Integrative taxonomy of Nearctic and Palaearctic Aleocharinae: new species, synonymies, and records (Coleoptera, Staphylinidae) Adam J. Brunke1, Mikko Pentinsaari2, Jan Klimaszewski3 1 Agriculture and Agri-Food Canada, Canadian National Collection of Insects, Arachnids and Nematodes, 960 Carling Avenue, Ottawa, Ontario, K1A 0C6, Canada 2 Centre for Biodiversity Genomics, 50 Stone Road East, University of Guelph, Guelph, Ontario, N1G 2W1, Canada 3 Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, 1055 du PEPS, PO Box 10380, Stn. Sainte-Foy, Québec, QC, G1V 4C7, Canada Corresponding author: Adam J. Brunke (adam.brunke@canada.ca) Academic editor: Volker Assing  |  Received 19 February 2021  |  Accepted 29 March 2021  |  Published 31 May 2021 http://zoobank.org/EEE8490B-B41D-4A6C-A963-234C256C99BF Citation: Brunke AJ, Pentinsaari M, Klimaszewski J (2021) Integrative taxonomy of Nearctic and Palaearctic Aleocharinae: new species, synonymies, and records (Coleoptera, Staphylinidae). ZooKeys 1041: 27–99. https://doi. org/10.3897/zookeys.1041.64460 Abstract A long tradition of separate Nearctic and Palaearctic taxonomic studies of the diverse aleocharine rove beetles (Coleoptera: Staphylinidae) has obscured the recognition of Holarctic species and detection of adventive species in both regions. Recently, integrated study of the two regions through detailed morphological comparisons and development of an authoritatively identified DNA barcode reference library has revealed the degree to which these two aleocharine faunas are interconnected, both naturally and through human activity. Here this approach is adopted to recognize new species, reveal Holarctic species, and recognize adventive species in both North America and Europe. The following new species are described: Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov. from British Columbia; Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov., from Ontario, Maryland and North Carolina; Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov., from Ontario and Virginia; and Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. from eastern Canada. Dasygnypeta velata and Philhygra angusticauda are revealed to be Holarctic species, resulting in the following synonymies: Dasygnypeta velata (Erichson, 1839) = Gnypeta minuta Klimaszewski & Webster, 2008, syn. nov. and Philhygra angusticauda (Bernhauer, 1909) = Atheta (Philhygra) pinegensis Muona, 1983, syn. nov. The Nearctic species Hylota ochracea (and genus Hylota), Thecturota tenuissima, and Trichiusa robustula are newly reported from the Palaearctic region as adventive, resulting in the following synonymies: Hylota ochracea ZooKeys 1041: 27–99 (2021) doi: 10.3897/zookeys.1041.64460 https://zookeys.pensoft.net Copyright Adam J. Brunke et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE Launched to accelerate biodiversity research A peer-reviewed open-access journal mailto:adam.brunke@canada.ca http://zoobank.org/EEE8490B-B41D-4A6C-A963-234C256C99BF https://doi.org/10.3897/zookeys.1041.64460 https://doi.org/10.3897/zookeys.1041.64460 https://zookeys.pensoft.net http://creativecommons.org/licenses/by/4.0/ http://creativecommons.org/licenses/by/4.0/ Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)28 Casey, 1906 = Stichoglossa (Dexiogyia) forticornis Strand, 1939, syn. nov.; Thecturota tenuissima Casey, 1893 = Atheta marchii Dodero, 1922, syn. nov.; and Trichiusa robustula Casey, 1893 = T. immigrata Lohse, 1984, syn. nov. The Palaearctic species Amarochara forticornis, Anomognathus cuspidatus, Oligota pumilio, and Parocyusa rubicunda are newly confirmed from the Nearctic region as adventive, resulting in the following synonymies: Parocyusa rubicunda (Erichson, 1837) = Chilopora americana Casey, 1906, syn. nov. and Anomognathus cuspidatus (Erichson, 1839) = Thectura americana Casey, 1893, syn. nov. The genus Dasygnypeta, sensu nov. is newly reported from North America, Paradilacra is newly reported from eastern North America, and Haploglossa is newly reported from Canada, resulting in the following synonymy: Paradilacra densissima (Bernhauer, 1909) = Gnypeta saccharina Klimaszewski & Webster, 2008, syn. nov. Native Cyphea wallisi is newly reported from across Canada and C. curtula is removed from the Nearctic fauna. The status of both Gyrophaena affinis and Homalota plana is uncertain but these species are no longer considered to be adventive in North America. Three new combinations are proposed: Dasygnypeta baranowskii (Klimaszewski, 2020) and D. nigrella (LeConte, 1863) (both from Gnypeta) and Mocyta scopula (Casey, 1893) (from Acrotona). Dolosota Casey, 1910, syn. nov. (type species Eurypronota scopula Casey), currently a subgenus of Acrotona, is therefore synonymized with Mocyta Mulsant & Rey, 1874. Additionally, four new Canadian records and 18 new provincial and state records are reported. Keywords Canada, DNA barcodes, faunistics, morphology, North America, rove beetles, United States Introduction Historically, taxonomic research on the hyperdiverse aleocharine rove beetle (Coleop- tera: Staphylinidae) faunas of North America and better-known Europe has been con- ducted separately, with a few exceptions (e.g., Klimaszewski et al. 1979). More recent- ly, a closer examination of Aleocharinae in these two regions has demonstrated that a number of species are shared between the Nearctic and Palaearctic, either naturally (Holarctic) or through human activity (adventive) (e.g., Muona 1984; Klimaszewski et al. 2007; Klimaszewski et al. in press). The interconnectedness of these assemblages, combined with the sheer diversity of the subfamily, have made it difficult to avoid describing synonyms of taxa from other regions, especially when those taxa have been described in entirely different genera (e.g., Gusarov 2003a). One strategy to broadly address this challenge is the publication of detailed illustrations of habitus and geni- talia in comprehensive faunal treatments such as the recently available ‘Aleocharinae of Eastern Canada’ (Klimaszewski et al. 2018) and ‘the Danish Beetle Bank’ website (Hansen et al. 2017), the latter an online resource for the Danish beetle fauna. In the past few years, resources such as these have made it possible to efficiently cross-check Nearctic and Palaearctic aleocharines without consulting a comprehensive reference collection for each region. In combination with careful morphological study, large-scale DNA barcoding (e.g., deWaard et al. 2019) has accelerated the discovery of Holarctic species, and the detec- tion of new adventive species and potential synonyms in the Canadian beetle fauna (e.g., Pentinsaari et al. 2019) by algorithmically flagging potential taxonomic issues Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 29 and novelties, and connecting authoritatively identified specimens to unidentifiable females, damaged specimens, or other life stages. This integrated taxonomic approach, as applied to Aleocharinae, has already resulted in the detection of adventive species of genera Amischa, Atheta, and Myllaena in North America (Pentinsaari et al. 2019), and has refined the classification of Holarctic species in Atheta (Klimaszewski et al. in press), Boreophilia (Klimaszewski et al. 2019), and Gnathusa (Klimaszewski et al. in press). Here we broadly compare morphological and molecular data across the Nearctic and West Palaearctic Aleocharinae in order to better integrate the taxonomic knowl- edge 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 subse- quently 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 ste- reoscopic microscope; Nikon Digital Camera DXM 1200F) and processed in Adobe Photoshop. Terminology mainly follows that used by Lohse et al. (1990) and Kli- maszewski et al. (2018). The ventral part of the median lobe of the aedeagus is consid- ered to be the part of the bulbus containing the foramen mediale, the entrance of the ductus ejaculatorius, and the adjacent venter (ventral part of the tubus of the median lobe) of the tubus; the opposite side is referred to as the dorsal part. Depository abbreviations: CBG Centre for Biodiversity Genomics, University of Guelph, Guelph, Ontario, Canada; CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ot- tawa, Ontario, Canada; cRW Personal collection of Reginald P. Webster, Charters Settlement, New Brun- swick, Canada (also known as RWC); LFC Laurentian Forestry Centre, Québec, Quebec, Canada; MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Mas- sachusetts, United States (C. Maier); NHMD Natural History Museum of Denmark, Copenhagen University, Copenha- gen, Denmark (A. Solodovnikov); NMNH National Museum of Natural History, Washington D.C., United States (F. Shockley); UAM University of Alaska Museum Insect Collection, Fairbanks, Alaska, United States (D. Sikes); Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)30 ZFMK Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany; ZMHB Museum für Naturkunde, Berlin, Germany (B. Jaeger); ZMUO Zoology Museum, University of Oulu, Oulu, Finland (M. Mutanen); ZSM Zoologische Staatssammlung, Munich, Germany (F. Koehler). We have examined all DNA barcode data for Aleocharinae previously generated by a variety of projects in both Europe and North America (e.g., Rulik et al. 2017; Sikes et al. 2017; McClenaghan et al. 2019; and other studies summarized by Pentinsaari et al. 2019). Fifty-three barcode sequences, the majority of which are Canadian sequence records originating from various projects coordinated by CBG, are published here for the first time. All sequences were analyzed using the workbench tools of the BOLD platform (http://www.boldsystems.org) after applying filters to exclude those flagged as misidentifications, those with sequence lengths under 100 bp, those with stop codons, and those flagged as contaminated. Sequences were generally visualized as clusters in neighbour-joining trees (using the Taxon ID Tree tool). In addition, BIN Discordance Reports, which compare the taxonomy of the specimen records to their BIN assign- ments, 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 Gen- Bank (accessions provided in Suppl. material 1: Table S1). Taxonomic accounts Tribe Aleocharini Fleming, 1821 Amarochara forticornis (Lacordaire, 1835) BOLD:ACF6186 Fig. 1A–H Material (DNA barcoded specimens). Canada: Ontario: Fergus, Centre Welling- ton District High School, 43.704, -80.358, Malaise trap, 3.V.2013, M. Cottrill (1, CBG); Guelph, Biodiversity Institute of Ontario, 43.528, -80.229, Malaise trap, 25.VII.2013, BIO Collections Staff (1, CBG); Rouge National Urban Park, west of http://www.boldsystems.org https://doi.org/10.5883/DS-ALEO2020 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACF6186 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 31 Figure 1. Amarochara forticornis (Lacordaire) A habitus B median lobe of aedeagus in lateral view C me- dian 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). Glen Rouge campground, 43.804, -79.146, marsh scrub along riverside, pitfall trap, 9.VI.2013, BIObus 2013 (1, CBG); Cambridge, rare Charitable Research Reserve, Preston Flats, 43.3908, -80.3747, grassy wetland, pitfall trap, 31.V.2015, BIO Col- lections staff (2, CBG); Peterborough, 44.318, -78.372, farm, malaise trap, B. Mc- Clenaghan (1, CBG). Distribution. Origin: Palaearctic (adventive in Nearctic). Canada: ON [new record]. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)32 Diagnosis. Amarochara forticornis may be easily recognized among the other Ca- nadian species of the genus by the distal antennomeres, which are less than twice as wide as long. The species is also unique within the genus by having a distinct basal impression on abdominal tergite VI. Bionomics. In its native range, A. forticornis occurs in a variety of open and forested habitats, including forests, edges of waterways, grasslands, agricultural fields, and gar- dens (Assing 2002). It has been mostly collected by pitfall traps in the spring and sum- mer, 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, subter- ranean 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). The key to Eastern Canadian Amarochara in Klimaszewski et al. (2018) should be modified as follows 1A Antennomere 10 only weakly transverse (Fig. 1A); abdominal tergite VI with distinct basal impression in addition to coarse punctures .............................. ..........................................................Amarochara forticornis (Lacordaire) – Antennomere 10 strongly transverse, at least twice as wide as long (native spe- cies); abdominal tergite VI with, at most, coarse punctures at base ........... 1B 1B Pronotum with strong microsculpture and coarse, dense punctation, surface almost matte .......................................................................A. duryi (Casey) – Pronotum without microsculpture or with fine microsculpture, and with fine sparse to moderately dense punctation, surface moderately to highly glossy....2 Tribe Oxypodini C.G. Thomson, 1859 Subtribe Microglottina Fenyes, 1918 Haploglossa nebulosa (Casey, 1906) BOLD:ACK6454 Fig. 2A–H Material (DNA barcoded specimens). Canada: Ontario: Rouge National Urban Park, Toronto Zoo, 43.8223, -79.1897, forest, malaise trap, 21.V.2013, L. Attard and K. Greenham (1, CBG). Distribution. Origin: Nearctic. Canada: ON [new record]. United States: OK, PA. Diagnosis. Haploglossa nebulosa may be easily distinguished from the other Nearc- tic species of the genus, H. barberi (Fenyes), by the bicolored elytra and fusiform body (Klimaszewski and Ashe 1991). Based on the shape of the spermatheca with its narrow capsule and broadly rounded apex, H. nebulosa may be most closely related to Pal- aearctic H. marginalis (Gravenhorst) as is suggested by barcode clustering. However, http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACK6454 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 33 Figure 2. Haploglossa nebulosa (Casey) A habitus B median lobe of aedeagus in lateral view (adapted from Klimaszewski and Ashe (1991)) C median lobe of aedeagus in dorsal view (adapted from Klimasze- wski 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). H. nebulosa can be readily distinguished by the pronotum, which is dark and paler only along the margins, while H. marginalis has broad pale areas laterally. Haploglossa nebulosa was compared to Palaearctic H. villosula (Stephens) by Klimaszewski and Ashe (1991; as H. pulla (Gyllenhal)), but the species is quite different externally (much Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)34 darker, finer pronotal punctation) and the spermatheca of the latter species is of the type with a large, rounded capsule. Bionomics. All members of Haploglossa are nidicolous, mostly in bird nests but also in mammal and ant nests (summarized by Staniec et al. 2010). Some species with well-known life histories appear to specialize on particular types of host nests, such as birds of prey (H. picipennis (Gyllenhal)) or bank swallows (H. nidicola (Fairmaire)) (Staniec et al. 2010). The genus is very rarely collected in North America. The Nearctic species H. barberi (Fenyes) was collected in long series from bank swallow nests (Kli- maszewski and Ashe 1991). One specimen of H. nebulosa has been found in a rodent nest within a hollow tree (Klimaszewski and Ashe 1991) but bird and mammal nests have been poorly sampled in the Nearctic and more collecting is needed to determine the biology of the Nearctic Haploglossa (Brunke and Buffam 2018). Comments. The genus Haploglossa and H. nebulosa are newly reported from Can- ada, from a single locality in southern Ontario. The species is also known from Okla- homa and Pennsylvania, United States (Klimaszewski and Ashe 1991). The key to genera of Oxypodini in Eastern Canada in Klimaszewski et al. (2018) should be modified as follows 8A Pronotum strongly converging anteriad; posterolateral margin of elytra with strong sinuate emargination ..................................................................... 8B – Pronotum not or, at most, weakly converging anteriad ...............................9 8B Pronotum with fine punctures, not clearly visible at moderate magnification, shape strongly transverse, ~ 1.5 × wider than long .......Crataraea Thomson – Pronotum with coarse punctures, clearly visible with low magnification, shape weakly transverse, no more than 1.4 × wider than long ....................... ......................................................................................Haploglossa Kraatz Subtribe Oxypodina C.G. Thomson, 1859 Hylota cryptica Klimaszewski & Webster, 2016 BOLD:ACN2725 Fig. 3A–G Material (DNA barcoded specimens). Canada: Ontario: Guelph, Hanlon Preserva- tion Park, 43.506, -80.213, mixed forest, dead wood and beating, 11.VI.2017, M. Pentinsaari (1, CBG); Hartington, Eel Lake Cottage, 44.5628, -76.553, Lindgren funnel, 12.VII.2017, G. Blagoev (1, CBG); Kawartha Lakes, 44.28, -78.529, farm, malaise trap, 5.V.2016, B. McClenaghan (1, CBG); Murphy’s Point Provincial Park, 44.7812, -76.2336, forest, malaise trap, 19.VI.2014, CBG Collections staff (1, CBG); Newfoundland: Terra Nova National Park, Blue Hill Road, 48.598, -53.9702, malaise trap, old balsam fir forest, 2.VII.2013, E. Perry (1, CBG). Distribution. Origin: Nearctic. Canada: AB, NB, NF [new record], ON [new record]. http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACN2725 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 35 Figure 3. Hylota cryptica Klimaszewski & Webster A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Webster et al. (2016). Bionomics. Little is known about the microhabitat preferences of this species, but it likely occurs in in nests or cavities within trees as does H. ochracea (Klimaszewski et al. 2018). Comments. This recently described species, previously known from New Brunswick and Alberta (Klimaszewski et al. 2018) is newly recorded from Ontario and Newfound- land. It is likely to be widely distributed in North America east of the Rocky Mountains. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)36 Hylota ochracea Casey, 1906 BOLD:ABW9176 Fig. 4A–H Stichoglossa (Dexiogyia) forticornis Strand, 1939, syn. nov. Material (DNA barcoded specimens). Canada: Ontario: Guelph, Dovercliffe Road, 43.51, -80.254, backyard, compost and mouldy hay piles, 6.VI.2018, M. Pentinsaari (3, CBG); Guelph, Hanlon Preservation Park, 43.51, -80.221, mixed forest, at UV light, 30.VI.2018, M. Pentinsaari (1, CBG); Whitby, Julie Payette Public School, Ma- laise trap, 43.886, -78.934, 22.IV.-03.V.2013, Z. Turner (1, CBG). Quebec: Montreal, Montreal Botanical Garden, 45.559, -73.566, Malaise trap, 24.VII-02.VIII.2014, M. Larrivee (1, CBG). Finland: Al: Lemland, Äspholm, 60.0675, 19.9583, 9.X.2011, M. Pentinsaari (1, ZMUO); Al: Lemland, Nåtö, 60.046, 19.981, 26.VI.2014, M. Pentin- saari (2, ZMUO); N: Sipoo, Sipoonkorpi, 60.304, 25.202, window trap, 2.VIII.2013, S. Karjalainen and P. Martikainen (1, ZMUO). Distribution. Origin: Nearctic (adventive in Europe). Canada: NB, NS, NT, ON, QC, SK. United States: NY, VT. Bionomics. Hylota ochracea is strongly associated with bird nests in forested habi- tats. It has also been collected from artificial analogs such as a pigeon coup, manmade nest boxes, and a plastic composter bin containing carrion and decaying vegetables (Klimaszewski et al. 2018). The specimens recently collected in Ontario, Canada were found in compost and at UV light. Comments. Hylota ochracea, a widespread Nearctic species (Klimaszewski et al. 2018), is newly reported from the Palaearctic region and had been previous- ly known from Finland, Denmark, Germany, Norway, Sweden, and Switzerland (Lundberg 2006; Schülke and Smetana 2015; Newton 2019) under the synonym Dexiogyia forticornis. Hylota is also a new genus record for the Palaearctic region. Nearctic Hylota ochracea and Palaearctic D. forticornis share a BIN and do not form separate clusters. One of the DNA barcode haplotypes is shared between Finnish and Canadian specimens. Nearctic and Palaearctic populations also have identical male and female genitalia. Based on its specialization on microhabitats in forests, we do not consider H. ochracea to be a naturally occurring Holarctic species. Holarctic beetles are generally those that occur north of the treeline and have crossed treeless Beringia in the last 2.8 Mya (reviewed in Brunke et al. 2020). Hylota ochracea may have been introduced to the Palaearctic region with the nest material of poultry or domestic pigeons, or with another form of decaying plant matter. A similar situation has occurred with the bird nest-associated staphylinid Bisnius palmi (Smetana), which was originally described from Italy but later found to be a native Nearctic species (Smetana 1995). With the above synonymy, the genus Dexiogyia is now known only from exter- nally similar sister species D. angustiventris (Casey) (Nearctic) and D. corticina (Erich- son) (West Palaearctic), plus Afrotropical D. congoensis (Scheerpeltz). As in the former D. forticornis, D. congoensis is probably misplaced due to superficial similarity. Hylota http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ABW9176 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 37 Figure 4. Hylota ochracea Casey A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H  female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), reproduced with permission. is readily separated from Dexiogyia by the shape of the pronotum, which is strongly convergent anteriad, such that its apical width is subequal to the width of the head. In Dexiogyia, the head is distinctly narrower than the pronotum. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)38 Isoglossa Casey, 1893 Rheobioma Casey, 1906; Klimaszewski and Pelletier (2004), syn. of Neoisoglossa Athetalia Casey, 1910 (in part); Klimaszewski and Pelletier (2004) syn. of Neoisoglossa Neoisoglossa Klimaszewski & Pelletier, 2004; Gouix and Klimaszewski (2007), syn. of Isoglossa, unnecessary replacement name; Klimaszewski et al. (2020) as valid genus, incorrectly attributed to Casey (1893). Comments. In Klimaszewski et al. (2020), Neoisoglossa was incorrectly attributed to Casey but was actually proposed by Klimaszewski and Pelletier (2004), apparently as an unnecessary replacement name for Isoglossa Casey 1893. The previous treatment of these generic names and two other synonyms in the catalog of Gouix and Klimasze- wski (2007) is correct and followed here. Blackwelder (1952) was wrong and there is no Isoglossa Newman that preoccupied Casey’s name, so Isoglossa Casey stands as valid with Neoisoglossa as a synonym. Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov. http://zoobank.org/A8A0402E-2950-4394-B9CF-25E3DD629804 BOLD:ACU5806 Fig. 5A–H Type material. Holotype. (male): Canada, British Columbia, Prince George, Nukko Lake Elementary EQP-CLL-574, 54.0831°N, 122.988°W, 764 m asl, Holly Sapun 04/20/2015 to 05/08/2015, Barcode of life, DNA voucher specimen, Sample ID: BIOUG22036-B02, Process ID: SMTPM2682-15 (CNC). Paratypes (3, CBG): Canada, British Columbia, Prince George, Nukko Lake Elementary EQP-CLL-574, 54.0831°N, 122.988°W, 764 m asl, Holly Sapun 04/20/2015 to 05/08/2015, Barcode of life, DNA voucher specimen, Sample ID: BIOUG22036-B07, Process ID: SMT- PM2682-15 (1 male, CBG); same label data except: Sample ID: BIOUG22035-H08, Process ID: SMTPM2665-15 (1 female, CBG); Sample ID: BIOUG22036-A04, Pro- cess ID: SMTPM2672-15 (1 female, CBG). Etymology. The species epithet refers to the remarkably separated triangular apex of the median lobe of the aedeagus, distinguishing it from all other members of the Ocalea group. Distribution. Origin: Nearctic. Canada: BC. Diagnosis. Isoglossa triangularis can be easily distinguished from all Nearctic spe- cies of the Ocalea group of genera by a combination of the strongly transverse and sparsely punctate pronotum, transverse antennomere 4, distinct triangular apex of the median lobe in lateral view (Fig. 5B), and distinct and simple ‘walking cane’ shape of the spermatheca (Fig. 5D). 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 http://zoobank.org/A8A0402E-2950-4394-B9CF-25E3DD629804 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACU5806 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 39 Figure 5. Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). 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 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)40 directed laterad from midline of disc forming arcuate lines on both sides; elytra trans- verse (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); me- dian lobe of aedeagus in lateral view with narrowly elongate crista apicalis at base of bulbus, tubus moderately long, strongly produced ventrally, apex narrowly triangu- lar constricted baso-dorsally in lateral view (Fig. 5B), internal sac structures not pro- nounced (Fig. 5B, C). FEMALE. Tergite VIII truncate apico-medially (Fig. 5G); ster- nite VIII arcuate apically (Fig. 5H); spermatheca with capsule approximately spherical with short neck, stem narrow, long and straight (Fig. 5D). Bionomics. The specimens were collected in a Malaise trap on an open field sur- rounded by mixed forest. Comments. Based on a combination of small size (< 4.5 mm), superficial, meshed microsculpture, sparse pronotal punctation, with punctures separated by more than two puncture diameters, pronotum transverse, shorter and narrower than elytra, and the transverse antennomeres 5–10, I. triangularis keys to genus Isoglossa Casey in Klimaszewski and Pelletier (2004). However, barcode sequences of this species do not cluster with Isoglossa agnita but rather form a cluster with Gennadota canadensis and the species of Neothetalia which bear a spermatheca with broad, circular loops, similar to those of Gennadota. Isoglossa triangularis has a sim- ple spermatheca with a long straight stem and is not externally similar to these taxa (see above), and the barcode divergence between these species and I. triangularis is 11–12%. It is likely that I. triangularis belongs in a separate genus, but this is outside of the scope of this study. We here place I. triangularis tentatively in Isoglossa as not to disturb the existing morphological diagnoses of the genera and identification keys (e.g., Klimaszewski and Pelletier 2004; Klimaszewski et al. 2020), pending generic revision of the Ocalea group. Parocyusa rubicunda (Erichson, 1837) Fig. 6A–D Tachyusa rubicunda Erichson, 1837 Chilopora americana Casey, 1906, syn. nov. Tetralaucopora americana: Klimaszewski et al. 2018 (as valid species) Parocyusa americana: Assing 2021 (possible synonym of P. rubicunda) Material (DNA barcoded specimens). Austria: Innervillgraten, Arntal, 46.8362, 12.3348, 1580 m, 22.VIII.2010, F. Koehler and J. Koehler (2, ZSM). Canada: Ontar- io: Northumberland County, Peter’s Woods Protected Natural Area, 44.124, -78.039, under rock in streambed,12.IX.2011, A. Brunke and S. Paiero (1, DEBU); Crieff Bog, 3 km W Puslinch, sedge meadow, 26.VI.1987, D. Blades (1, DEBU). United States: Connecticut: East Hartford, Two Rivers Magnet Middle School, 41.757, -72.655, 4.VI.2005, J. DeWaard (1, CBG). Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 41 Figure 6. Parocyusa rubicunda (Erichson) A habitus B spermatheca C female tergite VIII D female ster- nite VIII. Scale bars: 1 mm (A); 0.2 mm (B–D). Illustrations after Klimaszewski et al. (2016c). Additional non-barcoded material. Canada: Ontario: Ancaster, 21.X.1967 (1, CNC); Rondeau Prov. Pk., Tulip Tree Trail, Carex and moss on logs in pond, 5.VI.1985, A. Davies and J.M. Campbell (1, CNC); Quebec: Montreal, 20.IX.1969, E.J. Kiteley, 1 (CNC); Mt. Orford Park, 20.IX.-11.X.1972, Dondale and Redner, 1 (CNC). Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)42 Distribution. Origin: West Palaearctic (adventive in North America). Canada: BC, ON, QC, NB, NF. United States: CT, NY, PA. Bionomics. In North America, most specimens of this species have been collected from near water, including a sandy creek bank, in a dried streambed and in moss near the splash zone of a waterfall (Klimaszewski et al. 2018). Nearctic populations of this species are only known from female specimens and the species may be parthenogenetic in North America. In its native distribution, the northern and northwestern popu- lations are also parthenogenetic (Assing 2021) and most likely represent the source population for the Nearctic introduction. Comments. Parocyusa rubicunda is a widespread West Palaearctic species (Europe, European Russia, Turkey, Georgia, Iran, Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan) (Assing 2021). It is confirmed as established in the Nearctic region and had been previ- ously known from North America under the synonym Tetralaucopora americana (Casey) (Klimaszewski et al. 2018). Assing (2021) recently reported this species from BC and treated T. americana as a tentative synonym based on the results presented in this paper. Although all available sequences of this species are partial (382–407 bp) and a BIN has not been established as that would require at least one founding member with a minimum sequence length of 500 bp, Nearctic and Palaearctic sequences form a distinct cluster with only a single variable nucleotide site. External morphology and that of the spermatheca are identical. As spermathecae are of generally poor diagnostic value (especially the distal part) in Parocyusa (Assing 2021), the barcode evidence was quite critical for the resolution of this issue. Based on this evidence and a distribution centered around populated areas in northeastern and western North America, we here consider this species to be adventive in the Nearctic region. At the moment, it is not yet possible to determine whether the population in BC is a separate introduction from the northeastern population. Recently, Assing (2021) revalidated Parocyusa as a genus separate from Tectusa after the discovery that Tectusa was not a monophyletic group. The type species of Parocyusa was found to be congeneric with that of Tetralaucopora, and the latter became a junior synonym of the former. Tribe Tachyusini C.G. Thomson Revised key to the Canadian genera of Tachyusini Adapted from Klimaszewski et al. 2018. 1 Elytra at humerus only slightly broader than pronotum at base (Figs 7, 8); impressions of abdominal tergites shallow, with punctation similar to that of disc (Figs 7, 8) ............................................................................................2 – Elytra at humerus distinctly broader than pronotum at base (Figs 9–13); im- pressions of abdominal tergites with at least a few coarse punctures and glossy areas, punctation distinctly different from that of disc (Figs 9–13) ..............3 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 43 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 impres- sions shallower, gradually sloping to disc at base (Fig. 13) ............................. .......................................................................................Gnypeta Thomson Paradilacra densissima (Bernhauer, 1909) BOLD:ACF7668 Fig. 7A–H Atheta (Paradilacra) densissima Bernhauer, 1909 Gnypeta saccharina Klimaszewski & Webster, 2008, syn. nov. Material (DNA barcoded specimens). Canada: Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane for- est, 27.VI.2012, BIOBus 2012 (1, CBG); British Columbia: 10 km W Kamloops, New Afton Mine, Wetland Protected Area, 50.662, -120.504, 702 m, malaise trap, 22.VIII.2013, C. Simon (1, CBG); New Brunswick: York Co., Fredericton at Saint John River, 45.959, -66.625, margin of river in flood debris, 7.VII.2005, R.P. Webster [note: paratype of G. saccharina] (1, LFC); Ontario: Guelph, University of Guelph Arboretum, 43.53, -80.21, 12.VI.2019, M. Pentinsaari (1, CBG). Distribution. Origin: Nearctic. Canada: AB, BC, NB, ON [new record], SK [new record]. United States: CA, MT, NV, ND, OR, UT. Bionomics. This species has been collected from various wetland microhabitats including the edges of lakes, rivers, and a beaver pond (Gusarov 2003a). Comments. Paradilacra densissima and the genus Paradilacra, widespread in west- ern and central North America (Gusarov 2003a), are newly reported from SK (records in Klimaszewski et al. 2016a, as G. saccharina) and eastern North America based on http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACF7668 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)44 Figure 7. Paradilacra densissima (Bernhauer) A habitus B median lobe of aedeagus in lateral view C me- dian 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. records from NB (Klimaszewski et al. 2008) and ON (this study), including one se- quenced paratype of synonym Gnypeta saccharina. Under the present concept, only one widespread species of this genus is known. Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 45 Figure 8. Brachyusa helenae (Casey) A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2016a). Dasygnypeta Lohse, 1974, sensu nov. In his key to the genera of Tachyusini, Paśnik (2010) distinguished Dasygnypeta, with its single Palaearctic species Dasygnypeta velata (Erichson), from the Nearctic genera by the following features: the narrow and slender abdomen with base ap- proximately as wide as head, the ‘very deep’ tergal impressions, abdominal pubes- Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)46 Figure 9. Tachyusa obsoleta Casey A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view (adapted from Paśnik 2006) D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission. cence of tergites III–V directed posteriad, basal segment of metatarsus shorter than following two segments. Through an analysis of barcode data, we have discovered that the recently described Gnypeta minuta Klimaszewski & Webster is a synonym Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 47 of D. velata (see below). A re-examination of other Nearctic Gnypeta species re- vealed two others that are closely related to D. velata: G. baranowskii Klimaszewski, and G. nigrella (LeConte). Their morphological divergence from other Gnypeta was represented by an earlier placement in the ‘Nigrella species group’ of Gnypeta by Kli- maszewski et al. (2008), together with G. saccharina (now a synonym of Paradilacra densissima). In corroboration with morphology, DNA barcodes of D. velata and G. nigrella form sister clusters (sequences of G. baranowskii not available). Transfer of these two Gnypeta species to Dasygnypeta required a new concept for this genus as most of the distinguishing features were apomorphies of D. velata or found not to be of diagnostic value due to variability or overlap with other genera. Here we distinguish members of Dasygnypeta by their characteristic abdomen (Figs 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 Gnypeta); tergites III–V with very deep basal impressions, each creating strongly convex areas on the disc. Members of Dasygnypeta could be confused with Tachyusa, which also bears an elongate basal abdomen, but in the latter genus tergites III–V are far more elongate and the abdominal base is narrower than the head (Fig. 9). Dasygnypeta nigrella was even originally described by LeConte (1863) in Tachyusa, likely based on this similarity. Dasygnypeta baranowskii (Klimaszewski, 2020), comb. nov. Fig. 10A–H Gnypeta baranowskii Klimaszewski, 2020 Distribution. Origin: Nearctic. Canada: BC. Bionomics. The type series was collected by sifting litter (Klimaszewski et al. 2020). Comments. We here transfer this species to Dasygnypeta sensu nov. based on morphology illustrated by Klimaszewski et al. (2020). This recently described western species is most similar to eastern D. nigrella based on the moderately elongate anten- nae, more robust body and the distinctive deep emargination of female sternite VIII. However, it is easily distinguished by the coarser pronotal punctation, and male and female genitalia (Fig. 10). The aedeagus of D. baranowskii is superficially similar to P. densissima but these taxa are externally quite different. Dasygnypeta nigrella (LeConte, 1863), comb. nov. BOLD:ACS6831 Fig. 11A–H Tachyusa nigrella LeConte, 1863 Gnypeta nigrella: Klimaszewski et al. 2008 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACS6831 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)48 Figure 10. Dasygnypeta baranowskii (Klimaszewski) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2020), used with permission. Material (DNA barcoded specimens). Canada: New Brunswick: York Co., Freder- icton at St. John River, 45.959, -66.625, margin of river in drift (mostly maple seeds), 4.VII.2004, R.P. Webster (1, LFC). Additional non-barcoded material. Canada: Manitoba: 5 miles SW of Shilo, 5.VI.1958, J.F. McAlpine (1, CNC); Quebec: Montreal, 30.VIII.1968, E.J. Kiteley Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 49 Figure 11. Dasygnypeta nigrella (LeConte) A habitus B median lobe of aedeagus in lateral view C me- dian 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. (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). Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)50 Distribution. Origin: Nearctic. Canada: MB [new record], NB, NF, ON, QC [new record]. United States: IL, MA, MD, NJ, NY, PA, VT, WV. Bionomics. Collected along the edge of a variety of running and standing water- based habitats. Comments. We here transfer this species to Dasygnypeta sensu nov. based on mor- phology and close clustering of DNA barcode sequences with D. velata. Dasygnypeta nigrella is a widespread species in eastern North America and is here newly reported from Manitoba and Quebec. Dasygnypeta velata (Erichson, 1837) BOLD:ACZ0581 Fig. 12A–H Homalota velata Erichson, 1837 Gnypeta minuta Klimaszewski & Webster, 2008, syn. nov. Material (DNA barcoded specimens). Germany: Thuringia, Ufergehoelze am Spei- cher Loessau, 50.5665, 11.894, 460 m, 1.I.2013, GBOL-Team ZFMK (1, ZFMK); Thuringia, NE, Freibad, Werraufer, 50.9768, 10.0963, 20.X.2014, GBOL-Team ZFMK (1, ZFMK). United States: Alaska: Selawik NWR, Kugarak River, 66.561, -158.996, mud bank, shore washing, 23.VI.2010, D.S. Sikes (3, UAM). Additional non-barcoded material. Canada: Manitoba: 5 miles SW of Shilo, 5.VI.1958, J.F. McAlpine (2, CNC); Northwest Territories: Inuvik, 24.VI.1972, A. Smetana (1, CNC). United States: Alaska: Hess Creek, mi. 24 Wales Hwy, 1.VII.1978, J.M. Campbell and A. Smetana (1, CNC); Kenai Peninsula, Anchor River at Hwy 1, 450’, 4.VI.1978, A. Smetana and E. Becker (1, CNC); mi. 1259 Alaska Hwy, 7.VII.1968, J.M. Campbell and A. Smetana (1, CNC). Distribution. Origin: Holarctic. Canada: MB [new record], NB, NF, NT, SK. United States: AK. Bionomics. Nearctic specimens have been collected most frequently along the margins of running water but also along the margins of a forest pool (Klimaszewski et al. 2018). Comments. Dasygnypeta velata is newly reported from North America and was previously known in the Nearctic region under the synonym Gnypeta minuta (Kli- maszewski et al. 2018). We here newly record this taxon from MB. Dasygnypeta velata is here considered a Holarctic species as it has a broad, transpalaearctic distribution (Europe to Siberia; Newton 2019), occurs along rivers and has been collected north of the treeline in Alaska. This species has been collected together with D. nigrella in southern Manitoba (see above). The barcode sequences of the specimens from Alaska are all partial (386 to 407 bp), but the overlapping parts of the sequences are identical to the two German sequences. http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACZ0581 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 51 Figure 12. Dasygnypeta velata (Erichson) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after Klimaszewski et al. (2018), used with permission. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)52 Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. http://zoobank.org/C6DC72D8-A182-43D3-9691-A352F0AF5D0E BOLD:ADH7347 Fig. 13A–H Type material. Holotype. (male, CNC): Canada, Ontario, Hartington, Eel Lake, South Frontenac, Paul Hebert’s cottage property, 44.563°N, 76.549°W, 6.13.2017, Mikko Pentinsaari, Barcode of Life DNA voucher specimen, Sample ID: BIOUG34206- H01, Process ID: MPCAN465-17. Paratypes (3 CBG, 4 CNC): Canada, labelled as the holotype except: Sample ID: BIOUG34206-H02, Process ID: MPCAN466-17 (1 male, CBG); Sample ID: BIOUG34206-G12, Process ID: MPCAN464-17 (1 male, CBG); Sample ID: BIOUG34206-G11, Process ID: MPCAN463-17 (1 female, CBG). United States: North Carolina: Haywood Co., 3 mi N Dellwood, 19.VIII.1972, A. Smetana (3, CNC); Maryland: Patuxent Wildl. Res. Ctr., 5 km E Montpelier, tread- ing pond vegetation, 16.VI.1982, Bousquet & Davies (1, CNC). Etymology. The species epithet refers to the longitudinal impression on the pro- notum, most strongly developed in males. Diagnosis. Gnypeta impressicollis can be easily distinguished from all Nearctic spe- cies of the genus (except eastern G. baltifera (LeConte)) by the hexagonal pronotum with a longitudinal impression in the basal half (females) to nearly entire pronotal length (males). Males also have an impression on the vertex of the head. We have examined the female type of G. baltifera and it is externally similar but differs by the shorter, less angulate hexagonal pronotum, reddish and longer elytra and spermatheca with an elongate stem (C-shaped in G. impressicollis). Description. Body length 3.2–3.4 mm; colour dark brown, elytra brown with ir- regular 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 api- cal 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); http://zoobank.org/C6DC72D8-A182-43D3-9691-A352F0AF5D0E http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ADH7347 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 53 Figure 13. Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). 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 sub- spherical with broad apical invagination, stem tubular and C-shaped (Fig. 13D). Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)54 Distribution. Origin: Nearctic. Canada: ON. United States: MD, NC. Gnypeta impressicollis is probably broadly distributed in eastern North America. Bionomics. Specimens were collected by sifting leaf litter along a lake margin and by treading pond vegetation. Comments. It was challenging to place this species in either Gnypeta or Ischnopoda Stephens based on the concepts of Pašnik (2010). The extremely long legs, pronotal shape, C-shaped spermatheca and superficial punctation of the pronotum and abdo- men are consistent with at least some Neotropical members of Ischnopoda but the ligula of G. impressicollis is divided to the base, which is considered to be a feature of Gnypeta (Pašnik 2010). The C-shaped spermatheca of Gnypeta impressicollis also bears some similarity to the G. crebrepunctata group of Klimaszewski et al. (2008) but it is rather different in external morphology. We place this species in Gnypeta pending fu- ture systematic research. Tribe Hypocyphtini Laporte, 1835 Oligota parva Kraatz, 1862 BOLD:AAP9955 Fig. 14A–G Material (DNA barcoded specimens). Germany: Bornheim-Hemmerich, Ortslage, 50.7596, 6.93151, 30.VII.2010, F. Koehler (2, ZSM); Bornheim-Hemmerich, Ort- slage, 50.7596, 6.93151, 25.VIII.2013, GBOL-Team ZFMK (1, ZFMK); Wutha- Farnroda, Wartburgkreis, 50.947, 10.4214, 25.VIII.2012, GBOL-Team ZFMK (2, ZFMK). Canada: Ontario: Kawartha Lakes, 44.296, -78.452, farm, malaise trap, 24.VII.2015, B. McClenaghan (1, CBG); same except 19.VII.2016 (1, CBG); Guel- ph, Arboretum, Urban Organic Farm, 43.5381, -80.222, compost heaps and mouldy hay pile, 17.IX.2017, M. Pentinsaari (4, CBG). Distribution. Origin: West Palaearctic (adventive in North America). Canada: NB, ON [new record], PE. United States: CA, MA, MO, NV, TX. Bionomics. This species is generally found in anthropogenic habitats, including compost, dung, and old hay and grass (Klimaszewski et al. 2018). In Canada, it has been collected in compost and in ocean coastline drift at the top of the littoral zone (Klimaszewski et al. 2018). Comments. Oligota parva is a cosmopolitan species that is adventive in Canada. Here we newly report it from Ontario. Oligota pumilio Kiesenwetter, 1858 BOLD:AAN4271 Fig. 15A–C Material (DNA barcoded specimens). Belgium: Blanden, BR Meerdaalboos, 50.7976, 4.71622, 8.V.2010, F. Koehler (1, ZSM). Germany: Neuburg, Altrheine, http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAP9955 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAN4271 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 55 Figure 14. Oligota parva Kraatz A habitus B median lobe of aedeagus in lateral view C male tergite VIII D male sternite VIII E female tergite VIII F female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–F). Illustrations after Webster et al. (2016). 48.9943, 8.24412, 29.IX.2011, F. Koehler (2, ZSM); Erftstadt-Bliesheim, NWZ Al- twald Ville, 50.792, 6.844, 4.X.2010, F. Koehler (1, ZSM); Jockgrim, Sandmagerra- sen, 49.0805, 8.26568, 14.XI.2010, F. Koehler (1, ZSM); Ochtendung, Michelsberg, 50.3631, 7.3889, 17.III.2012, F. Koehler (1, ZSM); Edenkoben-Rhodt, Villa Ludwig- shoehe, 49.2767, 8.08991, 20.V.2012, F. Koehler (1, ZSM); Bad Muenster-Traisen, Rotenfels, 49.822, 7.832, 20.V.2012, F. Koehler & J. Koehler (1, ZSM); Osterholz bei Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)56 Figure 15. Oligota pumilio Kiesenwetter A habitus (image by A. Bogri – www.BilleBank.dk) B median lobe of aedeagus in lateral view (drawn from Kapp 2019) C spermatheca (drawn from Kapp 2019). Scale bars: 1 mm (A); 0.2 mm (B, C). Blankenburg, 51.9519, 11.0526, 18.III.2015, GBOL-Team ZFMK (2, ZFMK). Can- ada: Alberta: Waterton Lakes National Park, Red Rock Parkway, 49.088, -113.883, Moraine grassland, intercept trap, 1328 m, 11.VIII.2012, BIOBus 2012 (1, CBG). United States: Montana: Missoula County, Florence, MPG Ranch, 46.702, -114.034, grassland, pitfall trap, 05.VI.2019, M. Seidensticker (1, CBG). Distribution. Origin: West Palaearctic (adventive in North America). Canada: AB [new record]. United States (all except MT need verification): DC, IL, OH, MT [new record]. Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 57 Diagnosis. Among Canadian species of Oligota, O. pumilio is extremely similar to O. pusillima in the narrow, parallel body (Fig. 15A) and in male and female genitalia (Klimaszewski et al. 2018). However, it can be distinguished by the more abruptly truncate apex of the median lobe in lateral view (Fig. 15B), differently shaped sclerites of the internal sac (Fig. 15B), medially projected apex of male sternite VIII, and the transverse capsule of the spermatheca (Fig. 15C) (Kapp 2019). Bionomics. This species occurs in a wide variety of habitats across a broad eleva- tional range, including hollow trees, plant debris, old hay in cattle barns, moldy sub- strates and in mushrooms (Kapp 2019). The barcoded Nearctic specimens were collect- ed from grassland habitats by an intercept trap (Alberta) and a pitfall trap (Montana). Comments. Oligota pumilio is a West Palaearctic species that is adventive in Canada. Although it has been previously reported from the United States (OH, IL, DC) (Newton 2019), these records need confirmation as they are in the east, some distance away from the present records. This species’ presence in North America is thus verified here for the first time, from both Canada (AB) and United States (MT). It has also been reported as adventive from Argentina, Chile, and New Zea- land (Newton 2019). The barcoded specimens of O. pumilio from Canada and the United States share the same DNA barcode haplotype, which is also shared by some of the specimens from Germany. Oligota pusillima (Gravenhorst, 1806) BOLD:ABW7320 Fig. 16A–G Material (DNA barcoded specimens). Finland: Oba: Oulu, Hietasaari, 65.0225, 25.4247, 22.IV.2011, M. Pentinsaari (1, ZMUO); Germany: Edenkoben-Rhodt, Vil- la Ludwigshoehe, 49.277, 8.09, 20.V.2012, F. Koehler (2, ZSM); Edenkoben-Rhodt, Villa Ludwigshoehe, 49.277, 8.09, 23.VI.2012, F. Koehler (2, ZSM); Zweibrueck- en-Mauschbach, Monbijou-Wald, 49.2038, 7.39891, 16.X.2011, F. Koehler & W. Koehler (1, ZSM). Canada: Alberta: Two Hills, Two Hills School EQP-CLL-553, 53.7104N, 111.7437W, 613 m, Malaise trap, 21.IX.–2.X.2015, K. Warawa (2, CBG). Additional non-barcoded material. Ontario: Ottawa, Ottawa River, Deschênes Lookout, Berlese flood debris, 1.V.1985, A. Davies (1, CNC). Distribution. Origin: West Palaearctic (adventive in North America). Canada: AB [new record], NB, ON [new record]. United States: MA, NY. Bionomics. This species occurs in a variety of moist to dry, decaying organic mat- ter including rotting hay, compost, hollow trees, and ant nests (Kapp 2019). Canadian specimens were collected in compost (Webster et al. 2016), and in malaise traps and flood debris in a suburban setting (present study). Comments. Oligota pusillima is a Palaearctic species that has been introduced to North America, South America, Australia, Africa, and southeast Asia (Kapp 2019, Newton 2019). It is here reported from Ontario and Alberta for the first time, the latter representing the westernmost record in North America. Specimens from Alberta http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ABW7320 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)58 Figure 16. Oligota pusillima (Gravenhorst) A habitus B median lobe of aedeagus in lateral view C sper- matheca 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). were sequenced and their barcodes match those of Palaearctic specimens with no diver- gence. These specimens were also morphologically consistent with O. pusillima. Homalotini Heer, 1839 Anomognathus athabascensis Klimaszewski, Hammond & Langor, 2016 Fig. 17A–G Material. Non-sequenced specimens. Canada: Manitoba: Winnipeg, under bark of rotten ‘N. aceroides’ [= Acer negundo], 27.VIII.1918, J.B. Wallis (2, CNC). Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 59 Figure 17. Anomognathus athabascensis Klimaszewski, Hammond & Langor A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G fe- male sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2016b). Distribution. Origin: Nearctic. Canada: AB, MB [new record]. Bionomics. The MB specimens were collected under bark, confirming that this species lives in a way similar to other members of the genus. Comments. Anomognathus athabascensis, recently described from Alberta (Kli- maszewski et al. 2016b), is newly reported from Manitoba. This native Nearctic spe- cies is likely transcontinental but rarely reported due to its small size and elusive habits. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)60 Anomognathus cuspidatus (Erichson, 1839) BOLD:AAO0339 Figs 18A–D, 19A–D, 20A–D. Homalota cuspidata Erichson, 1839 Thectura americana Casey, 1893, syn. nov. Anomognathus americanus: Seevers (1978) (as valid species) Type material. Homolota cuspidata Erichson, 1839. Lectotype, male, here designated (ZMHB): cuspidata Er: [handwritten label] / 5387 [typed label] / Hist.-Coll. (Coleop- tera), Nr. 5387, Homalota cuspidata, Erichs., Europa, Zool. Mus. Berlin [typed white label] / Lectotype Homalota cuspidata des. J. Klimaszewski 2019 [white printed label]. Paralectotypes (3, ZMHB, without original labels): Hist.-Coll., (Coleoptera), Nr. 5387, Homolota cuspidata Erichs., Europa, Zool. Mus. Berlin; Syntype Homolota cus- pidata Erichson, 1837, labelled by MNHUB 2010; Paralectotype Homalota cuspidata des. J. Klimaszewski 2019 [white printed label] [1 female, spermatheca and terminalia dissected in Canada balsam on microslide attached to specimen]; same labels except: SYNTYPUS, Homalota cuspidata Erichson, 1837 [typed red label, added by MNHUB 2010] [1 female, spermatheca and terminalia dissected in Canada balsam on microslide attached to specimen]; same labels as before [1 damaged specimen, sex undetermined]. Males and females of the syntype series were morphologically consistent with the specimens forming molecular cluster BOLD:AAO0339, including those sequenced from Ontario, Canada. As the most obvious difference between A. cuspidatus and the potential new Central European species (see Diagnosis) was the shape of the median process on male tergite VIII (in lateral view) (Fig. 18B, D), a male syntype (see above) was designated as the lectotype of this species to fix its identity. Morphology of the ae- deagus itself was difficult to study due to its small size and obvious differences between molecular clusters (see below) were not observed (Fig. 19). Thectura americana Casey, 1893, syn. nov. Holotype (male) (NMNH): NY/ TYPE USNM 39614/ Thectura americana Casey (handwritten by Casey). Casey (1893) gave numerous characters to distinguish A. americanus from A. cus- pidatus but all of these were observed to be highly variable within populations in the material studied, including the shape of apical antennomeres, shape of the pronotum, position of the abdominal tubercles in the male, and the type of dorsal expansion of the median process of male tergite VIII. We could not find the depression at the base of tergite VIII on the holotype of A. americanus mentioned by Casey (1893). Although the aedeagus of the holotype was not studied (not extracted from partly damaged and frag- ile pygidium), male tergite VIII was intact and its median process in lateral view bears an apical hook, matching the present concept for A. cuspidatus (Fig. 20). Therefore, in corroboration with Fenyes (1918), we treat A. americanus as a synonym of A. cuspidatus. Non-type material (sequenced specimens indicated in square brackets). Cana- da: 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]). http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAO0339 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAO0339 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 61 Figure 18. A, B Anomognathus cuspidatus (Erichson), and C, D Anomognathus sp., putative undescribed spe- cies (Europe) A, C habitus and B, D male tergite VIII in lateral view. Scale bars: 1 mm (A, C); 0.2 mm (B, D). A photo record of this species from Ontario is available on bugguide.net (/view/1816108): Toronto, 19.V.2020, under bark, O. Strickland. Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 28.IV.2010, F. Koehler (1, ZSM [1 sequenced]). Czech Republic (all CNC): Bohemia, Poděbrady Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)62 Figure 19. Aedeagi of A–C Anomognathus cuspidatus (Erichson) and D potential undescribed species, in lateral view (top row) and dorsal view (bottom row) A sequenced non-type (Ontario, Canada) B lectotype of A. cuspidatus (‘Europe’) C, D sequenced non-types (Finland). Scale bar: 0.2 mm. 50 km, Smetana, 1959, car net trap (1); Bohemia, Chvojno, Smetana (1); Moravia, Drnholec, Smetana (1). Denmark (all NHMD): Staksrode, EJ, 24.IX.1983 (1); Æbelø F, 18.V.1997 (1); Faested Mose, SJ, 12.IV.1986 (1); Dyrehaven, 14.4.1934 (1); same except 21.3.1923 (1); same except 21.10.1932 (1); 30.4.1922 (1); same except 19.5.1911 (1); Lyng Huse, 29.3.1997 (1). Germany: Nationalpark Mueritz, Babke-Zartwitz-Speck-Schwarzenhof, 53.4125, 12.8463, car net, 20.VI.2015, GBOL-Team ZFMK (2, ZFMK [2 sequenced]); Hoenningen bis Insul, Ahrtal, 50.45, 6.942, 24.IV.2010, F. Koehler (1, ZSM [1 se- quenced]); Oberheimbach, Franzosenkopf, 50.004, 7.805, 27.V.2012, W. Koehler (1, Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 63 Figure 20. Male tergite VIII of A–C Anomognathus cuspidatus (Erichson) and D potential undescribed spe- cies, in dorsal (top row) and lateral (middle and bottom rows) A lectotype of A. cuspidatus (‘Europe’) B holo- type of A. americanus (Casey) (= A. cuspidatus) C, D sequenced, non-types (Finland). Scale bar: 0.2 mm. ZSM [1 sequenced]). Finland (all ZMUO): N: Espoo, Saunalahti, 60.1643, 24.6263, 17.IX.2012, fungusy aspen logs, E. Helve (1) [barcoded]; Al: Bjoerkoe, 59.9769, 20.1879, sifting, 24.IX.2014, M. Pentinsaari (1) [barcoded]; Ta: Lammi, R. Linna- vuori leg. (1); Ab: Naantali, R. Linnavuori leg. (1); Kb: Lieksa, R. Linnavuori (1); Rynmattyla, 24.VI.1945, Karvonen (2); same except 14.VIII.1945 (2). Slovakia (all CNC): Cenkov, Smetana, 1963 (11); Nová Sedlica, Smetana, 1961 (2); Ruská Poruba, Smetana, 1956 (2). United Kingdom (all CNC): Essex (6). Putative undescribed Anomognathus (corresponding to BIN BOLD:ACA9191): Finland (all ZMUO): N: Espoo, E. Helve, 1978 (1); same except 1976 (1); same except 1977 (1); same except 1979 (1); same except 1981 (1); same except 1982 (1); Ks: Taivalkoski, 728.53 Window trap, 2003, E. Hurme (2); same except Polyporus trap (1); Kb, Kitee, 23.05.2016, M. Pentinsaari leg., [1 sequenced]; Obb: Rovaniemi, Rovajärvi, 16.6– 8.7.2010, M. Pentinsaari and E. Kuusela [1 sequenced]. Germany: Schleiden-Wolfgarten, Dachsloecher, 50.6098, 6.42237, 26.VII.2012, F. Koehler (1, ZSM [1 sequenced]). Diagnosis. Anomognathus cuspidatus is distinctive for its trident-shaped apex of male and female tergite VIII (Fig. 20A–C) and can be distinguished from all described http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACA9191 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)64 species by this feature alone. However, in the course of this study, specimens repre- senting a remarkably divergent barcode cluster (BOLD:ACA9191; 9.63% uncorrected p-distance to A. cuspidatus) were investigated and found to likely represent an unde- scribed species of Anomognathus in Europe (confirmed specimens from Finland and Germany). Although most morphological characters of A. cuspidatus and the putative new species are highly variable, including the median lobe of the aedeagus, males can be dependably separated based on the shape of their median process of tergite VIII in lateral view: A. cuspidatus bears a minute to distinct hook at the apex (Fig. 20A–C), while in the undescribed species, the median process converges evenly to a single point, creating an elongate, turnip-shape (Fig. 20D). The shape of tergite VIII in females was observed to be extremely variable and no features were deemed to be diagnostic. Externally, most specimens can be recognized as either species (especially males) by the relative proportions of the head versus the pronotum, with A. cuspidatus generally bearing a small pronotum, narrower than the head (Fig. 18A) and the undescribed species bearing a wider, longer pronotum, wider than the head (Fig. 18C). The limits of this taxon need further investigation and should include morphological study of a much wider range of sequenced material. Distribution. Origin: West Palaearctic (adventive in North America). Canada: AB, NB, ON. United States: NY. Bionomics. This species occurs under the bark of dead trees. One specimen (NB) was collected from a Lindgren funnel. Comments. Anomognathus cuspidatus is a widespread West Palaearctic species that is known from Europe, European Russia and Algeria (Newton 2019) and has been previously known in North America under the synonym A. americanus. The record from Beijing, China should be verified. The species has become introduced in North America (before 1893) and it is unclear whether the population in Alberta represents a separate introduction from Europe, a secondary introduction from eastern North America or a broad adventive distribution across Canada. After the results of the present study, two species of Anomognathus are known to oc- cur in North America: native A. athabascensis Klimaszewski, Hammond & Langor and the adventive A. cuspidatus. These are easily separated by the drastically different shapes of male and female tergites VIII (Figs 17D, 20A–C). Previously, only females of A. cuspidatus (as A. americanus) were available from Canada (Klimaszewski et al. 2016b; Webster et al. 2016). Here we demonstrate that all available Nearctic Anomognathus specimens with a trident-shaped tergite VIII correspond to Palaearctic A. cuspidatus. Cyphea wallisi Fenyes, 1921 BOLD:ACK1459 Figs 21A–G, 22A–D Cyphea wallisi Fenyes, 1921 Agaricomorpha vincenti Klimaszewski & Webster, 2016, syn. nov. Agaricomorpha vincenti: Klimaszewski et al. 2018 (as synonym of C. curtula) http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACA9191 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACK1459 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 65 Figure 21. Cyphea wallisi Fenyes A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after Klimaszewski et al. (2018), used with permission. Type material. Cyphea wallisi Fenyes, 1921. Paratype, male (MCZ). Winnipeg, Man. [handwritten label] / Wallis [handwritten label] / 25490. / Cyphea, Wallisi, Feny [handwritten label] / Type., 9989, 9983 [typed red label]. Agaricomorpha vincenti Klimaszewski & Webster, 2016, syn. nov. Holotype, male (LFC). Canada, New Brunswick, Carleton Co., Jackson Falls, “Bell Forest”, 46.2200°N, 67.7231°W, 7–21.VI.2012, C. Alderson & V. Webster, coll. [white typed label] / Rich Appalachian hardwood forest, Lindgren funnel trap in canopy of Fa- gus grandifolia [white typed label] / Holotype Agaricomorpha vincenti Klimaszewski & Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)66 Webster, 2016 [red typed label] / Cyphea curtula (Erichson) det. Klimaszewski 2017 [white typed label] / Cyphea wallisi Fenyes det. A. Brunke 2020. The aedeagi of the male paratype (holotype in collection of the California Acad- emy of Sciences) of C. wallisi and holotype of A. vincenti are identical and both dif- fer from that of Palaearctic Cyphea curtula (image by V. Assing) by the broader distal lobe in lateral view, which only slightly extends beyond the distal plate (Fig. 22A–C). Therefore, we transfer Agaricomorpha vincenti from synonymy with Cyphea curtula to synonymy with Cyphea wallisi. Other material (DNA barcoded specimens). Canada: Ontario: Rouge National Urban Park, Toronto Zoo, 43.8223, -79.1897, forest, malaise trap, 25.VI.2013, L. Attard and K. Greenham (2, CBG); Hartington, Eel Lake Cottage, Lindgren funnel trap, 44.5628, -76.553, 25.VII.2017, G. Blagoev (1, CBG); Nova Scotia: Clyburn Valley Road, near golf course, Cape Breton National Highlands NP, forest, Malaise trap, 46.6553, -60.4285, 28.VI.2013, CBH staff (1, CBG). Non-barcoded specimens. Canada: Quebec: Mont St. Bruno Prov Park, 45.541, -73.319, Lindgren funnel, trap 5, tree 2, beech-maple canopy, 21.VII-3.VIII.2005 (1, CNC); Oka Prov Park, Lindgren funnel, trap 3, tree 1, beech-maple canopy, 27.VII.30. VIII.2005 (1, CNC). Distribution. Origin: Nearctic. Canada: AB, MB, NB, NS [new record], ON [new record], QC. Bionomics. Specimens have been collected in Malaise traps, window traps and Lindgren funnels placed in forests. Both the closely related West Palaearctic C. curtula and C. latiuscula Sjöberg have been consistently collected under bark, where they occur in the larval burrows of various longhorn beetles (Cerambycidae), bark beetles (Curcu- lionidae: Scolytinae) and the carpenter moth (Cossus L.) (Palm 1968). Comments. Cyphea wallisi is a broadly distributed native Nearctic species, report- ed from AB east to NS. Here we treat Nearctic records of Cyphea as C. wallisi (previ- ously treated as Palaearctic C. curtula, e.g., Klimaszewski et al. 2018) and newly report the genus from ON and NS. Cyphea wallisi is probably far more broadly distributed in North America than currently known and has been underreported due to its small size. Sequenced Nearctic specimens of Cyphea from ON and NS formed a barcode cluster that was nearly 5% divergent from those of Palaearctic specimens of C. curtula (BOLD:AAO1175, one published sequence record from Belgium and three unpub- lished records from the Netherlands). Northern European C. latiuscula, the only other species of the genus, has a broader body outline, different male genitalia and is quite differently colored (bicolored pronotum and pale elytra). No barcode sequence data are currently available for C. latiuscula. Based on the study of one paratype of C. wal- lisi, described from Manitoba and not reported since, it was discovered that Nearctic specimens of Cyphea correspond to this species and differ from Palaearctic C. curtula by the broader distal lobe of the median lobe of the aedeagus in lateral view, which only slightly extends beyond the distal plate (Fig. 22A–C). The shape of the median lobe of the aedeagus in dorsal view may also be diagnostic (Fig. 22D) but a prepara- tion in this view was unavailable for C. curtula. The illustration in Palm (1968) of the http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAO1175 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 67 Figure 22. Aedeagi of Cyphea wallisi Fenyes (A, B, D) and C. curtula (Erichson) (C), in lateral (A–C) and dorsal (D) view. Paratype of C. wallisi (A, D); holotype of Agaricomorpha vincenti Klimaszewski and