Research Article |
Corresponding author: Adam J. Brunke ( adam.j.brunke@gmail.com ) Academic editor: Volker Assing
© 2021 Adam J. Brunke, Mikko Pentinsaari, Jan Klimaszewski.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
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
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A long tradition of separate Nearctic and Palaearctic taxonomic studies of the diverse aleocharine rove beetles (Coleoptera: Staphylinidae) has obscured the recognition of Holarctic species and detection of adventive species in both regions. Recently, integrated study of the two regions through detailed morphological comparisons and development of an authoritatively identified DNA barcode reference library has revealed the degree to which these two aleocharine faunas are interconnected, both naturally and through human activity. Here this approach is adopted to recognize new species, reveal Holarctic species, and recognize adventive species in both North America and Europe. The following new species are described: Isoglossa triangularis Klimaszewski, Brunke & Pentinsaari, sp. nov. from British Columbia; Gnypeta impressicollis Klimaszewski, Brunke & Pentinsaari, sp. nov., from Ontario, Maryland and North Carolina; Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov., from Ontario and Virginia; and Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. from eastern Canada. Dasygnypeta velata and Philhygra angusticauda are revealed to be Holarctic species, resulting in the following synonymies: Dasygnypeta velata (Erichson, 1839) = Gnypeta minuta Klimaszewski & Webster, 2008, syn. nov. and Philhygra angusticauda (Bernhauer, 1909) = Atheta (Philhygra) pinegensis Muona, 1983, syn. nov. The Nearctic species Hylota ochracea (and genus Hylota), Thecturota tenuissima, and Trichiusa robustula are newly reported from the Palaearctic region as adventive, resulting in the following synonymies: Hylota ochracea Casey, 1906 = Stichoglossa (Dexiogyia) forticornis Strand, 1939, syn. nov.; Thecturota tenuissima Casey, 1893 = Atheta marchii Dodero, 1922, syn. nov.; and Trichiusa robustula Casey, 1893 = T. immigrata Lohse, 1984, syn. nov. The Palaearctic species Amarochara forticornis, Anomognathus cuspidatus, Oligota pumilio, and Parocyusa rubicunda are newly confirmed from the Nearctic region as adventive, resulting in the following synonymies: Parocyusa rubicunda (Erichson, 1837) = Chilopora americana Casey, 1906, syn. nov. and Anomognathus cuspidatus (Erichson, 1839) = Thectura americana Casey, 1893, syn. nov. The genus Dasygnypeta, sensu nov. is newly reported from North America, Paradilacra is newly reported from eastern North America, and Haploglossa is newly reported from Canada, resulting in the following synonymy: Paradilacra densissima (Bernhauer, 1909) = Gnypeta saccharina Klimaszewski & Webster, 2008, syn. nov. Native Cyphea wallisi is newly reported from across Canada and C. curtula is removed from the Nearctic fauna. The status of both Gyrophaena affinis and Homalota plana is uncertain but these species are no longer considered to be adventive in North America. Three new combinations are proposed: Dasygnypeta baranowskii (Klimaszewski, 2020) and D. nigrella (LeConte, 1863) (both from Gnypeta) and Mocyta scopula (Casey, 1893) (from Acrotona). Dolosota Casey, 1910, syn. nov. (type species Eurypronota scopula Casey), currently a subgenus of Acrotona, is therefore synonymized with Mocyta Mulsant & Rey, 1874. Additionally, four new Canadian records and 18 new provincial and state records are reported.
Canada, DNA barcodes, faunistics, morphology, North America, rove beetles, United States
Historically, taxonomic research on the hyperdiverse aleocharine rove beetle (Coleoptera: Staphylinidae) faunas of North America and better-known Europe has been conducted separately, with a few exceptions (e.g., Klimaszewski et al. 1979). More recently, a closer examination of Aleocharinae in these two regions has demonstrated that a number of species are shared between the Nearctic and Palaearctic, either naturally (Holarctic) or through human activity (adventive) (e.g.,
In combination with careful morphological study, large-scale DNA barcoding (e.g.,
Here we broadly compare morphological and molecular data across the Nearctic and West Palaearctic Aleocharinae in order to better integrate the taxonomic knowledge of these two regions. We describe four new Nearctic species, propose revised generic concepts, report new distributional records, and propose a number of new synonyms that impact our understanding of Holarctic and adventive species.
