A review of Canadian and Alaskan species of the genus Liogluta Thomson, and descriptions of three new species (Coleoptera, Staphylinidae, Aleocharinae)

Abstract Fourteen species of Liogluta Thomson are reported from Canada and Alaska. Three of these are described as new to science: Liogluta castoris Klimaszewski & Webster, sp. n.; Liogluta microgranulosa Klimaszewski & Webster, sp. n.; and Liogluta pseudocastoris Klimaszewski & Webster, sp. n. The previously unknown male of Liogluta gigantea Klimaszewski & Langor, Liogluta quadricollis (Casey), Liogluta wickhami (Casey), and female of Liogluta granulosa Lohse are described, and illustrated. Liogluta aloconotoides Lohse is synonymized with Liogluta terminalis (Casey). New provincial and state records are provided for six Liogluta species. A key to species, revised distribution with new provincial records, and new natural history data are provided.


Terminalis species group
This group of species is characterized by: small eyes, with diameter of eye distinctly shorter than postocular area of head in dorsal view (Figs 1,8,16,23,30); integument of forebody moderately to highly glossy (Figs 1,8,16,23,30); elytra short, at suture about as long as pronotum (Figs 1,8,16,23,30); male tergite VIII truncate apically, with or without minute crenulations (Figs 3,11,25), except with obtuse triangular projection in L. quadricollis (Fig. 18); median lobe of aedeagus with tubus usually arched and moderately narrow apically in lateral view (Figs 2,9,10,17,24); capsule of spermatheca with apical part compressed, with apical invagination small and short (Figs 7,15,22,29,34). ( Diagnosis. This species may be distinguished by the following combination of characters: body subparallel, entirely reddish-brown or with head and posterior abdomen chestnut brown (Fig. 1); length 3.9-4.5 mm; integument of forebody with meshed microsculpture, moderately glossy; head about one-quarter narrower than maximum width of pronotum; pronotum transverse, narrower at base and widest in apical third; elytra at suture about as long as pronotum; basal three articles of metatarsus elongate, first longest, second about as long as third, fourth shorter than either of preceding articles; apical margin of male tergite VIII with broad, short, truncate projection with rounded lateral angles, apical margin smooth or slightly crenulate (Fig. 3); female tergite VIII with apical margin broadly, just visibly emarginate (Fig. 6); genital structures as illustrated (Figs 2, 7). Natural history. Klimaszewski et al. (2011) reported this species (as L. aloconotoides) from various forest types and on coastal limestone barrens in Newfoundland. Specimens from New Brunswick were collected from dung in a coastal red spruce forest, by treading sedges along a small lake margin, from a Lindgren funnel trap deployed in a rich Appalachian hardwood forest with some conifers, and from a pitfall trap (Webster et al. 2012). In Alberta, adults were reared from well-decayed white spruce logs (Klimaszewski et al. 2015). Elsewhere, adults were captured in various forest types including a recently burned forest. The type specimens of L. aloconotoides were captured in August (Lohse et al. 1990). Klimaszewski et al. (2015) reported adults from July to October.

Liogluta terminalis
Distribution. Recorded from LB, NB, NF, NS, ON, QC, AB, YT, and BC (Casey 1906, Lohse et al. 1990, Majka and Klimaszewski 2008, Webster et al. 2012, Bousquet et al. 2013, Klimaszewski et al. 2015, and newly in USA from MT and NH. Comments. We have examined the female holotype of L. terminalis (Casey) from Glenora, British Columbia, and compared it with the specimens of L. aloconotoides Lohse east of the Rocky Mountains. We found no external or genital differences between the holotype of L. terminalis and the other female specimens identified as L. aloconotoides and therefore we consider L. aloconotoides as a new synonym of L. terminalis. Lohse, 1990 Figs 8-15 Liogluta (Anepsiota) trapezicollis Lohse, in Lohse et al. 1990: 165 Diagnosis. This species may be distinguished by the following combination of characters: body subparallel, slender, dark brown to black with pronotum brown and paler than head, elytra yellowish or reddish-brown (Fig. 8); length 3.8-4.4 mm; integument of forebody with meshed microsculpture moderately pronounced, surface moderately glossy; head about one-quarter narrower than maximum width of pronotum; pronotum transverse, narrower at base and widest at middle (width of pronotum variable, some specimens have pronotum markedly narrower than base of elytra and some have pronotum nearly as wide as base of elytra); elytra at suture slightly shorter than pronotum; basal three articles of metatarsus elongate, subequal in length and each slightly longer than fourth article; male tergite VIII with apical margin truncate, bordered by two short lateral teeth, variably sculptured and ranging from smooth to crenulate, or denticulate along margin (Fig. 11); genital structures as illustrated (Figs 9-15).

