Quedius molochinoides Smetana, 1965

Hansen, Aslak Kappel, Brunke, Adam, Simonsen, Thomas & Solodovnikov, Alexey, 2022, Revision of Quedius sensu stricto (Coleoptera: Staphylinidae), Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 62 (1), pp. 225-299 : 278-280

Quedius molochinoides Smetana, 1965

( Figs 1 , 4 , 5 , 10D,E , 16E , 17 , 21 , 22 )

Quedius molochinoides Smetana, 1965a: 35 [Type locality: Newfoundland: Ha-Ha Bay]

Quedius molochinoides miquelonensis Smetana, 1965a: 36 [Type locality: Miquelon], syn. nov.

References. Sආൾඍൺඇൺ (1971): 141 (characters and distribution); (1976): 174, (1978): 826, (1981): 640 (distribution); Dඈඐඇංൾ & Aඋඇൾඍඍ (1996): 391 (characters and distribution).

Type material examined. Quedius molochinoides : Hඈඅඈඍඒඉൾ: J (CNC), labelled: “NW Newfoundland Ha-Ha Bay N: o 103. 18.VII.49 Lindroth / Holotypus Quedius molochinoides m. / Quedius molochinoides m. 63 Smetana det. 19 / Holotype CNC No. 9474 Quedius molochinoides s. s. Smet. / CNC”. Pൺඋൺඍඒඉൾ (labelled as allotype): same collection data as holotype (1 ♀ CNC).

Quedius molochinoides miquelonensis : Hඈඅඈඍඒඉൾ: J (CNC), labelled: “ Miquelon N:o 353 8-10.VIII.51 Lindroth / Holotypus ”. Pൺඋൺඍඒඉൾ (labelled as allotype): “ Quedius molochinoides miquelonensis m. / 1963 Smetana det. / Quedius molochinoides miquelonensis m. Smetana det. 1963/ Allotype CNC No. 9475 Quedius molochinoides miquelonensis Smet. ” (1 ♀ CNC).

