ACA9191):

Finland (all : 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 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]).

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 ) were investigated and found to likely represent an undescribed species of in Europe (confirmed specimens from Finland and Germany). Although most morphological characters of 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: 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 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.

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

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

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

Fenyes, 1921

0EBFFF0B-32F6-5C94-89D4-73616E174085

BOLD:ACK1459

Figs 21A–G , 22A–D

Figure 21.

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.

Figure 22.

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

Fenyes, 1921

Klimaszewski & Webster, 2016, syn. nov.

:

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

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

The aedeagi of the male paratype (holotype in collection of the California Academy of Sciences) of and holotype of are identical and both differ from that of Palaearctic (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 from synonymy with to synonymy with .

Other material

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

Non-barcoded specimens.

Canada: Quebec: Mont St. Bruno Prov Park, , 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).

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

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

is a broadly distributed native Nearctic species, reported from AB east to NS. Here we treat Nearctic records of as (previously treated as Palaearctic , e.g., Klimaszewski et al. 2018) and newly report the genus from ON and NS. 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 from ON and NS formed a barcode cluster that was nearly 5% divergent from those of Palaearctic specimens of (BOLD:AAO1175, one published sequence record from Belgium and three unpublished records from the Netherlands). Northern European , 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 . Based on the study of one paratype of , described from Manitoba and not reported since, it was discovered that Nearctic specimens of correspond to this species and differ from Palaearctic 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 . The illustration in Palm (1968) of the aedeagus of in dorsal view appears to be less angulate than that of but this needs verification. Based on these differences in male genitalia (Fig. 22A–C) and the COI barcodes, is morphologically and genetically distinct from Palaearctic , 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 (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.

Mannerheim, 1830

C933A12D-0B8F-51BE-8FAE-7A344E583D3C

BOLD:ACF7981 [Nearctic]; BOLD:ABW9049 and BOLD:AAO0291 [both Palaearctic]

Fig. 23A–G

Figure 23.

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.

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

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.

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 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.

Seevers, 1951

A955BCE4-D571-59B3-A8DA-CCF2E57F884F

Fig. 24A–H

Figure 24.

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.

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

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

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

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

Seevers, 1951

7FD8B947-6FC3-5A0E-BDDE-4943A6B64484

BOLD:ACY8004

Fig. 25A–G

Figure 25.

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).

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

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

is extremely similar to and 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 but shorter and broader than that of . The emargination of male tergite VIII in appears to be shallower and broader than that of but more specimens are needed to confirm this.

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

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 (female) and 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.

(Gyllenhal, 1810)

8C64540B-A2E8-5F40-A4E4-4F1EF11047F0

BOLD:ADH5714 [Nearctic]; BOLD:AAO0434 [Palaearctic]

Fig. 26A–G

Figure 26.

(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.

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

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

Specimens occur under bark of dead trees.

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 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.

Casey, 1893

CB885C22-DF9B-587F-8888-136D80C98EA5

BOLD:AAO0406

Fig. 27A–G

Figure 27.

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).

Casey, 1893

Dodero, 1922, syn. nov.

Machulka, 1941, syn. nov.

:

:

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

Non-sequenced material.

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

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

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

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 (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 is a ‘recent’ introduction to Europe but from an unknown source. is primarily a New World genus, with ten described species in North and South America (Newton 2019). (Machulka) is currently treated as a synonym of (Newton 2019) and we simply transfer this name to synonymy with . The only Palaearctic species remaining is poorly known (Bernhauer, 1936), known only from the type collected in Great Britain and probably a synonym of . The characters Bernhauer (1936) gave to separate his species from (as ) are slight differences in coloration and body proportions, which are both highly variable in the Palaearctic specimens of studied. Therefore, we consider to be a native Nearctic species that has become adventive in the West Palaearctic and suggest that genus is naturally restricted to the Nearctic and Neotropical regions.

Seevers, 1978

Klimaszewski, Brunke & Pentinsaari sp. nov.

5EBBE4DD-F416-5D20-97E9-CB17191D5A7C

http://zoobank.org/A72E1E63-3D1B-4CC8-882C-6921E5FC3D33

BOLD:AAY6554

Fig. 28A–G

Figure 28.

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).

