Body moderately broad, forebody slightly and abdomen strongly glossy (Fig. 6); length 3.0–3.6 mm; dark brown to almost black, appendages light brown or reddish brown; antennomeres VIII-X subquadrate to slightly elongate; pronotum as long as elytra at suture, maximum width of pronotum slightly less than maximum width of elytra, pronotum in males is longer and less transverse that in females. Male. Tubus of median lobe of aedeagus with two, sharp basolateral projections on both sides in dorsal view (Figs 9, 10), and in lateral view ventral part of tubus produced angularly at base (Figs 7, 8), bulbus broadly oval in dorsal view, with two elongate narrow sclerites of internal sac (Fig. 9); tergite VIII arcuate apically (Fig. 11); sternite VIII elongate, parabolic apically (Fig. 12). Female. Spermatheca: capsule pitcher-shaped basally with elongate narrowly conical apical part without apical invagination, stem sinuate, narrowly looped posteriorly (Fig. 15); tergite VIII arcuate apically (Fig. 13); sternite VIII rounded apically, antecostal suture moderately sinuate (Fig. 14).

The median lobe of aedeagus of B. eremita is similar to that of B. islandica, but tubus is more elongate, narrower, less arcuate, and with basal projection angular in lateral view (Figs 7, 8), in dorsal view tubus with two pointed latero-basal projections (Figs 9, 10). In B. islandica, the tubus is broader, shorter, more arcuate ventrally, and with basal arcuate projection rounded in lateral view (Fig. 17), in dorsal view tubus with two smaller and less angular basal projections (Fig. 18). Spermathecae of the two species is variable in shape and very similar, female tergite and sternite VIII are similar in shape but sternite VIII in B. eremita has feebly arcuate medial part of antecostal suture (Fig. 14), which is strongly sinuate in B. islandica in the majority of examined specimens (Fig. 22). On average, the body of B. eremita is narrower and elytra shorter than that of B. islandica.

Figures 6–15. 

Boreophilia eremita (Rye): 6 habitus 7, 8 median lobe of aedeagus in lateral view 9, 10 median lobe of aedeagus in dorsal view 11 male tergite VIII 12 male sternite VIII 13 female tergite VIII 14 female sternite VIII 15 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Holarctic species; recorded from north and central Europe, Ireland, Ukraine, Russia (west and east Siberia) and the Russian Far East; Canada: LB, NB, MB; USA: AK.

Habitat: in NB – old silver maple forest with green ash and seasonally flooded marsh; silver maple swamp, margin of vernal pond, found in moist leaves. In AK - creekside/ocean beach confluence, under boards and drift wood; black and white spruce, willow; subalpine habitat with Veratrum, and Calamagrositis. Collecting methods: sifting litter, Lindgren funnel trap, Malaise trap, hand collecting. Collecting period: May to August.

NEARCTIC: Canada, New Brunswick, Queens Co., Grand Lake Meadows P.N.A., 45.8227N, 66.1209W, 19–31.V.2010, old Silver Maple forest with Green Ash and seasonally flooded marsh, Lindgren funnel trap (LFC) 1 male; New Brunswick, Queens Co., Canning Grand Lake near Scotchtown, 45.8762N, 66.1817W, 25.V.2006, Silver maple swamp, near lake margin, margin of vernal pond in moist leaves, RP Webster coll. (LFC) 1 female.

USA, Alaska, mi 110 Denali Hwy., Seattle Creek, 15.VII.1978, JM Campbell and S Smetana, coll. GA Lohse MHNG-1994 (MHNG) 1 male.

Females without male association, tentatively included in B. eremita [they may represent B. eremita or extreme narrow forms of B. islandica]: USA, Alaska, Naknek, 58.73973N, 157.0636W, creekside/ocean beach confluence, under boards and drift wood, hand collected, 10.VI.2007, D. Sikes, UAM100012293, UAM100012313, UAM100012315 (UAM) 3 females; Kenai Pen., Kalifonsky Bch. Near Kenai, 8.VI.78, A Smetana and E Becker (MHNG) 1 female.

PALEARCTIC: Finland: Muonio, Renkonen (MHNG) 6 females. Germany: b. Grönau, Lübeck, 10.V.34, G Benick (MHNG) 1 female [ident. uncertain]. Russia: Siberia, Tschimilcan, FI. Lenam. [Lena River], B. Poppius (MHNG) 1 male.

We have examined several European specimens identified as B. hercynica, which have the shape of median lobe of aedeagus and spermatheca similar to B. eremita, but the body color and the shape and proportions of forebody were different: body brown with dark brown head and pronotum and particularly elytra paler, and pronotum strongly transverse with sides broadly and evenly arcuate, elytra at suture slightly shorter than pronotum along midline. These specimens may represent extreme variation of B. eremita or a different and distinct species. Additional material is needed, and possibly DNA studies, to establish clear status of these specimens.

These specimens were labelled as follows: Belgium, Elsenborn, VIII.1931, Hohes Venn, coll. Benick (MHNG) 1 male; Germany, Brocken, Heinemann, hercynica s. mihi, det L Brundin (MHNG) 1 male; same label as before except: islandica Kr. (MHNG) 1 male; Brocken, Harz, 10.VI.1912, islandica, col. G Benick (MHNG) 1 male, Brocken, Heinemann (MHNG) 1 female, Brocken, 10–12.V.1913, Linke leg., A. islandica v. hercynica Renk., (MHNG) 1 female, Brockenkuppe, 26.VII.1916, coll. G Bennick (MHNG) 1 male; Norway, Kongberg [= Konsberg?], Münster, islandica Kr., coll. G Benick (MHNG) 1 female, Hammerfest, A Strand, islandica s. mihi, det. L Brundin (MHNG) 1 male, Norge, Skibotu, 22.VIII.1960, leg. Puthz, coll. Benick (MHNG) 1 male; Scotland, D Sharp, hercynica s. mihi det. L Brundin, Metaxya eremita Rye (MHNG) 2 males; Shetland Is., Mainland, Stany Fields nr. Wal Is., 13.VII.1963, ME Bacchus BM 1963-471, coll. G Benick (MHNG) 1 female; Sweden, Dorotea, Sa. Lappland, 30–31.V.1935, Bruce, A. islandica, coll. Benick (MHNG) 1 female.