Additional material examined. CANADA:Aඅൻൾඋඍൺ: Castle river valley, SW corner of township, [49.49, -114.22], along the road, under rocks etc., 23.V.1990, leg. B.F.& J.L.Carr (1 CNC);Cirrus Mountain, Banff National Park, [52.16, -116.97], 1676 m, 16.VI.1968, leg. Campbell & Smetana (1 CNC); Crow’s Nest Pass, Rocky Mount, [49.59, -114.53] (1 CNC); Kananaskis Forest Experiment Station, [51, -115.1], 1.VIII.1971, leg. J. M. & B.A. Campbell (1 CNC); Mile S Seebe, [51.09, -115.06], 26.VII.1968, leg. A. G. Raske (1 CNC); Tp. 5 Rge. 3 W. 5 Mer, [49.39, -114.34], leg. 23.VI.1994, leg. B. F. & J. L. Carr (1 CNC); Waterton, [49.05, -113.9], 10.VII.1930, leg. Washur (1 CNC); Waterton Lks. Nat. Pk., Cameron Lk., 49.0833, -113.9164, 12.VI.1973, leg. J. Redner & C. Starr (1 CNC); Waterton Lakes National Park, picnic area on Cameron Creek near Cameron Lake, [49.05, -113.98], 1630 m, pitfall trap, 7-11.VI.1980, leg.I. M.Smith (5 CNC); Waterton Lake National Park, Cameron Lake, [49, -114.04], berlese trap, grassy tussock, seepage area, 7.VI.1980, leg. I. M. Smith (1 CNC);Waterton Lake National Park, Crandell Lake Trail,[49.06,-113.94], berlese trap, Alder litter, 13.VI.1980, leg. I. M. Smith (1 CNC); Waterton Lakes National Parks, Lower Waterton Lake, 49.09, -113.85, 1370 m, ex moss, 26.VI.1980, leg. J. M. Campbell (7 CNC);Waterton Lakes National Park, Cameron Lake, [49, -114.04], pitfall trap, wet meadow, 8.VI.1980, leg.I. M.Smith (14 CNC); Waterton Lakes National Park, Cameron Creek picnic ground,[49.04, -113.98], pitfall trap, edge meadow, 17-28.VI.1980, leg. I. M. Smith (2 CNC); Waterton Lakes National Park, Maskinonge Lake,[49.1,-113.83], 1310 m, sifting litter, Populus litter, 14.VI.1980, leg. J. M. Campbell (1 CNC); Waterton Lakes National Park, near Highway 6, 4.4 km SE Junction Highway 5, [49.08, -113.79], pitfall trap, edge of beaver pond, 12-19.VI.1980, leg. I. M. Smith (2 CNC); Waterton Lakes National Park, Crypt Lake,[49, -113.84], 1981 m,under rocks, 22.VI.1980, leg. J. M. Campbell (1 CNC); Waterton Lakes National Park, near Lower Rowe Lake, [49.05, -114.05], berlese trap,moss from spring seepage area, 18.VI.1980, leg. I. M. Smith (4 CNC); Waterton Lakes National Park, Lookout Butte Picnic Site, [49.72, -113.56], flight interception trap, 9-14. VI.1980,leg. J. M.Campbell (1 CNC);Waterton Lakes National Park, Km 9 Chief Mountain Highway, [48.91, -113.43], 1371 m, flight interception trap, 7.VII.1980, leg. H. J. Teskey (38 CNC); Waterton Lakes National Park, near Highway 6 4.4 km SE Junction Highway 5, [49.08, -113.79], pitfall trap, in meadow by beaver pond, 7-11.VI.1980, leg. I. M. Smith (3 CNC); Waterton Lake National Park, Rowe Lakes Trail, [49.05, -114.06], 1828 m, under bark, 12.VI.1980, leg. J. M. Campbell (2 CNC); Waterton NP, Rowe Creek, 49.0833, -113.9164, 1664 m, 2.VIII.1976, leg. J. M. Campbell (1 CNC). BඋංඍංඌH Cඈඅඎආൻංൺ: Amiskwi River, Yoho National Park, [51.38, -116.55], 7.VIII.1971, J. M. & B. A. Campbell (1 CNC); Bowron Lake, [53.23, -121.37], 15.X.1953, leg. G. S. Smith (1 J CNC); Creston, [49.09, -116.51], 29.VII.1956, leg. G. S. Smith, (1 CNC); 8 mi W Creston, [49.09, -116.72], ex river debris, 10.VI.1968, leg. Campbell & Smetana, (1 CNC); 21 mi W Creston, [49.09, -117.04], ex river debris, 9.VI.1968, leg. Campbell & Smetana (1 CNC); Fish Lake, Summerland, [49.84, -115.44], 27.X.1931, leg. A. N. Gartell (2 CNC); Highway 37, 30km S Dease Lake, [58.43, -129.99], 2.VII.1987, leg. L. Carr (2 CNC); Highway 37A, Bear Glacier, [56.09, -129.66], 6-7.VII.1987, leg.Larson & Carr (1 CNC); Koutenay National Park near Dog Lake, [50.78, -115.92], 24.V.1981, leg. I. Askevold (1 CNC); Mt. Revelstoke, [51.09, -118.04], 