Liogluta trapezicollis
Natural history. The holotype was collected in July (Lohse et al. 1990). The Alaskan specimens were collected in July only from alpine zones between 453 and 1071 m elevation, none were collected in lowland forests. Habitats include alpine flood meadows, under rocks, herbaceous heath with Luetkea, Cassiopes, and Lupinus, low rocky tundra with Dryas, meadow heath with Phyllodoce, Senecio, and Luetkea, shrubby krummholz with Elliottia and Tsuga, wet meadows with Carex, Petasites, Senecio and Ranunculus.
Comments. We were not able to compare types of L. trapezicollis Lohse with the specimens we examined, and our determinations are based on the published description by Lohse in Lohse et al. (1990). The types of L. trapezicollis housed in the Canadian National Collection of Insects were borrowed several years ago by V. Gusarov (Oslo, Norway) and our persistent efforts to have these specimens returned to Canada have failed.  body with weak meshed microsculpture, surface highly glossy; head slightly narrower than pronotum; pronotum subquadrate; antennae enlarged and black to brown; elytra about as wide as pronotum and at suture about as long as pronotum; basal two articles of metatarsus distinctly elongate, subequal in length, each longer than third article. New description of male. Apical margin of tergite VIII with broad, obtusely triangular projection in almost middle half with rounded lateral angles (Fig. 18); sternite VIII elongate, rounded apically, with antecostal suture arcuate, well separated from basal margin (Fig. 19); median lobe of aedeagus with tubus short and sinuate, its subapical part narrowly elongate in lateral view, internal sac structures distinct (Fig. 17). Fe male. Tergite VIII with apical margin obtusely angulate, broadly rounded at middle (Fig. 20); sternite VIII shallowly, broadly emarginated medially (Fig. 21); spermatheca with a short club-shaped capsule and short apical invagination, stem long and highly sinuate (Fig. 22).
Natural history. The holotype and the other BC specimen were collected in September, the Carrs collected a male under a rock in a poplar stand.
Distribution. Vancouver Island, British Columbia (Casey 1894 Diagnosis. This species may be distinguished by the following combination of characters: body broadly subparallel (Fig. 23); pronotum, elytra, legs and basal antennal article reddish-brown, head and abdomen chestnut brown (Fig. 23); length 4.0-4.2 mm; integument of forebody with meshed microsculpture, moderately glossy; head about one-third narrower than maximum width of pronotum; pronotum more or less evenly arcuate laterally (Fig. 23); elytra at suture about as long as pronotum; basal three articles of metatarsus missing in holotype (Fig. 23). New description of male. Apical margin of tergite VIII with very broad, short, subtruncate projection with rounded lateral angles, with apical margin faintly crenulate (Fig. 25); sternite VIII broadly rounded apically (Fig. 26); median lobe of aedeagus with tubus slightly arcuate ventrally, with apex narrow and rounded (Fig. 24). Female. Tergite VIII truncate apically (Fig. 27); sternite VIII very slightly broadly emarginate apically (Fig. 28); spermatheca with tubular capsule and deep apical invagination, stem thin, long, and highly sinuate (Fig. 29). This species is similar to L. terminalis but has dark brown antennae, head and pronotum (antennae, head, and pronotum are uniformly reddish-brown or only slightly darker than remaining parts of the body in L. terminalis). Spermatheca is differently shaped in each species; L. wickhami has smaller and differently shaped capsule with a deep apical invagination and has a shorter and differently looped posterior stem (Fig. 29). Liogluta wickhami is also very similar to L. vasta but can be distinguished by the shape of pronotum which has evenly arcuate sides and is broadest at middle (Fig. 23), while it is trapezoidal in shape and is broadest in apical third in the latter species (Fig. 30).