Our data included six sequences of B. eremita from Finland (four from Lapland and two from Northern Ostrobothnia), which grouped into BIN BOLD:ABW4331. BOLD reports these sequences have an average distance of 0.06%, a maximum distance of 0.18% and are 4.1% distant from their nearest neighbor.

Body broad, forebody moderately and abdomen strongly glossy (Fig. 16); length 2.8–3.5 mm; uniformly black with paler, reddish brown appendages, or head, pronotum and VI-VII basal segments of abdomen dark brown, remainder of the body reddish brown, appendages light brown, sometimes elytra with some reddish tinge; antennomeres VIII-X subquadrate; pronotum as long as elytra at suture, maximum width of pronotum distinctly less than maximum width of elytra; elytral length variable, as long as pronotum or slightly longer. Male. Tubus of median lobe of aedeagus with two basolateral and slightly angular apically projections in dorsal view (Fig. 18), and one rounded projection in lateral view (Fig. 17); bulbus moderately broadly oval with two elongate sclerites of internal sac in dorsal view (Fig. 18); tergite VIII arcuate apically (Fig. 19); sternite VIII elongate, parabolic apically (Fig. 20). Female. Spermatheca: capsule pitcher-shaped basally with tubular apical projection moderately long and narrowed apically, stem coiled posteriorly, there is great variability in the shape of capsule (Fig. 23); tergite VIII arcuate apically (Fig. 21); sternite VIII rounded apically, antecostal suture strongly sinuate in most examined specimens (Fig. 22). For the differences between this and previous species see diagnosis of B. eremita.

Figures 16–23. 

Boreophilia islandica (Kraatz): 16 habitus 17 median lobe of aedeagus in lateral view 18 median lobe of aedeagus in dorsal view 19 male tergite VIII 20 male sternite VIII 21 female tergite VIII 22 female sternite VIII 23 spermatheca. Scale bars: 1 mm (habitus); 0.2 mm (remaining).

Holarctic species; recorded from Fennoscandia, Estonia, Faeroe Islands, Great Britain, Iceland, Russia (North European Territory); Canada: LB, NF, NT, NU, YT; USA: AK.

Habitat [new data]: Betula, Salix litter; Salix tundra hillside; Salix/Betula/Alnus/grasses; black/white spruce, willow; vegetation at lakeshore pond; subalpine habitat with Veratrum, Calamagrostis, and Leymus, Heracleum, Angelica. Collecting methods: hanging Malaise trap, pitfall traps, sweeping with net. Collecting period: May to August.

Females of this species may be confused with other species of Boreophilia and particularly those of closely related B. eremita. Associating females with males is considered here to be the most reliable way of identifying females of this and the previous species. At present, B. islandica is considered a somewhat variable species. Specimens vary from moderately robust and narrower to more robust and broader, with elytra as long as pronotum or slightly longer, all with the same morphology of genitalia. The BIN BOLD:ABX3767 formed a sister group to B. islandica in our analysis and was represented by a single female from Finland. The capsule of its spermatheca is curved at an angle of nearly 90 degrees and was among those shapes included in the illustrations of B. islandica by Palm (1970). BIN BOLD:ABX3767 may represent an undescribed species, or one of the described species not included in our DNA dataset, but corresponding males should be examined. A separate DNA based study is recommended to examine specimens of B. islandica sensu lato from a broad Holarctic distribution, including males and females.

NEARCTIC: Canada, Newfoundland, Long Range Mts., Portland Cr. Hill, 12–13.VIII.1982, Belland, Larson, McDonald (LFC) 1 female; Northwest Territories, 10.VI.1956, R.E. Leech (LFC) 1 male; Northwest Territories, Aklavik, 14.VI.1956, RE Leech (MHNG) 1 male, 1 female.

USA, Alaska: Anaktuvuk Pass, 647 m el., 68.14049N, 151.74091W, +/- 250 m, Salix, sweeping, 19.V.2016, D Sikes, K Daly, UAM100427773 (UAM) 1 male; Alaska, Aguttu Is., 15 m el., 52.473083N, 173.59065E, +/- 30 m, beach, Heracl. lanatum, 5 pitfalls, 5–19.VI.2008, L Kenney, R Kaler, UAM100029353 (UAM) 1 male; Aiktak Is., 10–20 m el., 54.18549N, 164.8432W, +/- 50 m, Leymus, Heracleum, maritime tundra, 5 pitfalls, 17–31.VIII.2009, A.L. Larned, UAM100321897, UAM100321780 (UAM) 2 males; same data except: Leymus, Heracleum, Angelica, 11–29.VII.2009, UAM100321769 (UAM) 1 female; UAM100321861 (UAM) 1 female; 54.18356N, 164.82793W, +/- 1 km, maritime tundra, Leymus, Heracleum, 5 pitfalls, 9–24.VI.2009, AL Larned, UAM100322546 (UAM) 1 female; 24.VI-11.VII.2009, S. Sapora, UAM100323153, UAM100323220 (UAM) 2 females; 54.11036N, 164.50500W, +/- 5 m, streamside, Leymus, 2 pitfalls, 26.VI-14.VII.2008, BA Drummond, UAM100030104, UAM100030117 (UAM) 2 females.

Females without male association, tentatively included as B. islandica:

USA, Alaska: White Mtns. Rec. Area, 180 m el., 65.33469N, 146.83969W, +/- 10 m, b. & w. spruce, willow, hanging Malaise, 10–17.VI.2016, J Hagelin, UAM100407456 (UAM) 2 females; S. Chilkat Pen., pl. 19, 652 m el., 58.42509N, 135.14229W, +/- 30 m, subalpine, Veratrum, Calamagrostis, hand coll., 9.VII.2010, DS Sikes, UAM100277138 (UAM) 2 females; Kasatochi, 42 m el., 52.16869N, 175.52475W, +/- 34 m, under rocks, Talus, cliff base, 11.VI.2008, DS Sikes, UAM100031453, UAM100031445 (UAM) 2 females.