Almost all specimens used in this study were dissected and their genitalia were subsequently examined on microslides. The genital structures were dehydrated in absolute ethanol, mounted in Canada balsam on celluloid microslides, and pinned with the specimens from which they originated. The photographs of the entire body and the genital structures were taken using an image processing system (Nikon SMZ 1500 stereoscopic microscope; Nikon Digital Camera DXM 1200F) and processed in Adobe Photoshop. Terminology mainly follows that used by
Depository abbreviations:
cRW Personal collection of Reginald P. Webster, Charters Settlement, New Brunswick, Canada (also known as RWC);
LFC Laurentian Forestry Centre, Québec, Quebec, Canada;
MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, United States (C. Maier);
NHMD Natural History Museum of Denmark, Copenhagen University, Copenhagen, Denmark (A. Solodovnikov);
We have examined all DNA barcode data for Aleocharinae previously generated by a variety of projects in both Europe and North America (e.g.,
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;
The DNA barcode sequences studied here, including both previously unpublished data and the sequences published in earlier studies, have been compiled into a publicly available dataset on BOLD (DS-ALEO2020, https://doi.org/10.5883/DS-ALEO2020) along with collecting data, images of the specimens (if available), and other metadata related to the specimens and sequences. The sequences are also available through GenBank (accessions provided in Suppl. material
(DNA barcoded specimens). Canada: Ontario: Fergus, Centre Wellington District High School, 43.704, -80.358, Malaise trap, 3.V.2013, M. Cottrill (1,
Origin: Palaearctic (adventive in Nearctic). Canada: ON [new record].
Amarochara forticornis may be easily recognized among the other Canadian species of the genus by the distal antennomeres, which are less than twice as wide as long. The species is also unique within the genus by having a distinct basal impression on abdominal tergite VI.
In its native range, A. forticornis occurs in a variety of open and forested habitats, including forests, edges of waterways, grasslands, agricultural fields, and gardens (
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 (
1A | Antennomere 10 only weakly transverse (Fig. |
Amarochara forticornis (Lacordaire) |
– | Antennomere 10 strongly transverse, at least twice as wide as long (native species); abdominal tergite VI with, at most, coarse punctures at base | 1B |
1B | Pronotum with strong microsculpture and coarse, dense punctation, surface almost matte | A. duryi (Casey) |
– | Pronotum without microsculpture or with fine microsculpture, and with fine sparse to moderately dense punctation, surface moderately to highly glossy | 2 |
(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,
Origin: Nearctic. Canada: ON [new record]. United States: OK, PA.
Haploglossa nebulosa may be easily distinguished from the other Nearctic species of the genus, H. barberi (Fenyes), by the bicolored elytra and fusiform body (
All members of Haploglossa are nidicolous, mostly in bird nests but also in mammal and ant nests (summarized by
The genus Haploglossa and H. nebulosa are newly reported from Canada, from a single locality in southern Ontario. The species is also known from Oklahoma and Pennsylvania, United States (
Haploglossa nebulosa (Casey) A habitus B median lobe of aedeagus in lateral view (adapted from
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 |
(DNA barcoded specimens). Canada: Ontario: Guelph, Hanlon Preservation Park, 43.506, -80.213, mixed forest, dead wood and beating, 11.VI.2017, M. Pentinsaari (1,
Origin: Nearctic. Canada: AB, NB, NF [new record], ON [new record].
Little is known about the microhabitat preferences of this species, but it likely occurs in in nests or cavities within trees as does H. ochracea (
This recently described species, previously known from New Brunswick and Alberta (
Stichoglossa (Dexiogyia) forticornis Strand, 1939, syn. nov.
(DNA barcoded specimens). Canada: Ontario: Guelph, Dovercliffe Road, 43.51, -80.254, backyard, compost and mouldy hay piles, 6.VI.2018, M. Pentinsaari (3,
Origin: Nearctic (adventive in Europe). Canada: NB, NS, NT, ON, QC, SK. United States: NY, VT.
Hylota ochracea is strongly associated with bird nests in forested habitats. It has also been collected from artificial analogs such as a pigeon coup, manmade nest boxes, and a plastic composter bin containing carrion and decaying vegetables (
Hylota ochracea, a widespread Nearctic species (
With the above synonymy, the genus Dexiogyia is now known only from externally similar sister species D. angustiventris (Casey) (Nearctic) and D. corticina (Erichson) (West Palaearctic), plus Afrotropical D. congoensis (Scheerpeltz). As in the former D. forticornis, D. congoensis is probably misplaced due to superficial similarity. Hylota is readily separated from Dexiogyia by the shape of the pronotum, which is strongly convergent anteriad, such that its apical width is subequal to the width of the head. In Dexiogyia, the head is distinctly narrower than the pronotum.
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
Rheobioma
Casey, 1906;
Athetalia
Casey, 1910 (in part);
Neoisoglossa
Klimaszewski & Pelletier, 2004;
In
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 (
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.
Origin: Nearctic. Canada: BC.
Isoglossa triangularis can be easily distinguished from all Nearctic species of the Ocalea group of genera by a combination of the strongly transverse and sparsely punctate pronotum, transverse antennomere 4, distinct triangular apex of the median lobe in lateral view (Fig.
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).
Body length 3.0–3.3 mm, dark brown with elytra, antennomeres 1–2 or 1–3, legs and apical part of abdomen yellow-brown, forebody moderately glossy and abdomen strongly so (Fig.