Natural history. Unknown. Distribution. The female holotype was captured in the Stickeen River Valley of British Columbia (Casey 1894), and one male was found in Three Guardsmen Pass, British Columbia.
The female lectotype is missing the spermatheca. The spermatheca of the Yukon specimen in CNC cited by Lohse and Smetana (1985), tentatively identified as belonging to this species, is illustrated in Fig. 34 (after Lohse and Smetana 1985). Males and more females from the type locality are needed to clearly define this species, which is here tentatively listed as a valid species. When more specimens of L. vasta become available for study and the morphological variation is known, we will be able to understand the relationship of this species to other nearctic Liogluta species. Liogluta vasta is also similar to some specimens (with broad pronotum) of L. trapezicollis. A DNA comparison between L. vasta and other Liogluta species is needed to clarify its identity and relationships.
Distribution. Canada: YT?. USA: AK. Discussion. The original type material of Homalota vasta Mӓklin, 1853 (ZMH) consists of two female specimens representing two different species in two genera, Atheta (as Boreophilia in Lohse and Smetana 1985) and Liogluta. Lohse and Smetana (1985) designated the female specimen belonging to Liogluta as the lectotype of Homalota vasta Mӓklin. However, the label data published by Lohse and Smetana (1985) for the Liogluta specimen corresponded to the Atheta specimen (see also discussion in Gusarov 2003b, who also mislabelled the specimens). We consider Lohse and Smetana's lectotype designation as valid regardless of the obvious mistake of publishing the wrong label data; therefore, the name vasta is affiliated with Liogluta. The female paralectotype belongs to Atheta keeni Casey, 1910. It is noteworthy that despite years of intensive collections made primarily between 2008-2013 in southeast Alaskan lowland forests and alpine zones, including in and around Sitka, which have resulted in 22,029 specimens of Staphylinidae (http://arctos.database.museum/saved/SE-AK-Staphylinidae), no specimens of Liogluta vasta were found.

Nigropolita species group
This group of species has a body shape non-typical for Liogluta and it resembles some of the Atheta (Dimetrota) species with elytra distinctly wider than head and pronotum (Figs 35,44). This group is characterized by large and bulging eyes, diameter of eye about as long as postocular area of head in dorsal view (Figs 35,44); integument of forebody highly glossy (Figs 35,44); elytra at suture at least as long as pronotum (Figs 35,44), and elytra about one-third broader than pronotum (Figs 35, 44); male tergite VIII truncate apically and apical margin entire or slightly produced with rounded lateral teeth (Figs 38,46); tubus of the median lobe of the aedeagus is arched, or almost straight and in lateral view moderately narrow apically (Figs 36, 45); spermatheca with spherical capsule with moderately long apical invagination and sinuate stem (Figs 42, 43). The spermatheca of L. nitens was not found and may be very small, not sclerotized, or absent.
Comments. This species is probably transcontinental in northern Canada.
Natural history. Adults were captured using pitfall traps in Carmanah Valley, Vancouver Island, from June to September, with the peak catch in June (Klimaszewski and Winchester 2002). They were found mainly in the interior and transition zones of a Sitka spruce forest (Klimaszewski and Winchester 2002). Several adults were collected from moss at the edge of an old road in the Queen Charlotte Islands, British Columbia. In Alberta, adults were collected in pitfall traps in various forest types in the Upper Cordilleran Ecoregion. Adults in Alaska were collected in a wide variety of habitats spanning lowland forests to alpine zones: alpine meadow litter, lowland forest clearcuts, floodplain meadows with Athyrium, Caltha, and Rubus, under rocks, in krummholz alpine habitats of Tsuga mertensiana, near bear dung in alpine habitats, old growth temperate rain coniferous forests, alpine heath with Empetrum, and Vaccinium, subalpine habitats with Salix, and Veratrum.

Comments.