PALEARCTIC: Czech Republic: Bohemia occ., Frant. Láznê-Soos, 1961, Smetana (MHNG) 2 females. Denmark: Helsinge, 6.V.1915, Häkan Lindberg (MHNG) 1 female. Finland: Kittilä, Renkonen (MHNG) 1 female; Lapp., Petsamo, 3.VII.1929, Häkan Lindberg (MHNG) 1 female; Oa Lappfiärd, 3.V.1944, Harald Lindberg (MHNG) 1 female. Germany: I. b. Grönau, Lübeck, 27.III.12, G. Benick (MHNG) 1 female; Brocken, 15.V.32, Foln.…[not clear] (MHNG) 1 female; Brocken, Heineman (MHNG) 1 female; Brocken Harz, 15.VII.1935, Fehse, Thale, G Benick (MHNG) 1 female. Island: Island, VII.1969, G Benick (MHNG) 1 male. Norvay: Lakselv Po, A Strand (MHNG) 3 females. Sweden: Lule Lappmark, Vietas 20, 27.VI.1967, TE Leiler (MHNG) 1 female; Moen M. elev., A Strand (MHNG) 1 female; Ő Torne, T Palm (MHNG) 1 female; Imtl. Frost-viken, Palm (MHNG) 1 female.

Our data included one sequence identified as B. islandica from Newfoundland and Labrador, Canada, and one sequence identified as Boreophilia sp. collected from Churchill, Manitoba, which are the only members of BIN BOLD:AAH0226. BOLD reports these sequences are 2.79% distant from their nearest neighbor.

Body broad, forebody moderately and abdomen slightly more glossy (Fig. 24); length 3.4–3.8 mm; head, pronotum and abdomen except for its apex dark brown, elytra dark-reddish brown, appendages light brown, or body entirely dark brown to almost black with tarsi reddish brown; antennomeres VIII-X subquadrate; pronotum shorter than elytra at suture, maximum width of pronotum distinctly less than maximum width of elytra. Male. Tubus of median lobe of aedeagus evenly arcuate laterally, apex narrow and slightly pointed in lateral view (Fig. 25), bulbus oval, broad basally and narrowed apically in dorsal view, and with two elongate sclerites of internal sac (Fig. 26); tergite VIII arcuate apically (Fig. 27); sternite VIII elongate, parabolic apically (Fig. 28). Female. Spermatheca: capsule pitcher-shaped basally with broadly tubular apical projection, moderately long and pointed apico-laterally, stem coiled posteriorly (Fig. 31); tergite VIII arcuate apically (Fig. 29); sternite VIII rounded apically, antecostal suture slightly sinuate (Fig. 30).

Figures 24–31. 

Boreophilia fusca (C.R. Sahlberg): 24 habitus 25 median lobe of aedeagus in lateral view 26 median lobe of aedeagus in dorsal view 27 male tergite VIII 28 male sternite VIII 29 female tergite VIII 30 female sternite VIII 31 spermatheca. Scale bars: 1 mm (for habitus); 2 mm (remaining).

Holarctic species; recorded from Fennoscandia, Russia (west and east Siberia) and the Far East; Canada: NT; USA: AK.

Habitat: tundra. Collecting methods: not recorded in Nearctic region. Collecting period: June and July.

NEARCTIC: Canada, NT, Aklavik, 16.VI.1956, EF Cashmann, fusca Sahlb. Det. Lohse (MHNG) 1 male; NT, Aklavik, 29.VI.1956, EF Cashmann, fusca Sahlb. Det. Lohse (MHNG) 1 female.

PALEARCTIC: Finland: Muonio, Renkonen, 2531, A. fusca Sahlb., Renkonen det., coll. G Benick (MHNG) 2 females. Country unspecified: Bjerkeng Schn. 21.VI.1912, coll. G Benick (MHNG) 1 male.

Our data included two sequences of B. fusca, one from Finland and one from Manitoba, Canada, which grouped into BIN BOLD: AAG4311. BOLD reports these sequences have an average and maximum distance of 0.54% and are 9.68% distant from their nearest neighbor.

Body broad, forebody glossy; length 2.8–3.5 mm; black with tarsi reddish brown (Fig. 32); antennomeres VIII-X subquadrate; pronotum as long as or slightly shorter than elytra at suture, maximum width of pronotum distinctly less than maximum width of elytra. Male. Tubus of median lobe of aedeagus narrow, broadly arcuate in lateral view, apex narrow and rounded (Figs 33, 34), bulbus large, oval, broad basally and narrowed apically in dorsal view, and with two elongate sclerites (Figs 35, 36); tergite VIII arcuate apically (Fig. 37); sternite VIII elongate, parabolic apically (Fig. 38). Female. Spermatheca: capsule pitcher-shaped basally with evenly, broadly tubular apical projection, moderately long and rounded apically, stem sinuate, half-looped posteriorly (Figs 41, 42); tergite VIII broadly rounded apically (Fig. 39); sternite VIII rounded apically and truncate medially, antecostal suture straight medially and slightly sinuate laterally (Fig. 40).

Figures 32–42. 

Boreophilia hyperborea (Brundin): 32 habitus 33, 34 median lobe of aedeagus in lateral view 35, 36 median lobe of aedeagus in dorsal view 37 male tergite VIII 38 male sternite VIII 39 female tergite VIII 40 female sternite VIII 41, 42 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Holarctic species; recorded from Fennoscandia, Greenland, Russia (North European Territory); Canada: NT, NU; USA: AK.

Habitat: tundra, under rocks. Collecting methods: hand collected from under rocks. Collecting period: June and July.