The specimens were collected in a Malaise trap on an open field surrounded by mixed forest.
Based on a combination of small size (< 4.5 mm), superficial, meshed microsculpture, sparse pronotal punctation, with punctures separated by more than two puncture diameters, pronotum transverse, shorter and narrower than elytra, and the transverse antennomeres 5–10, I. triangularis keys to genus Isoglossa Casey in
Tachyusa rubicunda Erichson, 1837
Chilopora americana Casey, 1906, syn. nov.
Tetralaucopora americana:
Parocyusa americana:
(DNA barcoded specimens). Austria: Innervillgraten, Arntal, 46.8362, 12.3348, 1580 m, 22.VIII.2010, F. Koehler and J. Koehler (2,
Canada: Ontario: Ancaster, 21.X.1967 (1,
Origin: West Palaearctic (adventive in North America). Canada: BC, ON, QC, NB, NF. United States: CT, NY, PA.
In North America, most specimens of this species have been collected from near water, including a sandy creek bank, in a dried streambed and in moss near the splash zone of a waterfall (
Parocyusa rubicunda is a widespread West Palaearctic species (Europe, European Russia, Turkey, Georgia, Iran, Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan) (
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 (
Recently,
Tribe Tachyusini C.G. Thomson
Adapted from
1 | Elytra at humerus only slightly broader than pronotum at base (Figs |
2 |
– | Elytra at humerus distinctly broader than pronotum at base (Figs |
3 |
2 | Pronotum with pubescence directed straight posteriad; hind tarsus subequal in length to hind tibia or longer (Fig. |
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. |
Paradilacra Bernhauer |
3 | Abdomen clavate, at base distinctly narrower than head (Fig. |
Tachyusa Erichson |
– | Abdomen at most slightly constricted at base, subequal to or wider than head (Figs |
4 |
4 | Abdomen at base elongate and moderately constricted, ca. as wide as head (Figs |
Dasygnypeta Lohse, sensu nov. |
– | Abdomen at base at most slightly constricted, wider than head (Fig. |
Gnypeta Thomson |
Atheta (Paradilacra) densissima Bernhauer, 1909
Gnypeta saccharina Klimaszewski & Webster, 2008, syn. nov.
(DNA barcoded specimens). Canada: Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane forest, 27.VI.2012, BIOBus 2012 (1,
Origin: Nearctic. Canada: AB, BC, NB, ON [new record], SK [new record]. United States: CA, MT, NV, ND, OR, UT.
This species has been collected from various wetland microhabitats including the edges of lakes, rivers, and a beaver pond (
Paradilacra densissima (Bernhauer) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after
Paradilacra densissima and the genus Paradilacra, widespread in western and central North America (
In his key to the genera of Tachyusini,
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
Gnypeta baranowskii Klimaszewski, 2020
Origin: Nearctic. Canada: BC.
The type series was collected by sifting litter (
We here transfer this species to Dasygnypeta sensu nov. based on morphology illustrated by
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
Tachyusa nigrella LeConte, 1863
Gnypeta nigrella:
(DNA barcoded specimens). Canada: New Brunswick: York Co., Fredericton at St. John River, 45.959, -66.625, margin of river in drift (mostly maple seeds), 4.VII.2004, R.P. Webster (1, LFC).
Canada: Manitoba: 5 miles SW of Shilo, 5.VI.1958, J.F. McAlpine (1,
Origin: Nearctic. Canada: MB [new record], NB, NF, ON, QC [new record]. United States: IL, MA, MD, NJ, NY, PA, VT, WV.
Collected along the edge of a variety of running and standing water-based habitats.
We here transfer this species to Dasygnypeta sensu nov. based on morphology and close clustering of DNA barcode sequences with D. velata. Dasygnypeta nigrella is a widespread species in eastern North America and is here newly reported from Manitoba and Quebec.
Dasygnypeta nigrella (LeConte) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in dorsal view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after
Homalota velata Erichson, 1837
Gnypeta minuta Klimaszewski & Webster, 2008, syn. nov.
(DNA barcoded specimens). Germany: Thuringia, Ufergehoelze am Speicher Loessau, 50.5665, 11.894, 460 m, 1.I.2013, GBOL-Team
Canada: Manitoba: 5 miles SW of Shilo, 5.VI.1958, J.F. McAlpine (2,
Origin: Holarctic. Canada: MB [new record], NB, NF, NT, SK. United States: AK.
Nearctic specimens have been collected most frequently along the margins of running water but also along the margins of a forest pool (
Dasygnypeta velata is newly reported from North America and was previously known in the Nearctic region under the synonym Gnypeta minuta (
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.
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
Holotype. (male,
The species epithet refers to the longitudinal impression on the pronotum, most strongly developed in males.