There is considerable variation in length and width of elytra in specimens from Vancouver Island, Oregon (having broader and longer elytra), and those with narrow and shorter elytra from the Queen Charlotte Islands, Alberta, and Alaska. The genitalic features were the same in those of the typical form with the longer and broader elytra, and those with narrower and shorter elytra. Therefore, we consider this as intraspecific variation. Additional studies, including DNA comparison, are needed to reveal the relationship between these two morphotypes. Two UAM Alaskan specimens (UAM:Ento:152502, UAM:Ento:232546) were DNA barcoded (UA-MIC2665-15, UAMIC2701-15) and they cluster closely with two specimens of this species DNA barcoded from Alberta, Canada.

Granulosa species group
This group of species is characterized by: body medium-sized and subparallel (Fig. 50), eyes large and bulging, diameter of eye about as long as postocular area of head in dorsal view (Fig. 50); integument of forebody highly glossy (Fig. 50); elytra about one-fifth broader than pronotum, at suture about as long as pronotum (Fig. 50); elytra sparsely and strongly granulose (Fig. 51); apical margin of male tergite VIII with broad, short obtusely angular projection medially (Fig. 53); median lobe of aedeagus with tubus slightly arched ventrad, moderately narrow apically in lateral view (Fig. 52). Lohse, 1990 Figs 50-57 Liogluta (Liogluta) granulosa Lohse, in Lohse et al. 1990: 164 Diagnosis. This species may be distinguished by the following combination of characters: body broadly subparallel, dark brown, with elytra, tarsi and tibiae often reddish-brown (Fig. 50) (one specimen from northern Yukon was entirely black); length 2.8-3.3 mm; integument of forebody with moderately pronounced meshed microsculpture; head about one-eighth narrower than maximum width of pronotum (Fig. 50); pronotum transverse, about evenly wide in basal one-third of its length, then strongly broadest at apical one-third and gradually narrowed apically (Fig. 50); elytra at suture about as long as pronotum, its surface coarsely granulose (Fig. 50); basal two articles of metatarsus about the same length, each shorter than fifth article. Male. Apical margin of tergite VIII with short, very obtusely angular projection in medial twothirds with rounded lateral angles, margin of projection smooth or micro-denticulate (Fig. 53); apical margin of sternite VIII broadly parabolic (Fig. 54); median lobe of aedeagus with tubus broadly arched, bent ventrad, apex narrow and rounded (Fig. 52). Female. Apical margin of tergite VIII truncate in middle one-third (Fig. 55); apical margin of sternite VIII arcuate, antecostal suture distinctly sinuate (Fig. 56); spermatheca highly sinuate as illustrated (Fig. 57).

Liogluta granulosa
Natural history. Adults were captured in June, July, August, and October. One Alaskan specimen was captured in tundra between Rubus species and another at a creekside/ocean beach confluence, under boards and driftwood.
Distribution. Canada: YT. USA: AK (Lohse et al. 1990. Comments. Only a few specimens of this species are known. Its distribution is nordic and the habitat is unknown. One specimen (UAM:Ento:29798) in UAM was DNA barcoded (UAMIC2693-15), the first and only for this species so far.
Description. This species may be distinguished by the following combination of characters: body narrowly subparallel; head, apical articles of antennae, and posterior part of abdomen black, elytra brownish and mottled with black, remaining parts reddish-brown (Fig. 58); length 4.6-5.1 mm; integument of forebody with moderately pronounced meshed microsculpture, surface moderately glossy (Fig. 58); head about one-quarter narrower than maximum width of pronotum (Fig. 58); pronotum transverse, about evenly wide in basal half of its length, then strongly narrowed apically (Fig. 58); elytra at suture about as long as pronotum, surface finely and densely microgranulose; basal three articles of metatarsus about equally elongate, each longer than fourth article. Male. Apical margin of tergite VIII with very broad, very obtusely angular projection, with obtuse lateral angles and small tooth medially, margin often micro-crenulate (Fig. 60); sternite VIII rounded apically (Fig. 61); median lobe of aedeagus with tubus distinctly arched ventrad in apical half, apical part narrow (Fig.  59). Female. Tergite VIII with apical margin obtusely angulate (Fig. 62); sternite VIII with apical margin slightly emarginate medially (Fig. 63); spermatheca with stem long, sinuate, spiral posteriorly, capsule club-shaped with apical invagination deep and narrow (Fig. 64).