NEARCTIC: Canada, NT, Barthurst Inl., Hiukitak River, 3.VIII.1966, GE Shewell, B. hyperborea Brn., det. GS Lohse (NHNG) 1 male.

USA, Alaska, Toolik Field Station, 724 m el., 68.6286N, 149.59772W, +/- 36 m, under rocks, 1- 3.VI.2008, D.S. Sikes, UAM100031281 (UAM) 1 male; Anaktuvuk Pass, 665 m el., 68.14059N, 151.74266W, +/- 200 m, Salix, tundra hillside, cobble, pollinator pitfall, 20.V.2016, D. Sikes, K. Daly, UAM100427687 (UAM) 1 female [tentative association].

PALEARCTIC: Norway, Vaalaasjö Andr. Strand, coll. G. Benick (NHNG) 1 female; Barviksmyren, W of Smelror, Varangerhalvøya, 22.VII.1998, V. Mahler (UCC) 1 female.

Greenland. Sdr. Strømfjord, 1.VII.1979, Brundin det. 1940 (NHMD) 1 female.

Our data included two sequences of B. hyperborea, one from Russia and one from Manitoba, Canada, which grouped into BIN BOLD:AAG4302. BOLD reports these sequences have an average and maximum distance of 0.16% and are 6.82% distant from their nearest neighbor.

Body moderately broad, forebody moderately glossy, abdomen slightly more so (Fig. 43); length 3.0–3.5 mm; head, pronotum and abdomen except for its apex dark brown, elytra reddish brown, appendages light brown, or body dark brown to almost black with elytra partially dark reddish brown and tarsi reddish brown; antennomeres VIII-X subquadrate to slightly elongate; pronotum ca. as long as elytra at suture, maximum width of pronotum distinctly less than maximum width of elytra. Male. Tubus of median lobe of aedeagus almost straight ventrally, apex narrow and pointed, slightly produced ventrally in lateral view (Figs 44, 45), bulbus broadly oval in dorsal view and with two elongate sclerites of internal sac (Fig. 46); tergite VIII arcuate apically, slightly pointed medially (Fig. 47); sternite VIII elongate, parabolic apically (Fig. 48). Female. Spermatheca: capsule pitcher-shaped basally with broadly tubular apical projection, moderately long and rounded apically, stem sinuate and without posterior loop (Figs 51, 52); tergite VIII arcuate apically (Fig. 49); sternite VIII truncate apically and slightly emarginate medially, antecostal nearly straight medially and slightly sinuate laterally (Fig. 50).

Figures 43–52. 

Boreophilia nearctica Lohse: 43 habitus 44, 45 median lobe of aedeagus in lateral view 46 median lobe of aedeagus in dorsal view 47 male tergite VIII 48 male sternite VIII 49 female tergite VIII 50 female sternite VIII 51, 52 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Nearctic species; recorded from Canada: AB [new record], LB, MB, QC, YT, NF; USA: AK.

Habitat [new data]: black spruce forest; alpine meadow. Collecting methods: hanging Malaise trap, pitfall traps, hand collecting under rocks and litter. Collecting period: July to September.

Canada, Alberta, Plateau Mountain, 50.226 -114.555, alpine meadow, under rocks and litter, 5.VII.2002, G Pohl and D Langor, 2 males, 1 female (CCDB-28541-D04, CCDB-28541-D05, CCDB-28541-D06) (NFRC); Newfoundland, Burnt Cape, site 10, 21U 0586332E, 5711616N, 51.54902N, 5.75489W, 24.VII.2003–7.VIII.2003 (LFC) 1 male; Burnt Cape, 55.755W, 51.549W, Coastal meadow, pitfall trap, 10–24.VII.2003, AM Hynes, Site 1–10 (LFC) 1 male; Yukon Territory, North Fork Pass, Ogilvie Mts.,3500’, 17.VI.1962, RE Leech, paratype CNC No. 20308 (CNC) 1 male; North Fork Pass, Ogilvie Mts., 4100’, 20.VI.1962, PJ Skitsko (CNC) 1 female; Quebec, Bonne Esperance, 14.VII.1929, WJ Brown, nearctica nov. sp. Lohse, Paratype (NHNG) 1 female.

USA, Alaska: Naknek, 58.73973N, 157.0636W, creek side/ocean beach confluence, under boards and drift wood, hand collected, 10.VI.2007, DS Sikes, UAM100012316 (UAM) 1 male; AK, Selawik NWR, 8 m el., 66.56137N, 158.99834W, +/- 304 m, Spirea, dwarf birch, 23–24.VI.2010, DS Sikes, UAM100283949 (UAM) 1 male; AK, Fairbanks, Bonanza Crk., 237 m el., 64.71922N, 148.38416W, +/- 10 m, birch, spruce, hanging Malaise trap, 6–13.V.2016, J Hagelin, UAM100407183 (UAM) 1 male; AK, Kenai Mts., Tern Lake Campground, 700’, 18.Vi.1978, Smetana and Becker (NHNG) 1 male.

The southernmost record of this species in the Rockies of southern Alberta suggests that B. nearctica probably occurs continuously along the western cordilleras, at successively higher elevation sites with decreasing latitude.

Our data included five sequences of specimens identified as B. nearctica, two from Alaska and three from Alberta, Canada, which grouped with two sequences identified as Boreophilia sp. into BIN BOLD:ACU9385. Our calculations indicate that the five sequences identified to species have an average distance of 0.14%, a maximum distance of 0.33% and are 6.37% distant from their nearest neighbor.

Body very broad, forebody moderately and abdomen strongly glossy (Fig. 53); length 3.0–3.5 mm; head, pronotum and abdomen except for its apex dark brown, elytra dark-reddish brown medially, appendages brown, or entire body dark brown to almost black and tarsi reddish brown; antennomeres VIII-X elongate; pronotum ca. as long as elytra at suture, maximum width of pronotum slightly less that the maximum width of elytra. Male. Unknown. Female. Spermatheca: capsule pitcher-shaped basally with broadly tubular and slightly pointed apical part, stem short, strongly sinuate and looped posteriorly (Fig. 56); tergite VIII arcuate apically (Fig. 54); sternite VIII rounded apically, antecostal suture strongly sinuate medially (Fig. 55). Females of this species may be confused with those of B. fusca, from which they differ by distinctly elongate antennomeres VIII-X, more deeply medially sinuate antecostal suture of sternite VIII, and spermathecal capsule more evenly elongate and apex less pointed laterad.