1737 m, under rocks in swamp, 1.VII.1952, leg. G. J. Spencer (1 CNC); Mount Revelstoke, [51.09, -118.04], 10.VII.1952, leg. G. J. Spencer (1 CNC); Radium, [50.62, -116.07], 26.IV.1994, leg. B.F. & J. L. Carr (1 CNC); Sinclair Creek, Kootenay National Park, [50.63, -116.06], 1828 m, 12.VI.1968, leg. Campbell & Smetana (1 CNC); Summerland, [49.6, -119.67], 27.X.1931, leg. A. N. Gartell (1 CNC); Trinity Valley, [50.4, -118.91], under old log, 16.IX.1943, leg. H. B. Leech, (1 CNC); Vernon, [50.31, -119.28], under stone, 18.IX.1939, leg.H.Leech (1 CNC);Vernon, Deep Lake, [50.79, -120.34], rotten log, hibernating, 10.IV.1989, leg. H. Leech, (1 CNC); Vernon, Monashee Mountains, [50.27, -119.27], 1371 m, under stone, 14.X.1934, leg. H. Leech (1 CNC). Oඇඍൺඋංඈ: 12 mi NE Moosonee, [51.44, -80.45], 1.VII.1973, leg. R. Parry & J. M. Campbell (1 CNC); Gregoire Mills, [49.29, -81.95], 22.VI.1973, leg. J. Redner & C. Starr (1 CNC). YඎKඈඇ: Mile 11 Canol River, [61.98, -132.45], 1219 m, 22.VIII.1962, leg. P. J. Skitsko (1 CNC); Slim’s River km 1705 Alaska Highway, [61.01, -138.51], ex marsh, 7.VII.1981, leg. Bright (1 CNC). Qඎൾൻൾർ: Lac Mitis, 48.30, -67.80, 9.-12.VII.1993, leg. Hébert & Jobín (1 LFC); Parc Gaspesie (Gaspe-O), Mont Logan, [48.89, -66.64], 1040 m, Spruce forest clearing with moss and ericaceae,pitfall traps, 2-14.VII.1993, leg. F. Landry (1 CNC). USA: AඅൺඌKൺ: Becharof NWR, Island Arm East, 57.76107, -155.98752, S facing slope, Salix , fern, & cow parsnips, pitfall traps, 29.VI.-5.VII.2007, leg. Savage (4 UAM); Chichagof Island, 57.98256, -135.21811, 970 m, Dryas , Salix, Cladonia portentosa, ground search, 15.VII.2013, leg. Ridling (1 UAM); Chichagof Island, 57.98708, -135.20357, 702 m, Nephrophyllidium crista-galli , Geranium erianthum , Artemisia arctica , pitfall traps, 14.-15.VII.2013, leg. Ridling (7 UAM); S Chilkat Peninsula, 58.42520, -135.15205, 687 m, subalpine, Vaccinum, Cornus, pitfall traps, 11.-12.VII.2010, leg. D. Sikes (1 ♀ UAM); Haines, Flower Mountain, 59.39353, -136.31129,alpine Sanguisorba,Carex , pitfall traps, 15.-16.VII.2010, leg. D. Sikes (3 UAM); Hope, Kenai Peninsula, [60.89, -149.65], 27.VII.1951, leg.W.J. Brown (1 CNC); Kenai Mountain CreekAbove Tern Lake Campground,[60.53, -149.53], 274 m, 13.VI.1978, leg. Smetana & Becker (1 CNC); Kenai Mountain, 2 mi S Moose Pass, [60.44, -149.37], 182 m, 30.V.1978, leg. Smetana & Becker (1 CNC); Kenai Mountain, 22 mi N Seward, [60.44, -149.45], 152 m, 27.V.1978, leg. Smetana & Becker (2 CNC); Lituya Bay, Cenotaph Island, [58.64, -137.57], 182 m, 22 Pitfalls in Alder Woods, 25.-26.V.1977, D. Mann, (1 CNC); Moose Pass, Kenai Peninsula, [60.49, -149.42], 16.VI.1951, leg. W. J. Brown (2 CNC); Mount Hawthorne, 58.23652, -134.23904, 748m, subalpine meadow, low ridge top, baited pitfall traps, 22.-23.VII.2013, leg. Ridling (4 UAM); Palmer, 61.60, -149.10, pasture, muskox dung, 1.VIII.1988, leg. Pilny (1 DEBU); Seward [60.12, -149.39], 14.V.1951, leg.W.J. Brown (2 CNC); Skagway, 59.50, -135.10, moist pasture,cow & horse dung, 15.VII.1988, leg.Pilny (1 DEBU);Valdez, [61.14, -146.35], on snow, 29.IV.1972, leg.Dick (1 UAM);Valdez, 61.1167, -146.2667, Picea & Populus forest, under wood and stones,wet vegetated gravel, 10.VIII.1988, leg.Pilny(2 DEBU). IൽൺHඈ: Moscow,[46.80,-116.85], 14.IV.1916,leg.A. C. Burrill (1 CNC). Nൾඐ HൺආඉඌHංඋൾ: Mount Washington, [44.27,-71.31], 1874, leg. Blanchard (4 MCZ); White Mountains, Peabody River, [44.39, -71.15], 1828 m, 12.VII.1925, leg. Quirsfeld (1 CNC). Nൾඐ YඈඋK: Indian Falls,Mount Marcy,[44.11, -73.92], 2.IX.1942, leg.H.Dietrich (1J CNC); Lake Tear, Mount Marcy, Essex County, [44.11, -73.93], 27.VII.1922, leg. H. Notman (1 CNC).