Webster (= C. wallisi) (B); non-type, C. curtula (image by V. Assing) (C). Scale bar: 0.2 mm. aedeagus of C. curtula in dorsal view appears to be less angulate than that of C. wallisi but this needs verification. Based on these differences in male genitalia (Fig. 22A–C) and the COI barcodes, Cyphea wallisi is morphologically and genetically distinct from Palaearctic C. curtula, and the latter species does not occur in North America as far as known. Both of these species have a median tubercle on male tergite VII, mentioned earlier by Fenyes (1921) but this structure was omitted from the illustrations in Kli- maszewski et al. (2018), though it was present in the original description of synonym Agaricomorpha vincenti (Webster et al. 2016). Previous differences between the two species given by Klimaszewski et al. (2018) (e.g., projecting pronotal angles, lighter/ darker body) proved to be highly variable. Gyrophaena affinis Mannerheim, 1830 BOLD:ACF7981 [Nearctic]; BOLD:ABW9049 and BOLD:AAO0291 [both Palae- arctic] Fig. 23A–G Material (DNA barcoded specimens). Belgium: Sint-Genesius-Rode, BR Zonien- woud, 50.7505, 4.423, 135 m, 16.VI.2010, F. Koehler (1, ZSM). Finland: Oba: Oulu, Linnanmaa, 65.0633, 25.4712, 7.VI.2011, M. Pentinsaari (1, ZMUO); Obb: Tornio, Kalkkimaa, 65.9014, 24.4711, 10.VII.2012, M. Pentinsaari (1, ZMUO); Al: Lemland, Herrövägen, 59.9796, 20.1954, car net, 5.VII.2012, M. Pentinsaari (1, ZMUO). Germany: Brohl-Luetzing, Brohltal, 50.4727, 7.31272, 22.V.2010, F. Koehler (1, ZSM), Riedlhuette, Diensthuettenstrasse, 48.937, 13.412, 09.VII.2011, F. Koehler & M. Koehler (1, ZSM), Waldhaeuser, Lusen- und Boehmstrasse, 48.93, http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACF7981 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ABW9049 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAO0291 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)68 Figure 23. Gyrophaena affinis Mannerheim A habitus B median lobe of aedeagus in lateral view C sper- matheca 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. 13.492, 09.VII.2011, F. Koehler & M. Koehler (1, ZSM). Canada: Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane forest, 27.VI.2012, BIOBus 2012 (2, CBG). Distribution. Origin. Uncertain. Canada: AB [new record], BC, MB, NB, NF, NS, ON, QC, SK. United States: AZ, DC, IL, IN, IA, KY, MA, ME, MI, MN, MO, NC, NH, NJ, NM, NY, OH, PA, TN, WA, WI, WV. Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 69 Comments. Gyrophaena affinis is newly reported from AB based on barcoded material. Sequenced Nearctic specimens from ON, AB, NB, and QC form a distinct bar- code cluster, separate from all sequenced Palearctic specimens and divergent by 4.65%. This pattern is inconsistent with a species that is adventive in North America and we remove G. affinis from the list of adventive species in Canada. In comparing images between those of Nearctic specimens (Fig. 23B) and those of Enushchenko and Se- menov (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. Gyrophaena gracilis Seevers, 1951 Fig. 24A–H Material (non-sequenced material). Canada: Quebec: Gatineau Park, wolf trail, near trail start, 45.541, -75.912, hardwood forest, Polyporus squamosus on large beech log, 8.VI.2019, A. Brunke & J. Smith (1, CNC). Distribution. Origin. Nearctic. Canada: NB, QC [new record]. United States: WI. Bionomics. Specimens have been collected from a partly dried Pleurotus mush- room, from within the pores of a Trametes polypore, and from the nest contents of a Barred owl (Strix varia Barton) (Klimaszewski et al. 2018). The specimen from QC was collected from Polyporus squamosus on a beech tree. Comments. The new record from QC, near the ON border, bridges the wide gap between previous records in NB and WI. Gyrophaena simulans Seevers, 1951 BOLD:ACY8004 Fig. 25A–G Material (DNA barcoded specimens). Canada: Ontario: Hartington, Eel Lake Cot- tage, 44.563, -76.549, deciduous forest, mushrooms, 4.X.2017, M. Pentinsaari (2, CBG). Distribution. Origin. Nearctic. Canada: ON [new record]. United States: IL, MD, PA. Diagnosis. Gyrophaena simulans is extremely similar to G. criddlei and G. pseudocrid- dlei but has a slightly more transverse and flatter pronotum, with straighter apical and basal margins, and differently shaped upper process of the median lobe in lateral view (Fig. 25B): longer than that of G. pseudocriddlei but shorter and broader than that of http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACY8004 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)70 Figure 24. Gyrophaena gracilis Seevers A habitus B, C median lobe of aedeagus in lateral view D sper- matheca 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. G. criddlei. The emargination of male tergite VIII in G. simulans appears to be shallower and broader than that of G. criddlei but more specimens are needed to confirm this. Bionomics. The Canadian specimens were collected by sifting mushrooms in a deciduous forest. No detailed data on the host fungus were recorded. Comments. Gyrophaena simulans is a native Nearctic species distributed in eastern North America and is newly reported from Canada. The barcode cluster Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 71 Figure 25. Gyrophaena simulans Seevers A habitus B median lobe of aedeagus in lateral view C sper- matheca 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). BOLD:ACY8004 also contains specimens identified as related species G. criddlei (fe- male) and G. pseudocriddlei but more research, with broader sampling of sequenced, identified males, is needed to determine whether these species share a BIN or these specimens are misidentified. As we were unable to verify the identifications at this time, these records are not published here. http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACY8004 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)72 Homalota plana (Gyllenhal, 1810) BOLD:ADH5714 [Nearctic]; BOLD:AAO0434 [Palaearctic] Fig. 26A–G Material (DNA barcoded specimens). Belgium: Sint-Genesius-Rode, BR Zonien- woud, 50.7505, 4.423, 28.IV.2010, F. Koehler (1, ZSM). Germany: Arnsberg-Bre- itenbruch, NWZ Hellerberg, 51.446, 8.135, 30.V.2011, F. Koehler (2, ZSM); Heim- bach-Blens, Linkheld, 50.648, 6.468, 29.VIII.2012, F. Koehler (2, ZSM); Erftstadt- Bliesheim, NWZ Altwald Ville, 50.7917, 6.84384, 03.VI.2011, F. Koehler (1, ZSM); westl. Klein-Quenstedt, 51.9239, 11.0478, 20.III.2015, GBOL-Team ZFMK (1, ZFMK). Finland: Al: Finström, Norrö, 60.2458, 19.822, 5.VII.2012, M. Pentinsaari (1, ZMUO); Ka: Joutseno, Kuurmanpohja, 61.071, 28.75, 3.VIII.2012, M. Pentin- saari (1, ZMUO). Canada: Ontario: Guelph, Eramosa River Trail, 43.539, -80.236, deciduous forest, 14.IV.2017, M. Pentinsaari (2, CBG). Distribution. Origin. Uncertain. Canada: AB, MB, NB, NF, NS, ON. United States: AZ, CA, CO, IA, ID, IN, MT, NY, OH, PA, TX. Bionomics. Specimens occur under bark of dead trees. Comments. Sequenced Nearctic specimens from ON form a distinct barcode cluster, separate from all sequenced Palearctic specimens and divergent by 7.58%. This pattern is inconsistent with a species adventive in North America and we remove H. plana from the list of adventive species in Canada. Preliminary comparisons between images of Palaearctic and Nearctic specimens revealed that there may be some slight differences in the shape of the spermatheca. More research is needed to determine the status of the Nearctic and Palaearctic populations, though the level of genetic diver- gence between discrete Nearctic and Palaearctic populations suggests that two sister species are involved. Thecturota tenuissima Casey, 1893 BOLD:AAO0406 Fig. 27A–G Thecturota tenuissima Casey, 1893 Atheta marchii Dodero, 1922, syn. nov. Pragensiella magnifica Machulka, 1941, syn. nov. Thecturota marchii: Muona 1984 (as valid species) Thecturota magnifica: Schülke and Smetana 2015 (as syn. of T. marchii) Material (DNA-barcoded specimens). Germany: Kobern-Gondorf, Ortslage/Wein- berge, 50.308, 7.460, 21.V.2010, F. Koehler (1, ZSM); Edenkoben-Rhodt, Villastrasse, 49.279, 8.092, 20.X.2012, F. Koehler (1, ZSM). Finland: Oba: Oulu, Linnanmaa, 65.0633, 25.4712, botanical garden, compost heap, flight-intercept trap, 7.VI.2011, M. Pentinsaari (1, ZMUO). http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ADH5714 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAO0434 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAO0406 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 73 Figure 26. Homalota plana (Gyllenhal) A habitus B median lobe of aedeagus in lateral view C sper- matheca 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. Non-sequenced material. Several males and females of T. tenuissima from Den- mark (NMHD) were compared with illustrations from Klimaszewski et al. (2017). Distribution. Origin. Nearctic (adventive in West Palaearctic). Canada: ON, QC. United States: RI. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)74 Figure 27. Thecturota tenuissima Casey A habitus B median lobe of aedeagus in lateral view C sper- matheca 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). Bionomics. Canadian specimens were collected by car-netting in mixedwood for- ests, while Palaearctic specimens are known from compost and other plant-based de- bris (Horion 1967). Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 75 Comments. Thecturota tenuissima is native to the Nearctic region and has become accidentally introduced to the West Palaearctic, including the Canary Islands, where it was previously known under the synonym T. marchii (Newton 2019). We expect this species to be broadly distributed in eastern North America and has been overlooked over much of its range because car-netting, an effective method for collecting small, obscure staphylinids, is rarely used in the Nearctic region. Nearctic and Palaearctic populations do not differ in male and female genitalia or in external morphology. Molecular data were unavailable for the Nearctic population, which was recently reported from Canada (Klimaszewski et al. 2017) but described from Rhode Island, USA in 1893 (Casey 1893). However, we are confident that these species are synonyms. Muona (1984) stated that T. marchii is a ‘recent’ introduction to Europe but from an unknown source. Thecturota is primarily a New World genus, with ten described species in North and South America (Newton 2019). Thecturota magnifica (Machulka) is currently treated as a synonym of T. marchii (Newton 2019) and we simply transfer this name to synonymy with T. tenuissima. The only Palaearctic species remaining is poorly known T. williamsi (Bernhauer, 1936), known only from the type collected in Great Britain and probably a synonym of T. tenuissima. The characters Bernhauer (1936) gave to separate his species from T. tenuissima (as T. marchii) are slight differences in coloration and body proportions, which are both highly variable in the Palaearctic specimens of T. tenuissima studied. Therefore, we consider T. tenuissima to be a native Nearctic species that has become adventive in the West Palaearctic and suggest that genus Thecturota is naturally restricted to the Nearctic and Neotropical regions. Geostibini Seevers, 1978 Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov. http://zoobank.org/A72E1E63-3D1B-4CC8-882C-6921E5FC3D33 BOLD:AAY6554 Fig. 28A–G Type material. Holotype (male) (CNC): Canada: ON: Waterloo County, Cam- bridge, Rare Charitable Research Reserve, 43.390, -80.374, soybean field, pitfall trap, 29.VI.2010, A. Brunke [white printed label] / HOLOTYPE Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari sp. nov., des A. Brunke 2020 [red printed label]. Paratypes (1 NMNH, 6 DEBU, 1 CNC): Canada: ON: Wellington County, Eramosa, 43.616, -80.215, soybean field, pitfall trap, 13.VII.2010, A. Brunke [white printed la- bel] / PARATYPE Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari sp. nov., des A. Brunke 2020 [yellow printed label] [CNC, DEBU, 7 specimens]. United States: VA: Arlington County, Marcey Creek, 38.9087, -77.1083, 70 m, suburban backyard, Malaise trap, 14–21.VI.2015, S. Miller [white printed label] / Barcode of life, DNA voucher specimen, Sample ID: BIOUG42376-E12, Process ID: GMUAF1698-18 [yellow printed label] / PARATYPE Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov., des A. Brunke 2020 [yellow printed label] [NMNH, 1 specimen]. http://zoobank.org/A72E1E63-3D1B-4CC8-882C-6921E5FC3D33 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:AAY6554 Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)76 Figure 28. Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII (structure ac- cidentally over-cleared in balsam preparation) F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Non-type material (barcoded specimens). Canada: Ontario: Guelph, 25 Divi- sion St., 43.554, -80.264, Malaise trap, 14.VII.2010, A. Smith (1, CBG); Guelph, John F. Ross CVI, 43.5621, -80.247, Malaise trap, 22.IV-03.V.2013, G. Staines (1, CBG); Milverton, Milverton Public School, 43.568, -80.928, Malaise trap, 22.IV- 03.V.2013, J. Van Bakel (1, CBG); Collingwood, Collingwood Collegiate Institute, Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 77 44.489, -80.215, 188 m, Malaise trap, 22.IV-05.V.2014, A. Breton (1, CBG); Cam- bridge, rare Charitable Research Reserve, 43.3736, -80.3652, 304 m, 04–11.VI.2015, BIO Collections Staff (1, CBG). Etymology. The species epithet refers to the similarity to related species A. gre- garia (Erichson), which was originally treated separately from other Aloconota under subgenus Glossola Fowler (e.g., Benick 1954) because it lacks obvious male secondary sexual characters. Diagnosis. Aloconota pseudogregaria can be easily distinguished from all other species of the genus occurring in eastern North America by the distinctly bicolored abdomen (Fig. 28A). Among Central European species, the spermatheca of A. pseu- dogregaria is most similar to that of Palaearctic A. gregaria but in the latter the apex is distally truncate, median lobe is distinctly sinuate and only weakly projected ventrad, the abdomen is darker and not distinctly bicolored, and the microsculpture of the forebody is much stronger, creating a dull reflection. Description. Body length 2.4–2.7 mm, moderately flattened (stronger so on elytra), narrowly subparallel, colour of head, pronotum, scutellar region of elytra, apical part of abdomen and antennomeres 5–11 dark brown to dark reddish brown, elytra and antennomeres 1–3 paler, red-brown and legs yellow; forebody finely and densely punctate, microsculpture shallow, consisting of meshes; head slightly elongate and with small, shallow impression medially, head slightly narrower than pronotum, postocular region elongate, ca. as long as maximum diameter of eye, tempora with carinae dorsally only; antennae slender, as long as pronotum and elytra combined, ba- sal three antennomeres strongly elongate, 4 subquadrate, 5–10 subquadrate to slightly transverse, and terminal one strongly elongate and ca. as long as two preceding anten- nomeres 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, hu- meri 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 emargina- tion, 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 mod- erately projecting ventrad apically in lateral view (Fig. 28B). FEMALE. Tergite VIII rounded apically and slightly pointed medially (Fig. 28F); sternite VIII rounded api- cally (Fig. 28G); spermatheca S-shaped, capsule pitcher-shaped with short neck, stem strongly sinuate and swollen apically (Fig. 28C). Distribution. Origin: Nearctic. Canada: ON. United States: VA. Bionomics. This species has only been collected by passive traps, including malaise and pitfall traps. All specimens have been collected from at least partly disturbed habi- tats, such as forest edges, agricultural fields, and suburban environments. This species corresponds to ‘Aleocharinae sp. 5’ in Brunke et al. (2014), which was collected in both soybean fields and adjacent forest edges by pitfall traps. Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)78 Comments. Aloconota pseudogregaria is probably broadly distributed in northeast- ern North America. We have compared the male and female genitalia of A. pseudogre- garia with all Central European and Nearctic species of Aloconota, and are confident that this taxon has not been previously described from Europe or North America, despite its occurrence in disturbed habitats in North America, which is typical for in- troduced species. Although Aloconota pseudogregaria clustered most closely with A. gre- garia (BOLD:ABU6164) in our barcode dataset, its BIN is ~ 8% different from that of the latter. Based on morphology of the aedeagus and spermatheca, Aloconota pseudogre- garia is probably even more closely related to East Palaearctic Aloconota described from Japan and Korea (e.g., Sawada 1970 [as Tomoglossa], Lee and Ahn 2017) rather than to A. gregaria. However, the described species all differ markedly in external morphology. Athetini Casey, 1910 Atheta (Datomicra) nigra (Kraatz, 1856) BOLD:ACO4408 Fig. 29A–G Material (DNA-barcoded specimens). Canada: Ontario: Peterborough, 44.253N, 78.415W, farm, malaise trap, 24–30.V.2015, B. McClenaghan (1, CBG). Germany: Koeln-Worringen, Worringer Bruch, 51.044, 6.87427, 01.VII.2010, F. Koehler & J. Koehler (1, ZSM). Distribution. Origin. Palaearctic (adventive in North America). Canada: ON [new record], SK. Bionomics. Canadian specimens have been collected on farmland and directly from horse manure. Comments. Atheta nigra is a Palaearctic species reported from across Europe, Eu- ropean Russia, Kazakhstan, North Korea and southern China (Newton 2019). It is adventive in North America and New Zealand (Newton 2019) and is here newly re- ported from Ontario. The new record from Ontario indicates that this species is far more widely distributed in North America than previously known. Mocyta scopula (Casey, 1893), comb. nov. BOLD:ACH8720 Fig. 30A–F Eurypronota scopula Casey, 1893 Pancota laetabilis Casey, 1906 Dolosota abundans Casey, 1910 Dolosota flaccida Casey, 1910 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ABU6164 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACO4408 http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ACH8720 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 79 Figure 29. Atheta (Datomicra) nigra (Kraatz) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in ventral view D spermatheca E apical part of dorsal male abdomen showing 4 dents on tergite VIII F female tergite VIII G female sternite VIII; A, D, F, G after Kli- maszewski 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). Adam J. Brunke et al. / ZooKeys 1041: 27–99 (2021)80 Dolosota redundans tergina Casey, 1910 Dolosota scopula: Casey (1910) (as type species of Dolosota) Dolosota secunda Casey, 1910 Dolosota sequax Casey, 1910 Acrotona (Dolosota) abundans: Seevers (1978) (as valid species) Acrotona (Dolosota) flaccida: Seevers (1978) (as valid species) Acrotona (Dolosota) scopula: Seevers (1978) (as valid species) (Dolosota syn. of Acrotona, in part; some species moved to Pancota) Acrotona (Dolosota) secunda: Seevers (1978) (as valid species) Acrotona (Dolosota) sequax: Seevers (1978) (as valid species) Pancota laetabilis: Seevers (1978) (as valid species) Pancota redundans tegrina: Seevers (1978) (implied, subspecies not directly mentioned) Acrotona abundans: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b)) Acrotona flaccida: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b)) Acrotona laetabilis: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b)) Acrotona redundans tergina: Majka and Sikes (2009) (syn. of A. scopula following Gusa- rov (2003b)) Acrotona scopula: Majka and Sikes (2009) (valid species following Gusarov (2003b)) Acrotona secunda: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b)) Acrotona sequax: Majka and Sikes (2009) (syn. of A. scopula following Gusarov (2003b)) Material (DNA-barcoded specimens). Canada: Ontario: Georgian Bay Islands Na- tional Park, Fairy Lake, 44.8929, -79.8514, mostly conifer forest with moss, Berlese funnel, 5.VIII.2015, BIObus 2015 (1, CBG). Distribution. Origin. Nearctic. Canada: ON [new record]. United States: IA, MO, MS, NY, PA, RI. Diagnosis. Mocyta scopula can be distinguished from bicolored Canadian species and paler specimens of M. fungi by its finely punctate pronotum that is almost as wide as the elytra and ca. as long, and the distinctly transverse antennomeres 6–10 (Fig. 30A). The barcode sequences of M. scopula forms a sister cluster with M. luteola (BOLD:ABW2813), with a sequence divergence of ~ 7.5%. These species can be easily separated using the above diagnosis. Bionomics. The Canadian specimen was collected from forest litter with a Berlese funnel but nothing specific is known about this species’ microhabitat preferences. Comments. Mocyta scopula is a native Nearctic species distributed in eastern North America. Here we newly report it from Canada based on one male specimen collected in southern Ontario. Its distribution in the United States is based on type material, including its putative synonyms, which should be verified. Mocyta scopula is the type species of Dolosota Casey, which has been treated as a subgenus of Acrotona since Seevers (1978). However, using the generic concepts of Kli- maszewski et al. (2015), this species best fits in genus Mocyta based on the following http://boldsystems.org/index.php/Public_BarcodeCluster?clusteruri=BOLD:ABW2813 Integrative taxonomy of Nearctic and Palaearctic Aleocharinae 81 Fi