Gnypeta impressicollis can be easily distinguished from all Nearctic species of the genus (except eastern G. baltifera (LeConte)) by the hexagonal pronotum with a longitudinal impression in the basal half (females) to nearly entire pronotal length (males). Males also have an impression on the vertex of the head. We have examined the female type of G. baltifera and it is externally similar but differs by the shorter, less angulate hexagonal pronotum, reddish and longer elytra and spermatheca with an elongate stem (C-shaped in G. impressicollis).
Body length 3.2–3.4 mm; colour dark brown, elytra brown with irregular rust-brown patches, first two or three basal tergites rust-brown with posterior edge yellow, apex of abdomen rust-brown, legs and antennae rust-brown; integument highly glossy (Fig.
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).
Origin: Nearctic. Canada: ON. United States: MD, NC. Gnypeta impressicollis is probably broadly distributed in eastern North America.
Specimens were collected by sifting leaf litter along a lake margin and by treading pond vegetation.
It was challenging to place this species in either Gnypeta or Ischnopoda Stephens based on the concepts of
(DNA barcoded specimens). Germany: Bornheim-Hemmerich, Ortslage, 50.7596, 6.93151, 30.VII.2010, F. Koehler (2,
Origin: West Palaearctic (adventive in North America). Canada: NB, ON [new record], PE. United States: CA, MA, MO, NV, TX.
This species is generally found in anthropogenic habitats, including compost, dung, and old hay and grass (
Oligota parva is a cosmopolitan species that is adventive in Canada. Here we newly report it from Ontario.
(DNA barcoded specimens). Belgium: Blanden, BR Meerdaalboos, 50.7976, 4.71622, 8.V.2010, F. Koehler (1,
Origin: West Palaearctic (adventive in North America). Canada: AB [new record]. United States (all except MT need verification): DC, IL, OH, MT [new record].
Among Canadian species of Oligota, O. pumilio is extremely similar to O. pusillima in the narrow, parallel body (Fig.
Oligota pumilio Kiesenwetter A habitus (image by A. Bogri – www.BilleBank.dk) B median lobe of aedeagus in lateral view (drawn from
This species occurs in a wide variety of habitats across a broad elevational range, including hollow trees, plant debris, old hay in cattle barns, moldy substrates and in mushrooms (
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) (
(DNA barcoded specimens). Finland: Oba: Oulu, Hietasaari, 65.0225, 25.4247, 22.IV.2011, M. Pentinsaari (1,
Ontario: Ottawa, Ottawa River, Deschênes Lookout, Berlese flood debris, 1.V.1985, A. Davies (1,
Origin: West Palaearctic (adventive in North America). Canada: AB [new record], NB, ON [new record]. United States: MA, NY.
This species occurs in a variety of moist to dry, decaying organic matter including rotting hay, compost, hollow trees, and ant nests (
Oligota pusillima is a Palaearctic species that has been introduced to North America, South America, Australia, Africa, and southeast Asia (
Non-sequenced specimens. Canada: Manitoba: Winnipeg, under bark of rotten ‘N. aceroides’ [= Acer negundo], 27.VIII.1918, J.B. Wallis (2,
Origin: Nearctic. Canada: AB, MB [new record].
The MB specimens were collected under bark, confirming that this species lives in a way similar to other members of the genus.
Anomognathus athabascensis, recently described from Alberta (
Anomognathus athabascensis Klimaszewski, Hammond & Langor A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after
Homalota cuspidata Erichson, 1839
Thectura americana Casey, 1893, syn. nov.
Anomognathus americanus:
Homolota cuspidata Erichson, 1839. Lectotype, male, here designated (
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.
Thectura americana Casey, 1893, syn. nov. Holotype (male) (
Male tergite VIII of A–C Anomognathus cuspidatus (Erichson) and D potential undescribed species, in dorsal (top row) and lateral (middle and bottom rows) A lectotype of A. cuspidatus (‘Europe’) B holotype of A. americanus (Casey) (= A. cuspidatus) C, D sequenced, non-types (Finland). Scale bar: 0.2 mm.
(sequenced specimens indicated in square brackets). Canada: Alberta: Peace River, 25 km NW Peace River, 17–23.VIII.1993, J. Hammond (2,
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,
Germany: Nationalpark Mueritz, Babke-Zartwitz-Speck-Schwarzenhof, 53.4125, 12.8463, car net, 20.VI.2015, GBOL-Team
Putative undescribed Anomognathus (corresponding to BIN BOLD:ACA9191):
Finland (all
Anomognathus cuspidatus is distinctive for its trident-shaped apex of male and female tergite VIII (Fig.
Origin: West Palaearctic (adventive in North America). Canada: AB, NB, ON. United States: NY.
This species occurs under the bark of dead trees. One specimen (NB) was collected from a Lindgren funnel.