Redescription. Measurements JJ (n = 2): HW = 1.56– 1.71 (1.63); HL = 1.36–1.44 (1.40); HL/HW 0.84–0.87 (0.86); PW = 2.00–2.22 (2.11); PL = 1.91–2.00 (1.96); PL/PW 0.90–0.96 (0.93); EW = 2.02–2.36 (2.19); EL = 1.84–2.29 (2.07); EL/EW 0.91–0.97 (0.94); EL/PL 0.97– 1.14 (1.05); PW/HW 1.48–1.54 (1.51); forebody length 5.11–5.73 (5.42). ♀♀ (n = 5): HW = 1.69–1.89 (1.77); HL = 1.38–1.56 (1.45); HL/HW 0.81–0.83 (0.82); PW = 2.20–2.38 (2.26); PL = 2.00–2.13 (2.07); PL/PW 0.89–0.93 (0.92); EW = 2.22–2.49 (2.36); EL = 2.09–2.22 (2.18); EL/EW 0.88–0.96 (0.93); EL/PL 1.02–1.11 (1.06); PW/ HW 1.52–1.63 (1.55); forebody length 5.53–5.87 (5.70).

Medium sized species; body dark brown to black ( Figs 10D,E ).

Head black, distinctly transverse with eyes medium sized (EyL/TL = 1.76–1.94 (1.87)), microsculpture of transverse waves, with two interocular punctures between anterior frontal punctures ( Fig. 6G ); antennae and palpi dark with joints reddish pale, all antennomeres elongate, except last and penultimate antennomere almost transverse.

Thorax: pronotum dark brown to black, slightly wider than long, clearly wider than head, with microsculpture of transverse waves, three punctures in dorsal row and one to two in sublateral row with its posteriormost puncture reaching just beyond level of first puncture of dorsal row; scutellum sparsely punctured and pubescent; elytra variable in coloration, from all black to reddish brown, occasionally dark with reddish sutural margins, uniformly pubescent, with punctures clearly separated, slightly wider than long, same length to slightly longer than pronotum; legs reddish brown with inner face of tibia and inner femur darkened and tarsi paler.

Abdomen dark brown to black, tergites uniformly punctured, with clear iridescence.

Male. Aedeagus ( Figs 16E , 17 ): paramere lanceolate, rather broad, parallel-sided or with slight basal attenuation, reaching apex of median lobe, with small sensory peg setae in two irregular rows meeting below apex to form a slightly pointed ‘arrowhead’ shape; median lobe lanceolate broad, sides slightly constricting until abrupt constriction near pointed apex, on parameral side with two large pointed teeth folded from lateral margin to form almost closed cavity; internal sac with two larger oval and two smaller rounder sclerites.