Distribution. Canada: Known only from New Brunswick, Canada. Natural history. Nearly all adults from New Brunswick were collected from American beaver (Castor canadensis Kuhl) dams. Most were collected from among sticks and debris near an overflow area of the dam, another from under overhanging sticks on the outer margin of the dam. One individual was collected from among leaves and sedges near a pond margin. Specimens were collected in July and August. Etymology. Castoris is a Latin adjective derived from the name of the American beaver (Castor canadensis Kuhl), in reference to beaver dams where some of the type specimens were captured.
Natural history. In New Brunswick, adults were collected using an aerial (butterfly) net in a mixed forest opening during evening flights (between 15:00 and 19:00 h) during April and May. A number of individuals were collected from among sticks and debris near the overflow area of a beaver dam during May. One individual was sifted from sphagnum and litter near a brook in an eastern white cedar swamp in May. In Nova Scotia, specimens were captured in flight interception, pan, and Malaise traps during the months of June and August. The single specimen from Ontario was captured in October. Comments. This species is similar to L. microgranulosa but in L. castoris the pronotum and elytra are more elongate and more reddish-brown (Fig. 65); the median lobe of the aedeagus has the apical part of the tubus broader and shorter in lateral view (Fig. 66); male tergite VIII is truncate and not at all angulate medially (Fig. 67); the sperma theca has a longer stem (Fig. 71); and female sternite VIII has an apical emargination which is much less noticeable and the antecostal suture is more distinctly sinuate (Fig. 70). Etymology. Pseudocastoris is the Latin prefix pseudo-, false, added to the species name castoris, reflecting the close similarity of the two species.

Liogluta pseudocastoris
Description. Body length 3.9-4.4 mm, subparallel; dark brown with irregularly shaped lighter areas on pronotum in some individuals, head and abdomen dark brown, antennae dark, and legs yellowish; integument moderately glossy, more so on posterior portion of abdomen; forebody with minute and sparse punctation and sparse pubescence (Fig. 72); elytra with micro-granulation (Fig. 72); head rounded and narrowed posteriorly, eyes large, each about as long as postocular area in dorsal view (Fig. 72); antennae with articles V-X subquadrate to slightly transverse (Fig. 72); pronotum transverse, broadly rounded laterally, slightly wider than head and narrower than elytra, pubescence directed latero-posteriad from midline of disc (Fig. 72); elytra transverse, at suture about as long as pronotum, slightly longer laterally, with pubescence directed posteriad; abdomen subparallel for most of its length, about as wide as elytra (Fig. 72). Male. Tergite VIII broadly rounded apically, margin smooth (Fig. 74); apical margin of sternite VIII broadly parabolic (Fig. 75); median lobe of aedeagus with bulbus narrowly oval, tubus almost straight with apical part narrowly rounded in lateral view (Fig. 73); internal sac without distinct sclerites but with some vaguelyshaped structures (Fig. 73). Female. Tergite VIII broadly rounded apically (Fig. 76); sternite VIII a little less broadly rounded apically, antecostal suture slightly sinuate, moderately separated from basal margin (Fig. 77); spermatheca with capsule clubshaped, [invagination not perceptible], stem sinuate, about equally narrow throughout with only posterior part enlarged but not twisted (Fig. 78). Natural history. Most individuals were collected from among sticks and debris near an overflow area of a beaver dam during May and June. Others were collected using an aerial (butterfly) net in a mixed forest opening during an evening flight (between 16:30 and 19:00 h) during May.
Distribution. Known only from New Brunswick, Canada.