Figures 53–56. 

Boreophilia ovalis Klimaszewski and Langor: 53 habitus 54 female tergite VIII 55 female sternite VIII 56 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Nearctic species, recorded only from Canada: NF.

Habitat: unspecified forest. Collecting methods: one female was captured in Malaise trap. Collecting period: June to September.

. Two specimens of this species, one being a paratype, were submitted for DNA barcoding but failed to generate DNA sequences (process IDs on BOLD: LFCAB222-15, NGSFT931-15).

Body narrow, subparallel, moderately glossy, abdomen slightly more so (Fig. 57); length 3.0–3.8 mm; head, pronotum and abdomen dark brown, elytra reddish brown, legs paler, yellowish brown; antennomeres VIII–X subquadrate; pronotum longer than elytra at suture, maximum width of pronotum ca. the same as maximum width of elytra. Male. Tubus of median lobe of aedeagus almost straight (slightly arcuate) in lateral view, apex triangular in shape, slightly pointed (Fig. 58), bulbus broad in dorsal view and with two elongate narrow sclerites as illustrated (Figs 59, 60); tergite VIII arcuate apically (Fig. 61); sternite VIII elongate, parabolic apically (Fig. 62). Female. Undescribed.

Figures 57–62. 

Boreophilia nomensis (Casey): 57 habitus 58 median lobe of aedeagus in lateral view 59, 60 median lobe of edeagus in dorsal view 61 male tergite VIII 62 male sternite VIII. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Nearctic species, recorded from Canada: YT, BC [new record]; USA: AK.

Habitat: spruce and aspen forest with horsetail/shrub/grass undergrowth; edge of snowfield. Collecting methods: pitfall trap, hand collecting under rocks. Collecting period: June, July and August.

Canada, British Columbia, Kinaskan Lake Provincial Park, Kinaskan Lake Trail, 57.532–130.202, 833 m, spruce and aspen forest, pitfall trap, 1.VIII.2014, BIObus 2014, (BIOUG24477-H04) (CBG). USA, Alaska, Thompson Pass, 61.137 -145.745, under rocks nr. snowfield, 28.VII.2010, DS Sikes and AB Sikes (UAM100288002) (UAM).

Our data included two sequences of specimens identified as B. nomensis, one from Alaska and one from British Columbia, Canada which grouped into BIN BOLD:ACU9384. BOLD reports these sequences have an average and maximum distance of 1.06% and are 4.64% distant from their nearest neighbor.

Canada, Yukon Territory, British Mts., Windy Ridge, 450 m, 69.27N, 140.26W, 2.VII.1984, 84–47, sifting Salix litter, JM Campbell (CNC).

labeled as the holotype (CNC) 3 sex undetermined.

Body narrowly subparallel, forebody moderately glossy, abdomen slightly more so (Fig. 63); length 3.0–3.4 mm; uniformly dark brown to black, appendages yellowish brown or reddish brown; antennomeres VIII-X subquadrate; pronotum ca. as long as elytra at suture, maximum width of pronotum slightly less than maximum width of elytra. Male. Tubus of median lobe of aedeagus arcuate basally, straight apically and produced ventrad in lateral view, apex broad and rounded in lateral view (Fig. 64), bulbus broad and with two large sclerites, each apically curved, sharply pointed, and with small median projection (Figs 64, 65); tergite VIII arcuate apically (Fig. 66); sternite VIII elongate, parabolic apically (Fig. 67). Female. Spermatheca: capsule club-shaped, tubular basally and rounded apically, with small apical invagination; stem sinuate and coiled posteriorly (Fig. 70); tergite VIII arcuate apically and truncate medially (Fig. 68); sternite VIII rounded apically and truncate medially, antecostal suture distinctly sinuate (Fig. 69). Spermatheca of this species is very similar to that of B. neoinsecuta (Figs 79, 80), but shape of apex of female sternite VIII are different in the two species (Figs 69, 78).

Figures 63–70. 

Boreophilia venti (Lohse): 63 habitus 64 median lobe of aedeagus in lateral view 65 median lobe of aedeagus in dorsal view 66 male tergite VIII 67 male sternite VIII 68 female tergite VIII 69 female sternite VIII 70 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Holarctic species, recorded from Europe, Finland; Asia, East and West Siberia, Mongolia; and North America: Canada: YT; USA: AK [new record].

Habitat [new data]: Salix litter; Salix tundra hillside, lakeshore debris. Collecting methods: sifting Salix litter, pitfall traps, hand/aspirator collecting from under rocks. Collecting period: May and July.

USA, Alaska, Anaktuvuk Pass, 640–680 m el., 68.14049N, 151.74091W,+/- 2 km in and around village, hand, forceps, 2–22.V.2016, D Sikes, K Daly UAM100427681 (UAM) 1 female; Anaktuvuk Pass, 665 el., 68.14059N, 151.74266W, +/- 200 m, Salix, tundra hillside, cobble, pollinator pitfall, 20.V.2016, D Sikes, K Daly, UAM100427683 (UAM) 1 male; UAM100427684 (UAM) 1 female; UAM100427689 (UAM) 1 female; UAM100427693 (UAM) 1 male.

Lohse, in Lohse et al. 1990, described Dimetrota venti from Yukon. This species is extremely similar externally and genitally to Boreophilia insecuta described from Europe. The genitalia of B. insecuta were illustrated by Brundin (1954), and here, based on a specimen from Siberia (Figs 82, 83).