Differential diagnosis. Quedius molochinoides can be distinguished from Q. laticollis and Q. strenuus by the presence of two additional setiferous punctures between the anterior frontal punctures. It can be distinguished from Q. labradorensis by the darker body including the clearly darkened basal three antennomeres. It can be distinguished from other members of the molochinoides -group ( Q. altanai , Q. horni , and Q. lanei ) only by the male genitalia, specifically by the broad median lobe with two large, flattened pointed teeth strongly folded to form an almost closed cavity. Quedius molochinoides can also be accidentally confused with Q. brunnipennis Mannerheim, 1843 from the subgenus Raphirus , but will easily be recognized by the entire labrum and by more parallel-sided abdomen.

Synonymic notes. The subspecies Q. m. miquelonensis was described from the islands of St. Pierre and Miquelon as a slightly broader and larger form with the male sixth sternite wider and more sharply triangular (Sආൾඍൺඇൺ 1971a) compared to the nominal subspecies. Generally, species within Quedius s. str. display notable intraspecific variation in body size. With the new material examined here the observed variation between the two subspecies falls within the limits of the variation observed within the widely distributed nominal subspecies. Thus, we consider Quedius molochinoides miquelonensis Smetana, 1965 syn. nov. a junior subjective synonym of Quedius molochinoides Smetana, 1965 .

Comments. One of the paratypes of Q. lanei from Fishlake in Summerland of British Columbia, Canada was found to belong to Q. molochinoides . Specimens from the Okanagan area (Summerland, Copper Mountain and Vernon) are slightly different from the nominal species of Q. molochinoides and others of Q. molochinoides- group 3, specifically their genitalia being more slender. Images of the genitalia for two of these are provided in the supplementary material ( Fig. S5 ). We are hesistant to describe them as a new species as we are lacking genetic barcodes as evidence for this. That said they may very well represent yet another species new for science.

Bionomics. Quedius molochinoides prefers meadows or open forests where it has mainly been collected in litter, grass tussock or under rocks from low elevations up to 2000 m. Specimens from Newfoundland were found on more or less open ground with rich vegetation or in leaf litter under bushes near river edges (Sආൾඍൺඇൺ 1971a), in British Columbia some specimens were collected in river flood-debris and by sifting wet debris and fallen leaves under bushes at mountain creeks, other specimens were found under rocks near snow in Kootenay National Park (Sආൾඍൺඇൺ 1971a). In Alaska the species was collected using pitfall traps in subalpine meadows and shrublands. Immature specimens were collected in July in Alberta and Idaho and from July to September on Mt. Washington in New Hampshire (Sආൾඍൺඇൺ 1971a).

Distribution. Quedius molochinoides is a northern species distributed transcontinentally across boreal North America from Labrador and Newfoundland to Alaska (ecoregions 4, 5, 6.1 and partly 6.2 and 7.1, Fig. 21 ). To the south it is found at higher elevation sites, e.g. in the northern Rockies in the west (north part of ecoregion 6.2) and Mount Washington and Mount Marcy in the East (ecoregion 5.3). The northernmost records are from Valdez in Alaska and Churchill in Manitoba, suggesting that the species extends throughout the boreal ecoregion of the continent. There is a single record from Moscow Mountains in Central Idaho, which represents the southernmost record in the Rockies (ecoregion 6.2.3). COI barcoding showed that Q. molochinoides forms three distinct clusters ( Figs 4 , 5 )). Quedius molochionide s-group 1 is known from the Canadian national parks of Banff and Waterton Lakes and is delimited by the Rocky Mountains Trench to the west and the Canadian plains to the East (part ecoregion 6.2.4, Fig. 22 ). Quedius molochionides -group 2 is found in the Canadian Rockies, west of the Rocky Mountains trench and towards the north continuing to the Pacific Coast in Alaska all the way to the Kenai Peninsula northwards (ecoregion 6.2.1, 6.2.2, 6.2.3 and partly 6.1 and 7.1). Quedius molochinoides -group 3 is present from the Becharof Peninsula of Alaska in the west across the North American boreal region to glacial refugia of New England, Quebec and Newfoundland in the east (ecoregions 4, 5, and partly 6.1 and 7.1).