Anomognathus cuspidatus is a widespread West Palaearctic species that is known from Europe, European Russia and Algeria (
After the results of the present study, two species of Anomognathus are known to occur in North America: native A. athabascensis Klimaszewski, Hammond & Langor and the adventive A. cuspidatus. These are easily separated by the drastically different shapes of male and female tergites VIII (Figs
Cyphea wallisi Fenyes, 1921
Agaricomorpha vincenti Klimaszewski & Webster, 2016, syn. nov.
Agaricomorpha vincenti:
Cyphea wallisi Fenyes, 1921. Paratype, male (MCZ). Winnipeg, Man. [handwritten label] / Wallis [handwritten label] / 25490. / Cyphea, Wallisi, Feny [handwritten label] / Type., 9989, 9983 [typed red label].
Agaricomorpha vincenti Klimaszewski & Webster, 2016, syn. nov. Holotype, male (LFC). Canada, New Brunswick, Carleton Co., Jackson Falls, “Bell Forest”, 46.2200°N, 67.7231°W, 7–21.VI.2012, C. Alderson & V. Webster, coll. [white typed label] / Rich Appalachian hardwood forest, Lindgren funnel trap in canopy of Fagus grandifolia [white typed label] / Holotype Agaricomorpha vincenti Klimaszewski & Webster, 2016 [red typed label] / Cyphea curtula (Erichson) det. Klimaszewski 2017 [white typed label] / Cyphea wallisi Fenyes det. A. Brunke 2020.
The aedeagi of the male paratype (holotype in collection of the California Academy of Sciences) of C. wallisi and holotype of A. vincenti are identical and both differ from that of Palaearctic Cyphea curtula (image by V. Assing) by the broader distal lobe in lateral view, which only slightly extends beyond the distal plate (Fig.
(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,
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,
Origin: Nearctic. Canada: AB, MB, NB, NS [new record], ON [new record], QC.
Specimens have been collected in Malaise traps, window traps and Lindgren funnels placed in forests. Both the closely related West Palaearctic C. curtula and C. latiuscula Sjöberg have been consistently collected under bark, where they occur in the larval burrows of various longhorn beetles (Cerambycidae), bark beetles (Curculionidae: Scolytinae) and the carpenter moth (Cossus L.) (
Cyphea wallisi is a broadly distributed native Nearctic species, reported from AB east to NS. Here we treat Nearctic records of Cyphea as C. wallisi (previously treated as Palaearctic C. curtula, e.g.,
Sequenced Nearctic specimens of Cyphea from ON and NS formed a barcode cluster that was nearly 5% divergent from those of Palaearctic specimens of C. curtula (BOLD:AAO1175, one published sequence record from Belgium and three unpublished records from the Netherlands). Northern European C. latiuscula, the only other species of the genus, has a broader body outline, different male genitalia and is quite differently colored (bicolored pronotum and pale elytra). No barcode sequence data are currently available for C. latiuscula. Based on the study of one paratype of C. wallisi, described from Manitoba and not reported since, it was discovered that Nearctic specimens of Cyphea correspond to this species and differ from Palaearctic C. curtula by the broader distal lobe of the median lobe of the aedeagus in lateral view, which only slightly extends beyond the distal plate (Fig.
(DNA barcoded specimens). Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 135 m, 16.VI.2010, F. Koehler (1,
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.
Gyrophaena affinis is newly reported from AB based on barcoded material.
Sequenced Nearctic specimens from ON, AB, NB, and QC form a distinct barcode cluster, separate from all sequenced Palearctic specimens and divergent by 4.65%. This pattern is inconsistent with a species that is adventive in North America and we remove G. affinis from the list of adventive species in Canada. In comparing images between those of Nearctic specimens (Fig.
(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,
Origin. Nearctic. Canada: NB, QC [new record]. United States: WI.
Specimens have been collected from a partly dried Pleurotus mushroom, from within the pores of a Trametes polypore, and from the nest contents of a Barred owl (Strix varia Barton) (
The new record from QC, near the ON border, bridges the wide gap between previous records in NB and WI.
Gyrophaena gracilis Seevers A habitus B, C median lobe of aedeagus in lateral view D spermatheca E male tergite VIII F male sternite VIII G female tergite VIII H female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–H). Illustrations after
(DNA barcoded specimens). Canada: Ontario: Hartington, Eel Lake Cottage, 44.563, -76.549, deciduous forest, mushrooms, 4.X.2017, M. Pentinsaari (2,
Origin. Nearctic. Canada: ON [new record]. United States: IL, MD, PA.
Gyrophaena simulans is extremely similar to G. criddlei and G. pseudocriddlei but has a slightly more transverse and flatter pronotum, with straighter apical and basal margins, and differently shaped upper process of the median lobe in lateral view (Fig.
The Canadian specimens were collected by sifting mushrooms in a deciduous forest. No detailed data on the host fungus were recorded.