In B. venti, the tubus of the median lobe is distinctly arcuate in lateral view (Fig. 64), but in B. insecuta it is always straight basally (Fig. 82) and the two main sclerites are slightly different in shape (Figs 64, 65, 82, 83). However, the remaining genital characters are similar and females of the two species are not distinguishable morphologically.

(male): USA, Alaska, North Slope, Atkasuk, 17.VII.1978, B Vogel coll., B. insecuta det. Lohse (CNC).

USA, Alaska, Anaktuvuk Pass, 647 m el., 68.14049N, 151.74091W, +/- 250 m, under rocks, forceps/aspirators, 19.V.2016, D Sikes, K Daly, UAM100413204, UAM100413205, UAM100413207 (UAM) 3 females; Anaktuvuk Pass, 640–680 m el., 68.14049N, 151.74091W, +/- 2 km in and around village, hand/forceps, 20–22.V.2016, D Sikes, K Daly, UAM100388381 (UAM) 1 female.

Figures 71–80. 

Boreophilia neoinsecuta Klimaszewski: 71 habitus 72, 73 median lobe of aedeagus in lateral view 74 median lobe of aedeagus in dorsal view 75 male tergite VIII 76 male sternite VIII 77 female tergite VIII 78 female sternite VIII 79, 80 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Derived from prefix neo- added to existing specific name insecuta, a closely related species.

Body moderately broad, subparallel, forebody moderately glossy, abdomen slightly more so (Fig. 71); length 3.2–3.5 mm; head, pronotum and abdomen except for its apex dark brown, elytra reddish brown, mottled with some darker spots, appendages reddish brown; antennomeres VIII–X subquadrate; pronotum ca. as long as elytra at suture, maximum width of pronotum ca. equals to maximum width of elytra. Male. Tubus of median lobe of aedeagus almost straight laterally, apex narrow and rounded, bulbus broad and with two elongate and posteriorly bent sclerites (Figs 72, 73), in dorsal view median lobe elongate oval (Fig. 74); tergite VIII arcuate apically (Fig. 75); sternite VIII elongate, parabolic apically (Fig. 76). Female. Spermatheca: capsule club-shaped, moderately long and rounded apically, with small apical invagination; stem sinuate and coiled posteriorly (Figs 79, 80); tergite VIII arcuate apically (Fig. 77); sternite VIII produced apically and pointed medially, antecostal suture slightly sinuate laterally (Fig. 78).

Nearctic, Canada: MB, YT: USA: AK.

Habitat: tundra, under rocks. Collecting methods: forceps/aspirator. Collecting period: May to July.

Lohse, in Lohse et al. (1990) reported Boreophilia insecuta (Eppelsheim) in North America from AK, MB, YT, as a Holarctic species. However, these records represented a different and undescribed Nearctic species, which is here described as B. neoinsecuta sp. n. The two species, B. insecuta and B. neoinsecuta, have similarly shaped genitalia, which explains why they were confused. The true B. insecuta (Eppelsheim) was illustrated by Brundin (1954), and here (Figs 82, 83), and has a distinctly dilated apex of median lobe of aedeagus in lateral view (Fig. 82), while it is narrower in B. neoinsecuta (Figs 72, 73), and the two sclerites of internal sac of median lobe of aedeagus are broader and differently shaped than those of B. neoinsecuta (Figs 82, 83). Spermathecae of both species are very similar in shape, but female sternite VIII in B. insecuta is apically rounded and truncate medially, while in B. neoinsecuta is triangularly produced apically and pointed medially (Fig. 78).

Figures 81–83. 

Boreophilia insecuta (Eppelsheim) [specimen from Siberia, Russia]: 81 habitus 82 median lobe of aedeagus in lateral view 83 median lobe of aedeagus in dorsal view. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Our data included two sequences of B. neoinsecuta paratypes, both from Alaska which grouped into BIN BOLD:ADR7545. These sequences are 0.00% distant from each other and BOLD reports they are 7.23% distant from their nearest neighbor.

USA, Alaska, Bering Land Bridge N. Pk., 413 m el, 65.83713N, 164.58995W, +/- 30 m snowfield, tundra, under rocks, on moss, 9.VII.2016, DS Sikes et al., UAM100418913 (UAM).

all labelled the same except: UAM100418886 (UAM) 1 male, UAM100418887 (UAM) 1 female, UAM100418892 (UAM) 1 female, UAM100418905 (UAM) 1 male, UAM100418912 (UAM) 1 female, UAM100418905 (UAM) 1 male.

Named after Danish explorer Vitus Bering, whose name is shared with the species’ type locality, Bering Land Bridge National Park, and ‘Beringia’, the area of adjacent Russia and Alaska that were previously connected multiple times during the past 1 million years.

Body narrow, subparallel, glossy, abdomen slightly more so; microsculpture of forebody strong (Fig. 84); length 2.2–2.4 mm; head, head, pronotum and abdomen dark brown, elytra with reddish brown tinge, legs paler, yellowish brown; antennomeres VIII–X subquadrate; pronotum slightly longer than elytra at suture, maximum width of pronotum ca. the same as maximum width of elytra. Male. Tubus of median lobe of aedeagus straight basally and produced ventrad apically in lateral view, apex triangular in shape, distinctly pointed ventrad (Fig. 85), bulbus broad in dorsal view and with two elongate sclerites, strongly narrowed at one end as illustrated (Fig. 86); tergite VIII broadly arcuate apically (Fig. 87); sternite VIII elongate, narrowly rounded apically (Fig. 88). Female. Spermatheca: capsule club-shaped, moderately long and rounded apically, with small apical invagination; stem sinuate and coiled posteriorly (Fig. 91); tergite VIII arcuate apically (Fig. 89); sternite VIII broadly rounded apically, antecostal suture strongly sinuate medially (Fig. 90).

Figures 84–91. 

Boreophilia beringi Klimaszewski and Brunke, sp. n.: 84 habitus 85 median lobe of aedeagus in lateral view 86 median lobe of aedeagus in dorsal view 87 male tergite VIII 88 male sternite VIII 89 female tergite VIII 90 female sternite VIII 91 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Nearctic, USA: AK.