(male) (CNC): Canada: ON: Waterloo County, Cambridge, Rare Charitable Research Reserve, , soybean field, pitfall trap, 29.VI.2010, A. Brunke [white printed label] / HOLOTYPE Klimaszewski, Brunke & Pentinsaari sp. nov., des A. Brunke 2020 [red printed label]. : Wellington County, Eramosa, , soybean field, pitfall trap, 13.VII.2010, A. Brunke [white printed label] / PARATYPE Klimaszewski, Brunke & Pentinsaari sp. nov., des A. Brunke 2020 [yellow printed label] [CNC, DEBU, 7 specimens]. United States: VA: Arlington County, Marcey Creek, , 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 Klimaszewski, Brunke & Pentinsaari, sp. nov., des A. Brunke 2020 [yellow printed label] [NMNH, 1 specimen].

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

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

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 is most similar to that of Palaearctic 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.

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).

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

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 ‘ sp. 5’ in Brunke et al. (2014), which was collected in both soybean fields and adjacent forest edges by pitfall traps.

is probably broadly distributed in northeastern North America. We have compared the male and female genitalia of with all Central European and Nearctic species of , 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 clustered most closely with (BOLD:ABU6164) in our barcode dataset, its BIN is ~ 8% different from that of the latter. Based on morphology of the aedeagus and spermatheca, is probably even more closely related to East Palaearctic described from Japan and Korea (e.g., Sawada 1970 [as ], Lee and Ahn 2017) rather than to . However, the described species all differ markedly in external morphology.

Casey, 1910

(Kraatz, 1856)

418FBDEF-289E-530A-8EFF-54B3793EA320

BOLD:ACO4408

Fig. 29A–G

Figure 29.

(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).

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

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

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

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.

(Casey, 1893) comb. nov.

9906C34F-CC1D-54D1-B1AA-4848C04A897E

BOLD:ACH8720

Fig. 30A–F

Figure 30.

(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).

Casey, 1893

Casey, 1906

Casey, 1910

Casey, 1910

Casey, 1910

:

Casey, 1910

Casey, 1910

:

:

:

:

:

:

:

:

:

:

:

:

:

:

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

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

can be distinguished from bicolored Canadian species and paler specimens of 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 forms a sister cluster with (BOLD:ABW2813), with a sequence divergence of ~ 7.5%. These species can be easily separated using the above diagnosis.

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

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.

is the type species of Casey, which has been treated as a subgenus of since Seevers (1978). However, using the generic concepts of Klimaszewski et al. (2015), this species best fits in genus 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 . Therefore, we synonymize Casey syn. nov. with Mulsant and Rey. The other species included in by Seevers (1978) were treated as synonyms of 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 are here comb. nov. in .

The aedeagus, coloration and punctation of the Canadian specimen are consistent with type material of , 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 by V. Gusarov.

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

(Bernhauer, 1909)

353C13CE-E3C2-5AD8-B9EC-0A6955E1EA3D

BOLD:ACG2845

Fig. 31A–G

Figure 31.

(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.

Bernhauer, 1909

Muona, 1983, syn. nov.

(DNA barcoded specimens). Canada: Alberta: Jasper National Park, Miette Hotsprings, , 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, , 11.VII.2012, M. Pentinsaari (2, ZMUO).

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

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

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 (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.

(Casey, 1910)

9054FA71-A57B-50B4-A7F4-0F4AB4589FED

Fig. 32A–H

Figure 32.

(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).

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

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

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).

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.

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 in the Casey collection at NMNH.

(Mäklin, 1852), sensu nov.

7178546D-E5F2-598E-BE22-1A01EA8C11B9

BOLD:ACU6301

Fig. 33A–I

Figure 33.

(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).

(DNA-barcoded specimens). Canada. Alberta: Waterton Lakes National Park, Highway 6 pulloff, , 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, , 764 m, 8.V.2015, H. Sapun (1, CBG). United States. Alaska: Dall Island, , 15.VII.2011, D. S. Sikes (1, UAM); Prince of Wales Island, Luck Point, , clear cut, berlese, 9.VIII.2011, J. Stockbridge and B. Wong (1, UAM).

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

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 but has a sinuate ventral face of the median lobe in lateral view and large spines in the internal sac (Fig. 33B).

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).

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

Neither this species nor 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 .

(Kiesenwetter, 1844)

17854478-C200-53E7-B229-61B2137A9C2E

BOLD:AAN6150

Fig. 34A–H

Figure 34.