Gyrophaena simulans is a native Nearctic species distributed in eastern North America and is newly reported from Canada. The barcode cluster BOLD:ACY8004 also contains specimens identified as related species G. criddlei (female) and G. pseudocriddlei but more research, with broader sampling of sequenced, identified males, is needed to determine whether these species share a BIN or these specimens are misidentified. As we were unable to verify the identifications at this time, these records are not published here.
(DNA barcoded specimens). Belgium: Sint-Genesius-Rode, BR Zonienwoud, 50.7505, 4.423, 28.IV.2010, F. Koehler (1,
Origin. Uncertain. Canada: AB, MB, NB, NF, NS, ON. United States: AZ, CA, CO, IA, ID, IN, MT, NY, OH, PA, TX.
Specimens occur under bark of dead trees.
Sequenced Nearctic specimens from ON form a distinct barcode cluster, separate from all sequenced Palearctic specimens and divergent by 7.58%. This pattern is inconsistent with a species adventive in North America and we remove H. plana from the list of adventive species in Canada. Preliminary comparisons between images of Palaearctic and Nearctic specimens revealed that there may be some slight differences in the shape of the spermatheca. More research is needed to determine the status of the Nearctic and Palaearctic populations, though the level of genetic divergence between discrete Nearctic and Palaearctic populations suggests that two sister species are involved.
Thecturota tenuissima Casey, 1893
Atheta marchii Dodero, 1922, syn. nov.
Pragensiella magnifica Machulka, 1941, syn. nov.
Thecturota marchii:
Thecturota magnifica:
(DNA-barcoded specimens). Germany: Kobern-Gondorf, Ortslage/Weinberge, 50.308, 7.460, 21.V.2010, F. Koehler (1,
Several males and females of T. tenuissima from Denmark (NMHD) were compared with illustrations from
Origin. Nearctic (adventive in West Palaearctic). Canada: ON, QC. United States: RI.
Canadian specimens were collected by car-netting in mixedwood forests, while Palaearctic specimens are known from compost and other plant-based debris (
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 (
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 (
Holotype
(male) (
(barcoded specimens). Canada: Ontario: Guelph, 25 Division St., 43.554, -80.264, Malaise trap, 14.VII.2010, A. Smith (1,
The species epithet refers to the similarity to related species A. gregaria (Erichson), which was originally treated separately from other Aloconota under subgenus Glossola Fowler (e.g., Benick 1954) because it lacks obvious male secondary sexual characters.
Aloconota pseudogregaria can be easily distinguished from all other species of the genus occurring in eastern North America by the distinctly bicolored abdomen (Fig.
Aloconota pseudogregaria Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C spermatheca D male tergite VIII E male sternite VIII (structure accidentally over-cleared in balsam preparation) F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G).
Body length 2.4–2.7 mm, moderately flattened (stronger so on elytra), narrowly subparallel, colour of head, pronotum, scutellar region of elytra, apical part of abdomen and antennomeres 5–11 dark brown to dark reddish brown, elytra and antennomeres 1–3 paler, red-brown and legs yellow; forebody finely and densely punctate, microsculpture shallow, consisting of meshes; head slightly elongate and with small, shallow impression medially, head slightly narrower than pronotum, postocular region elongate, ca. as long as maximum diameter of eye, tempora with carinae dorsally only; antennae slender, as long as pronotum and elytra combined, basal three antennomeres strongly elongate, 4 subquadrate, 5–10 subquadrate to slightly transverse, and terminal one strongly elongate and ca. as long as two preceding antennomeres combined; pronotum slightly transverse (width/length ratio 1.3), trapezoidal in shape, flattened, pubescence directed straight posteriad in central part of disc and obliquely posteriad laterally; elytra at suture ca. as long as pronotum along midline, flat, distinctly transverse (width/length ratio 1.5), ~ 1/3 broader than pronotum, humeri angular, posterior margins slightly sinuate laterally, pubescence directed straight posteriad forming slightly arcuate lines in sutural region of disc; abdomen subparallel, tergites III–VI distinctly impressed at base; basal metatarsomere ~ 1/3 longer than the following one. MALE. Tergite VIII rounded apically with minute median emargination, lacking apical teeth (Fig.
Origin: Nearctic. Canada: ON. United States: VA.
This species has only been collected by passive traps, including malaise and pitfall traps. All specimens have been collected from at least partly disturbed habitats, such as forest edges, agricultural fields, and suburban environments. This species corresponds to ‘Aleocharinae sp. 5’ in
Aloconota pseudogregaria is probably broadly distributed in northeastern North America. We have compared the male and female genitalia of A. pseudogregaria with all Central European and Nearctic species of Aloconota, and are confident that this taxon has not been previously described from Europe or North America, despite its occurrence in disturbed habitats in North America, which is typical for introduced species. Although Aloconota pseudogregaria clustered most closely with A. gregaria (BOLD:ABU6164) in our barcode dataset, its BIN is ~ 8% different from that of the latter. Based on morphology of the aedeagus and spermatheca, Aloconota pseudogregaria is probably even more closely related to East Palaearctic Aloconota described from Japan and Korea (e.g.,
(DNA-barcoded specimens). Canada: Ontario: Peterborough, 44.253N, 78.415W, farm, malaise trap, 24–30.V.2015, B. McClenaghan (1,
Origin. Palaearctic (adventive in North America). Canada: ON [new record], SK.