Habitat: snowfield, tundra, under rocks, on moss. Collecting methods: aspirating from moss. Collecting period: July.

We here compared Palaearctic Boreostiba piligera (J. Sahlberg), two males from Finland (ZMUO, NHMD), with our new species from Alaska. The two species are very similar externally but may be distinguished by the different morphology of the median lobe of aedeagus. In B. piligera, the apical part of the tubus is more weakly deflexed ventrad and distinctly more elongate. The sclerites of the internal sac are also more elongate and quite differently shaped (less like a talon and more even in thickness along their length). Therefore we conclude that these two populations represent sibling species.

Body moderately broad, strongly glossy, abdomen slightly more so (Fig. 92); length 2.8–3.5 mm; black, elytra with some reddish tinge, tarsi yellowish brown; antennomeres VIII-X slightly elongate; pronotum shorter than elytra at suture, maximum width of pronotum distinctly less than maximum width of elytra. Male. Tubus of median lobe of aedeagus almost straight and slightly produced ventrad in lateral view, apex broad and rounded (Fig. 93), bulbus broad and with two elongate sclerites of distinctive shape (Figs 93, 94); tergite VIII truncate or slightly concave apically (Fig. 95); sternite VIII elongate, parabolic apically (Fig. 96). Female. Spermatheca: capsule broadly club-shaped, moderately long and rounded apically, with small apical invagination; stem sinuate, straight medially, and looped posteriorly (Fig. 99); tergite VIII arcuate apically (Fig. 97); sternite VIII triangularly produced apically and pointed medially, antecostal suture arcuate, straight medially (Fig. 98).

Figures 92–99. 

Boreophilia subplana (J Sahlberg): 92 habitus 93 median lobe of aedeagus in lateral view 94 median lobe of aedeagus in dorsal view 95 male tergite VIII 96 male sternite VIII 97 female tergite VIII 98 female sternite VIII 99 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Holarctic species, recorded from Spitsbergen, Fennoscandia, Russia (west and east Siberia); Canada: NT, NU; USA: AK, NH.

Habitat: Salix leaf litter, tundra hillside, Black spruce forest, brackish shoreline, under rocks, wrack. Collecting methods: Malaise traps, aspirating from under rocks/cobbles, sweeping low vegetation, pitfall traps. Collecting period: June to August.

NEARCTIC: Canada, NT, Muskox L., NWT, 64.45N, 108.10W, 2.VIII.1953, JG Chillcott, Boreophilia subplana Sahlb. Det. GA Lohse (MHNG) 1 male; Spence Bay, NWT, 2.VII.1951, Boreophilia subplana Sahlb. Det. GA Lohse (MHNG) 1 sex?

USA, Alaska: Anaktuvuk Pass, 647 m el., 68.14049N, 151.74091W, 19.V.2016, +/- 250 m, Salix leaf litter, Berlese funnel, D Sikes, K Daly, UAM100432806, UAM100432826, UAM100432833, UAM100431905, UAM100431909, UAM100413051, UAM100413054, Salix, sweeping, UAM100427774 (UAM) 4 males, 4 females; Anaktuvuk Pass, 665 m el., 68.14059N, 151.74266W, 20.V.2016, +/- 200 m, Salix, tundra hillside, pollinator pitfall, UAM100427686, UAM100427688, UAM100427690, UAM100427691, UAM100427692 (UAM) 2 males, 3 females; Tetlin NWR, Alaska Hwy, 63.36124N, 141.96761W, +/- 10 m, 13–24.VII.2015, Black spruce, Malaise, J Hagelin, UAM100391356 (UAM) 1 male; Tetlin NWR, Alaska Hwy site 2, 523 m el, 63.08456N, 141.96761W, +/- 10 m, 13–24.VII.2015, Black spruce, hanging Malaise, J Hagelin, UAM100390752 (UAM) 1 female; Kotzebue, 1 m el, 66.89151N, 162.5933W, +/- 100 m el, brackish shoreline, under rocks, wrack, 6.VII.2016, D Sikes, K Daly, UAM100420026 (UAM) 1 female; Anaktuvuk Pass, Eleanor Lk., 649 m el., 68.14742N, 151.71854W, +/- 100 m lake shore, pond, sweep, dipnet, 20.V.2016, D Sikes, K Daly, UAM100427750, UAM100427751 (UAM) 2 males.

PALEARCTIC: Russia, Polarnyi Ural, c. Tobols. Gyub. [ernia], F Zajzew, 5.VI.1909, A. subplana, det. Benick (MHNG) 1 male.

Bernhauer (1907) described Atheta (Metaxya) angusticornis from Mount Washington, New Hampshire, USA. Gusarov (2003) transferred it to the genus Boreophilia, and considered it closely related to B. subplana, from which he differentiated it by the “shape of aedeagus, particularly a narrower apex of the median lobe in parameral view”. We have studied the median lobe of AK and NH specimens and found no differences warranting different species recognition. We therefore consider the two populations as belonging to the same species. The Mount Washington, NH, population represents a remnant, southernmost population of this species known only from higher elevations.

Our data included two sequences of B. subplana, both from Finland but because they are < 500 bp in length, they were not assigned to a BIN on BOLD. Our calculations indicate these sequences have an average and within-species maximum distance of 0.0% and are 6.37% distant from their nearest neighbor.

Body narrow, subparallel, moderately glossy, abdomen slightly more so (Fig. 100); length 3.5–3.8 mm; head, pronotum and abdomen, except for its apex, dark brown, elytra reddish brown, legs yellowish red-brown, or body uniformly piceous with tarsi and tibiae reddish brown; antennomeres VIII-X subquadrate (females) to slightly elongate (males); pronotum as long as elytra at suture or slightly shorter, maximum width of pronotum slightly less than maximum width of elytra. Male. Tubus of median lobe of aedeagus straight basally and strongly projecting ventrad at apex, apex broad and angular in lateral view (Fig. 101), in dorsal view bulbus broad and angular apico-laterally, with two elongate narrow sclerites of internal sac (Figs 102, 103); tergite VIII arcuate apically (Fig. 104); sternite VIII elongate, parabolic apically (Fig. 105). Female. Spermatheca: capsule pitcher-shaped basally with subspherical apical part bearing small invagination, stem sinuate, narrow, looped posteriorly (Fig. 108); tergite VIII arcuate apically (Fig. 106); sternite VIII rounded apically, antecostal suture straight medially and sinuate laterally (Fig. 107).