Comments. This species is closely related to L. castoris and L. microgranulosa but in L. pseudocastoris the body is darker, particularly the pronotum, the pronotum is strongly narrowed basally with more angular posterior angles (Fig. 72); the shape of the median lobe of the aedeagus is different in lateral view, with the apical part narrower and very slightly arched ventrad (Fig. 73); the apical margin of male tergite VIII is evenly rounded (Fig. 74); the apical margin of female sternite VIII is not emarginate, with the antecostal suture only slightly sinuate (Fig. 77), and the shape of the spermatheca is different, with the posterior part of the stem enlarged but not twisted (Fig. 78). Diagnosis. This species may be distinguished by the following combination of characters: length 4.2-4.5 mm; body dark reddish-brown, with head dark brown, and legs and at least basal three antennal articles reddish-yellow; integument glossy; pronotum with dense punctation and pubescence; elytra with dense punctation and pubescence with very fine micro-granulation (Fig. 79); head subquadrate, slightly narrower than pronotum, large eyes, each about as long as postocular region in dorsal view (Fig. 79); pronotum subquadrate, widest at apical third (Fig. 79); elytra subparallel, as wide as pronotum and at suture about as long as pronotum (Fig. 79); abdomen subparallel, about as wide as elytra (Fig. 79); Male. Apical margin of tergite VIII with broad, moderate projection in middle three-fifth, with apical margin crenulate (Fig.  81); apical margin of sternite VIII broadly parabolic (Fig. 82); median lobe of aedeagus with tubus short and straight, apical part narrowly rounded in lateral view (Fig. 80). Female. Tergite VIII rounded apically (Fig. 83); apical margin of sternite VIII with broad, shallow median emargination (Fig. 84); spermatheca short, S-shaped, capsule short, club-shaped, stem broad, sinuate, slightly twisted posteriorly (Fig. 85).
Natural history. Adults were collected in a conifer forest using pitfall traps, in a spruce-moss forest using carrion-baited traps, and in a highbush blueberry field. Others were collected by sifting litter and moss, sifting Alnus litter and Sphagnum moss near a pond, and treading flooded Carex and grasses. The flight period is from May to October. Diagnosis (based on male lectotype). This species may be distinguished by the following combination of characters: small body size, length 2.8 mm; head and abdomen dark brown, pronotum, elytra and legs reddish-yellow (Fig. 86); integument glossy with weak meshy microsculpture; pronotum and elytra with moderately dense punctation and pubescence, elytra with very fine micro-granulation (Fig. 86); head subquadrate, slightly narrower than pronotum; large eyes, each about as long as postocular region in dorsal view (Fig. 86); antennae of the holotype are partially damaged and cannot be completely described, but fifth and sixth articles suggest that missing funicle articles are subquadrate; pronotum slightly transverse, widest near the middle (Fig. 86); elytra wider and slightly longer than pronotum (Fig. 86); abdomen subparallel, about as wide as elytra (Fig. 86). Male. Apical margin of tergite VIII with broad, truncate projection in middle two-thirds bounded laterally by small tooth-like processes, apical margin crenulate (Fig. 88); apical margin of sternite VIII evenly broadly parabolic from base (Fig. 89); median lobe of aedeagus with tubus bent slightly ventrad at middle, apical part relatively broadly rounded in lateral view (Fig. 87). Female. Unknown. Liogluta atriventris may be distinguished from the other species of the granulosa group by the following combination of characters: body size small, length 2.8 mm; elytra slightly longer than pronotum (Fig. 86); pronotum glossy with weak microsculpture (Fig. 86); shape of median lobe of aedeagus different in lateral view (Fig.  87), and projection on apical margin of male tergite VIII crenulate, with tooth-like processes laterally (Fig. 88).
Natural history. Unknown. Distribution. Known only from Vancouver Island, British Columbia. Comments. This species is known only from one damaged male specimen. More specimens, including females, are needed for study to confirm the status of this species.

Gigantea species group
This group is characterized by: body broad, eyes large and bulging, diameter of eye about as long as postocular area of head in dorsal view (Fig. 90); integument of forebody glossy (Fig. 90); elytra not granulose, about one-fifth broader than pronotum, at suture about as long as pronotum (Fig. 90); apical margin of male tergite VIII rounded with broad crenulations (Fig. 92); median lobe of aedeagus with tubus arched slightly ventrad near apex, apical part narrow in lateral view (Fig. 91); spermatheca vaguely S-shaped (Fig. 96); female with apical margin of sternite VIII broadly truncate, with a row of microsetae (Fig. 95).