Canadian specimens have been collected on farmland and directly from horse manure.
Atheta nigra is a Palaearctic species reported from across Europe, European Russia, Kazakhstan, North Korea and southern China (
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
Eurypronota scopula Casey, 1893
Pancota laetabilis Casey, 1906
Dolosota abundans Casey, 1910
Dolosota flaccida Casey, 1910
Dolosota redundans tergina Casey, 1910
Dolosota scopula:
Dolosota secunda Casey, 1910
Dolosota sequax Casey, 1910
Acrotona (Dolosota) abundans:
Acrotona (Dolosota) flaccida:
Acrotona (Dolosota) scopula:
Acrotona (Dolosota) secunda:
Acrotona (Dolosota) sequax:
Pancota laetabilis:
Pancota redundans tegrina:
Acrotona abundans:
Acrotona flaccida:
Acrotona laetabilis:
Acrotona redundans tergina:
Acrotona scopula:
Acrotona secunda:
Acrotona sequax:
(DNA-barcoded specimens). Canada: Ontario: Georgian Bay Islands National Park, Fairy Lake, 44.8929, -79.8514, mostly conifer forest with moss, Berlese funnel, 5.VIII.2015, BIObus 2015 (1,
Origin. Nearctic. Canada: ON [new record]. United States: IA, MO, MS, NY, PA, RI.
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.
The Canadian specimen was collected from forest litter with a Berlese funnel but nothing specific is known about this species’ microhabitat preferences.
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
The aedeagus, coloration and punctation of the Canadian specimen are consistent with type material of M. scopula, previously examined and imaged by JK. The two other members of the BIN BOLD:ACH8720 originate from a study by
2a | Pronotum much broader than elytra; antennal articles 5–10 in specimens slightly elongate; spermatheca forming concentric circles posteriorly | M. discreta (Casey) |
– | Pronotum ca. as broad as elytra or slightly narrower (Fig. |
2b |
2b | Pronotum coarsely punctate and extremely transverse with weakly rounded base and apex; antennal articles 5–10 subquadrate; median lobe in lateral view strongly produced ventrad | M. luteola (Erichson) |
– | Pronotum finely punctate and transverse, but more rounded at base and apex (Fig. |
M. scopula (Casey) |
Atheta (Metaxya) angusticauda Bernhauer, 1909
Atheta (Philhygra) pinegensis Muona, 1983, syn. nov.
(DNA barcoded specimens). Canada: Alberta: Jasper National Park, Miette Hotsprings, 53.124, -117.7755, Malaise trap placed in valley with creek bed, sides rocky and mossy, 1439 m, 21.VII.2012, BIObus 2012 (1,
Origin. Holarctic. Canada: AB[new record], BC, NB. United States: AK, NH.
As with other species of the genus, P. angusticauda is associated with riparian habitats.
Philhygra angusticauda is a Holarctic species that was previously recognized in the Palaearctic (Finland, Norway, European Russia, Russian Far East) (
Philhygra angusticauda (Bernhauer) A habitus B median lobe of aedeagus in lateral view C female pygidium D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after
(non-sequenced specimens). Canada: Ontario: Algonquin Park, ~45.87, -77.33, car net, 20.VII.2016, T. Struyve (10,
Origin. Nearctic. Canada: ON [new record]. United States: MA, RI.
This species can be readily recognized by a combination of its small size, large eyes and relatively simple, ventrally projecting median lobe of the aedeagus in lateral view (Fig.
Nothing specific is known about this species’ microhabitat preferences but it probably occurs near water as do other species of the genus. The series of Ontario specimens was collected using a car net, which is typically effective for collecting small staphylinids.
Philhygra finitima is a native Nearctic species distributed in northeastern North America. Here, we newly report it from Canada. Canadian specimens were identified based on comparison with images (Fig.
(DNA-barcoded specimens). Canada. Alberta: Waterton Lakes National Park, Highway 6 pulloff, 49.065, -113.779, 1569 m, intercept trap, montane forest, 21–27.VI.2012, BIOBus 2012 (1,
Origin. Nearctic. Canada: AB [new record], BC. United States: AK, WA.
Philhygra laevicollis can be distinguished from most species of the genus by the general shape of the median lobe in lateral view. It is most similar to P. pseudolaevicollis but has a sinuate ventral face of the median lobe in lateral view and large spines in the internal sac (Fig.