(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).

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

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

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

Most specimens of this species were collected by passive traps in a variety of habitats. In Sweden, 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.).

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 (Casey, 1911) described from Manitoba and treated as a synonym of (Newton 2019), is the basis of this record. 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.

Klimaszewski, Brunke & Pentinsaari sp. nov.

3C47DBC7-7186-5657-8D36-0DA42F3B1270

http://zoobank.org/7D087111-AC74-4D0C-851D-801BA3206497

Fig. 35A–G

Figure 35.

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.

: NEW BRUNSWICK. YORK CO: New Maryland, Charters Settlement, , 22 April 2005, R.P. Webster coll. / mature spruce and cedar forest, seepage area, in saturated sphagnum and leaf litter / HOLOTYPE Klimaszewski, Brunke & Pentinsaari sp. nov., des. Klimaszewski 2021 [red printed label]. : same data as holotype (1 male, CNC). Canada, New Brunswick, York Co., New Maryland, Charters Settlement, , 14.04.2005, mixed forest in litter and sphagnum, R.P. Webster (1 male, LFC); York Co., New Maryland, Charters Settlement, , 18.04.2005, mixed forest under bark, R.P. Webster (1 male, LFC); York Co., New Maryland, Charters Settlement, , 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, , 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, , 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, , eastern white cedar swamp, in moss and litter near small ponds, 14.VI.2006, R.P. Webster (1, cRW).

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

This species is similar externally and genitally to 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 , Fig. 33B), apical sclerites of internal sac without large spike-like projections (Fig. 35B).

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 (Fig. 35C); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination and with short narrow stem.

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

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).

Although they were not re-examined here, the specimens reported by Majka and Klimaszewski (2008) as , certainly belong to This species is very similar externally and genitally to occurring in western North America (AK, AB, BC, WA). Previously, it was tentatively identified as 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 produced a 407 bp sequence and therefore, no BIN has been generated.

Casey, 1893

3DA98EAA-0B4A-5E7E-80FB-E1B0A447C7B9

BOLD:AAY6555

Fig. 36A–I

Figure 36.

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).

Casey, 1893

Lohse, 1984, syn. nov.

(DNA-barcoded specimens). Austria: Innervillgraten, Arntal, , mountain forest and alpine pastures, car net, 25.VIII.2013, GBOL-Team ZFMK (1, ZFMK). Germany: Nuernberg, N Flughafen, , sifting compost, date not provided, GBOL-Team ZFMK (1, ZFMK); Schoenau/Hoersel, W, Gewerbegebiet, , sifting compost, 25.VIII.2012, GBOL-Team ZFMK (2, ZFMK); Kahlenberg/Eisenach, Pferdekoppel, , in horse dung, 16.IX.2013, GBOL-Team ZFMK (1, ZFMK). Finland: Ka: Hamina, Meltti, , 12.X.2011, M. Pentinsaari (1, ZMUO); Al: Mariehamn, Dalen, , 9.X.2011, M. Pentinsaari (2, ZMUO). Canada: Ontario: Guelph, Division Street, , malaise trap, 14.VII.2010, A. Smith (1, CBG); Chelsey, Chelsey District Community School, EQP-CLL-581, , 281 m, malaise trap, 22.IX–3.X.2014, A. Grieve (1, CBG); Guelph, Arboretum, Urban Organic Farm, , 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.

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

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).

is a Nearctic species that is broadly distributed in eastern North America but not well collected. It was previously recognized under the synonym 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 is otherwise endemic to that region. Lohse (1984) stated that was compared with types of North American species described by Casey (1893), but this taxon is a morphological and molecular match to . 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. 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.

1A	Antennomere 10 only weakly transverse (Fig. 1A); abdominal tergite VI with distinct basal impression in addition to coarse punctures	Amarocharaforticornis (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

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

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 9–13); impressions of abdominal tergites with at least a few coarse punctures and glossy areas, punctation distinctly different from that of disc (Figs 9–13)	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 10–13); tergite III ca. twice as wide as long or wider; tergal impressions never with median carina (Figs 10–13)	4
4	Abdomen at base elongate and moderately constricted, ca. as wide as head (Figs 10–12); tergite III, at most, twice as wide as long; tergal impressions deep and sharply delineated from strongly convex disc (Figs 10–12)	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

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)