Figures

Fig. 1. Phylogenetic tree of Quedius s. str. based on preliminary target capture molecular dataset of Bඋඎඇĸൾ et al. (2021) and Hൺඇඌൾඇ et al. (in prep). Species groups shown in color. Q. altanai Hansen & Brunke, sp. nov. of the Q. molochinoides-group was not included in the dataset. Habitus sketch is of Q. fuliginosus (Gravenhorst, 1802). Lettering denotes species distribution within their respective groups.

Fig. 4. Phylogeny of Quedius s. str. species based on Bayesian Inference (BI) and Maximum Likelihood (ML) COI barcodes (part 2). BI posterior probabilities (PP), ML ultrafast bootstrap support (UFB) and ML SH-aLRT test are shown. Nodes coloured as: green, PP ≥ 0.90, and either UFB ≥ 95 or SH-aLRT ≥ 80; yellow, one of PP ≥ 0.90, UFB ≥ 95 or SH-aLRT ≥ 80; red, PP <0.90, UFB <95 and SH-aLRT <80.

Fig. 5. Network analysis of Quedius molochinoides-group: Quedius horni Hatch, 1957 (purple), Q. lanei Hatch, 1957 (green), Q. molochinoides-group 1 (dark blue), Q. molochinoides-group 2 (light blue), and Q. molochinoides-group 3 (teal).

Fig. 10. Habitus of Quedius s. str. A – Q. horni Hatch, 1957; B – Q. lanei Hatch, 1957; C – Q. altanai Hansen & Brunke, sp. nov.; D – Q. molochinoides Smetana, 1965 (red elytra variant); E – Q. molochinoides (black elytra variant); F – Q. labradorensis Smetana, 1965.

Fig. 16. Aedeagus of Quedius s. str. A – Q. labradorensis Smetana, 1965; B – Q. horni Hatch, 1957; C – Q. altanai Hansen & Brunke, sp. nov.; D – Q. lanei Hatch, 1957; E – Q. molochinoides Smetana, 1965. I – parameral view of whole aedeagus, II – apices of paramere in antiparameral view with peg setae, III – lateral view of whole aedeagus, IV – apices of median lobe at an angle between parameral and lateral view.

Fig. 17. Variation in genitalic morphology of species from Quedius molochinoides-group.From left to right for each specimen, first paramere then median lobe (parameral view) and lastly median lobe (lateral view) are given. For each specimen locality is noted underneath.

Fig. 21. Distribution of Quedius unicolor Kiesenwetter, 1847 (top),.Q. labradorensis Smetana, 1965 (middle), Quedius molochinoides-group (bottom). Barcoded specimens are denoted with +.

Fig. 22. Distribution of Quedius molochinoides Smetana, 1965 (top), and Q. laticollis (Gravenhorst, 1802) and Q. strenuus Casey, 1915 (bottom).

Fig.6. Head and pronotal chaetotaxy of Quedius s. str.A – general scheme; B – Q. curtipennis Bernhauer, 1908, C – Q. fuliginosus (Gravenhorst, 1802); D – Q. levicollis Brullé, 1832; E – Q. afrofuliginosus Gusarov, 1991; F – Q. molochinus- and Q. laticollis-groups; G – Q. molochinoides-group. Abbreviations as follows: anterior frontal puncture (AFP); basal punctures (BP); interocular punctures (IOP); lateral puncture (LP); lateral row (LR); posterior frontal puncture (PFP); parocular punctures (POP); sublateral row (SLR).