Philhygra laevicollis (Mäklin) A habitus B median lobe of aedeagus in lateral view C median lobe of aedeagus in ventral view D, E female pygidium F male tergite VIII G male sternite VIII H female tergite VIII I female sternite VIII. A, B, E–H after
Specimens have been collected from clear cut areas, transitional zone of a coniferous forest, seepages, and river and creek edges, from moss, leaf litter, gravel, dung, carrion and pitfall traps (
Philhygra laevicollis is a western Nearctic species that was previously considered to include eastern populations that we here treat as Philhygra pseudolaevicollis sp. nov. that differs in male genitalia but also by the divergent DNA barcode sequence.
Neither this species nor P. laevicollis are known from MB, this error was corrected by
(DNA-barcoded specimens). Canada: Ontario: Puslinch, Hanner property, 43.4464, -80.2512, Malaise trap in hardwood forest, 21.VIII.2008, T. Terzin (1,
Origin. Palaearctic (adventive in North America). Canada: MB, ON [new record]. USA: CT, MA, ME, NH, NY, PA, RI, SC, VT, WI.
Males of this species are easily recognized among other Canadian Philhygra by the simple, non-projecting median lobe in lateral view (Fig.
Most specimens of this species were collected by passive traps in a variety of habitats. In Sweden, P. palustris is considered a eurytopic species that occurs in various types of decaying plant matter, including compost, seaweed and hay piles, and along muddy shores of water bodies (
Philhygra palustris is a Palaearctic species that has become adventive and widespread in eastern North America. In the Palaearctic, it is very broadly distributed and reported from Europe, North Africa (Morocco), Russia (European and Siberia), Mongolia, North and South Korea (Lee and Ahn 2012), Japan, and northern China (
This species was reported from Canada (Manitoba) for the first time in the checklist by
Holotype (male) (
(DNA-barcoded specimens). Canada: New Brunswick: Restigouche Co., 9 km S of Saint Arthur, 47.818, -66.756, eastern white cedar swamp, in moss and litter near small ponds, 14.VI.2006, R.P. Webster (1, cRW).
Prefix -pseudo meaning false/not genuine, added to the sibling species name P. laevicollis (Mäklin).
This species is similar externally and genitally to P. laevicollis but may be distinguished from it by the following combination of characters: body on average narrower, antennomeres 6–7 more elongate (Fig.
Philhygra pseudolaevicollis Klimaszewski, Brunke & Pentinsaari, sp. nov. A habitus B median lobe of aedeagus in lateral view C female pygidium D male tergite VIII E male sternite VIII F female tergite VIII G female sternite VIII. Scale bars: 1 mm (A); 0.2 mm (B–G). Illustrations after
Body narrowly subparallel, moderately flattened, length 3.0–4.2 mm; colour dark brown, elytra dark brownish to brownish yellow, except for darker scutellar area and paler legs, basal antennomeres rust-brown (Fig.
Origin. Nearctic. Canada: NB, NS, ON, QC.
This species has been recorded from various wetland and riparian habitats in NB: in moss and leaf litter near brook and in litter, grasses, and moss on hummocks in old-growth eastern white cedar swamps and a wet alder swamp, in moist leaves along vernal pond margins in various mixed forests, and a red oak/red maple forest; also from pitfall traps in regenerating red spruce forests (NB) and from vernal pool litter in ON (summarized by
Although they were not re-examined here, the specimens reported by
Trichiusa robustula Casey, 1893
Trichiusa immigrata Lohse, 1984, syn. nov.
(DNA-barcoded specimens). Austria: Innervillgraten, Arntal, 46.8362, 12.3348, mountain forest and alpine pastures, car net, 25.VIII.2013, GBOL-Team
(non-barcoded). Numerous dissected specimens from Denmark were examined in the collection of NHMD.
Origin. Nearctic (adventive in Europe). Canada: ON, NB. United States: IA.
In its native range, this species has been collected in a variety of decaying plant matter, especially near water. This species was also common in compost in NB. In Europe, this species has been collected from similar microhabitats including grass clippings and compost (
Trichiusa robustula is a Nearctic species that is broadly distributed in eastern North America but not well collected. It was previously recognized under the synonym T. immigrata Lohse in the West Palaearctic (Europe, Canary Islands, Madeira;
When describing his new species,
We would like to thank the curators listed under Materials and methods for access to the material under their care. Thanks to J. Pedersen (NMHD) and V. Assing (Hanover, Germany) for their valuable input regarding the synonymy of Nearctic and Palaearctic species, and to V. Assing, A. Hansen (NHMD), and A. Bogri (NHMD) for generously sharing habitus and genitalia images for several species. We thank A. Newton (FMNH) for bringing the Neoisoglossa/Isoglossa issue to our attention. We also thank L. Hendrich, S. Schmidt (
Specimen voucher data and corresponding DNA barcode accession numbers
Data type: specimen data
Explanation note: All specimen data and sequence accession numbers associated with the DNA barcode dataset used in this study.