The spermatheca of B. caseyi was illustrated in Lohse et al. (1990). It is slightly deformed and based on a female captured in a different locality than that of the male holotype. It may belong to B. subplana, a species with very similar spermatheca. The spermathecal stem of B. caseyi has a broad posterior loop and female sternite VIII is rounded apically (Fig. 108), and not triangularly produced and pointed medially as in B. subplana (Fig. 99).

Figures 100–108. 

Boreophilia caseyi Lohse: 100 habitus 101 median lobe of aedeagus in lateral view 102, 103 median lobe of aedeagus in dorsal view 104 male tergite VIII 105 male sternite VIII 106 female tergite VIII 107 female sternite VIII 108 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Nearctic species, recorded from Canada: MB, NT, YT; USA: AK.

. Habitat: arctic tundra. Collecting methods: pitfall traps. Collecting period: June and July.

USA, Alaska, Toolik Lake Field Station, 724 m el., 68.6286N, 149.59772W, +/- 6m arctic tundra, 3 pitfalls, 2.VI–30.VII.2008, D Sikes UAM100044717, UAM100044680, UAM100044997 (UAM) 2 males, 1 female.

Lohse, in Lohse et al. (1990) described the new species B. manitobensis from MB and AK. The holotype from MB is represented by a male with a distorted median lobe of the aedeagus. We have studied the external and internal morphology of the two species, including the structures of internal sac, and found no significant differences between B. manitobensis and B. caseyi. Therefore, B. manitobensis is here synonymized with B. caseyi. The two species were published in the same paper, but B. caseyi has page priority and therefore was chosen as a valid species.

Four specimens of B. caseyi from UAM were submitted for DNA barcoding and three did not produce DNA sequences. The one which was successfully sequenced was flagged on BOLD as possibly contaminated so we excluded it from our analyses.

Body broad, narrowly oval, moderately glossy, abdomen slightly more so (Fig. 109); length 2.8–30 mm; uniformly dark brown to almost black, tarsi paler, yellowish-brown; antennomeres VIII-X subquadrate to slightly transverse; pronotum as long as elytra at suture, maximum width of pronotum distinctly shorter than maximum width of elytra. Male. Tubus of median lobe of aedeagus slightly arcuate in lateral view, apex narrowly rounded, bulbus broad and with two elongate narrow sclerites (Fig. 110), in dorsal view median lobe of aedeagus elongate oval (Fig. 111); tergite VIII truncate apically (Fig. 112); sternite VIII elongate, parabolic apically (Fig. 113). Female. Spermatheca: capsule broadly club-shaped, moderately long and rounded apically, with small apical invagination; stem short, sinuate and looped posteriad (Fig. 116); tergite VIII truncate apically (Fig. 114); sternite VIII rounded apically and truncate medially, antecostal suture arcuate and slightly sinuate laterally (Fig. 115).

Figures 109–116. 

Boreophilia vega (Fenyes): 109 habitus 110 median lobe of aedeagus in lateral view 111 median lobe of aedeagus in dorsal view 112 male tergite VIII 113 male sternite VIII 114 female tergite VIII 115 female sternite VIII 116 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

Holarctic species, known from West and East Siberia, Russian Far East, North Korea; and Canada: Herschel Island, YT. USA: not recorded.

Habitat: Yukon specimens were collected in an alluvial fan in June and July (Klimaszewski et al. 2012).

Boreophilia herschelensis is conspecific with B. vega and is here synonymized. Boreophilia vega has a median lobe of aedeagus similar to that of B. neoinsecuta (Fig. 72) and B. insecuta (Fig. 82), but the spermatheca of B. vega compared to the latter two species is differently shaped (Figs 79, 80, 116). Externally, B. vega differs from B. neoinsecuta in elytra distinctly broader that the maximum width of pronotum (Figs 71, 109). Female sternite VIII is differently shaped in the two species (Figs 78, 115).

Our data included one sequence of B. vega from Yukon Territory, Canada, but because this sequence was < 500 bp long it was not assigned a BIN on BOLD. We calculate that this sequence is 6.5 % distant from its nearest neighbor.

Body moderately broad, subparallel, moderately glossy, abdomen slightly more so (Fig. 117); length 2.8–2.9 mm; uniformly dark brown to almost black, basal sternite slightly reddish-brown, legs paler, yellowish-brown; antennomeres VIII–X subquadrate to slightly transverse; pronotum slightly shorter than elytra at suture, maximum width of pronotum distinctly less than maximum width of elytra. Male. Unknown. Female. Spermatheca: capsule ovoid apically and pitcher-shaped basally, without apical invagination; stem short, narrow, sinuate and looped posteriad (Fig. 120); tergite VIII truncate apically (Fig. 118); sternite VIII rounded apically, antecostal suture arcuate and slightly sinuate laterally (Fig. 119).

Figures 117–120. 

Boreophilia davidgei Klimaszewski and Godin: 117 habitus 118 female tergite VIII 119 female sternite VIII 120 spermatheca. Scale bars: 1 mm (for habitus); 0.2 mm (remaining).

This species may be distinguished by the unique shape of spermatheca.

Nearctic species, known only from Canada, YT.

Habitat: white spruce and feathermoss forest, mixed pine and willow forest, black spruce stand, mixed aspen and white spruce forest (Klimaszewski et al. 2012). Females from Yukon were collected in May to September using pitfall traps and litter sifting in mature forest.

This species is tentatively assigned to this group, because the male is unknown and morphology of median lobe of aedeagus could not be analysed.