Quedius curtipennis Bernhauer, 1908

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 : 241-248

Quedius curtipennis Bernhauer, 1908

( Figs 1 , 3 , 6B , 7B , 11C , 12 , 18 )

Quedius curtipennis Bernhauer, 1908 e: 335 [Type locality: Faroe; Romania; Buchara; Böhmen, Wran a. Moldau].

Quedius gracilis Stephens, 1832: 215 [Type locality: London], syn. rev. Quedius parallelus Hatch,1957:216 [Type locality:Washington, Seattle]

References. Gඋංൽൾඅඅං (1924): 79 (characters), SർHൾൾඋඉൾඅඍඓ (1933): 1441 (characters), Hൺඇඌൾඇ (1952): 141 (characters); Sආൾඍൺඇൺ (1958): 362 (characters and ecology); (1962): 133, (1965b): 38 (characters); (1971a): 129 (characters, synonymy of Q. parallelus ); (1965a): 12, (1970): 62, (1990): 98, (1993): 50 (distribution); (1978a): 826 (biology); KඈඋGൾ (1962b): 332 (suspected synonymy of Q. parallelus ); (1964): 119 (distribution); Pൺඅආ (1963): 141 (characters); LඈHඌൾ (1964): 211 (characters); Dඏඈෞගĸ (1965): 89 (distribution); Hඈඋංඈඇ (1965): 272 (distribution); Sඓඎඃൾർĸං (1968): 735 (distribution); Fඋൺඇĸ (1969): 267 (biology); Bඈඋൽඈඇං (1974b): 10, (1976b): 92 (characters); Pඈඉൾ (1977): 31 (distribution); Cඈංൿൿൺංඍ (1978): 191 (characters); Oඎඍൾඋൾඅඈ (1978): 281 (pupal characters); Bඎඋൺĸඈඐඌĸං et al. (1980): 134 (distribution); TඬඍH (1984): 118 (characters); DඋඎGආൺඇൽ (1987): 312 (distribution); Nඈඐඈඌൺൽ (1990): 145 (biology); BඈඋGൾඌ (1990): Table II (distribution); Cංർൾඋඈඇං & Zൺඇൾඍඍං (1995): 32 (distribution); Oඐൾඇ (1997): 149 (distribution); Oඎඍൾඋൾඅඈ et al. (1998): 129 (biology); AඌඌංඇG (2001): 75 (biology); UHඅංG et al. (2006): 54 (distribution); Tඋඈඇඊඎൾඍ (2006): 102 (distribution); Mൺඃĸൺ & Sආൾඍൺඇൺ (2007): 429 (characters and distribution); ÖඓGൾඇ (2011):204 (distribution); Bඋඎඇĸൾ & MൺඋඌHൺඅඅ (2011): 59 (characters and distribution); Wൾൻඌඍൾඋ et al. (2012): 313 (distribution); MൺGඎඋൺ et al. (2013): 719 (biology); TඬඍHආඣඋඣඌඓ et al. (2014): 688 (biology); Sൾආൾඇඈඏ et al. (2015): 127 (distribution); Sൾආංඈඇൾඇĸඈඏ et al. (2015): 330 (distribution); Sൺඅඇංඍඌĸൺ & Sඈඅඈൽඈඏඇංĸඈඏ (2018a): 125 (notes on types and distribution); (2019): 48 (distribution).

Type material examined. Quedius curtipennis : Lൾർඍඈඍඒඉൾ: (J, here designated): “Suderö Faroer Ins./ Dr. Cornu 1907/ v. curtipennis Brh. Typus / fuliginosus Grav. Scheerp. / Chicago NHMus M. Bernhauer Collection / D. Drugmand det. 1994 Quedius (s. str.) curtipennis Brnh. / Lectotype Quedius curtipennis Bernhauer, 1908 A.K. Hansen & A. Solodovnikov des. 2021” (FMNH). Pൺඋൺඅൾർඍඈඍඒඉൾඌ: “Nördl. Faroer Ins. / Dr. Cornu 1907 / v. curtipennis Brh. Typus / fuliginosus Scheerp. [sic!] det. [illegible] / Chicago NHMus M. Bernhauer Collection / D. Drugmand det. 1994 Quedius (s. str.) curtipennis Brnh / Paralectotype Quedius curtipennis Bernhauer, 1908 A.K. Hansen & A. Solodovnikov des. 2021” (1 J FMNH); “ v. curtipennis Buchara Bang Haas det. Bernh. / Chicago NHMus M. Bernhauer Collection” (1 J FMNH).

Quedius parallelus : Hඈඅඈඍඒඉൾ: “Seattle, Wash. U.W.Campus IV-12- 1949 / Del. 1954 H. Houk / Type J Quedius (s. str.) parallelus - 1951 M. Hatch / Quedius (s. str.) fuliginosus Grav.M. Hatch – 1947” (1J UWBM). Pൺඋൺඍඒඉൾඌ: “Seattle,Wash. April 12, 1934 ” (1J UWBM); “Seattle,Wash. U.W.Campus, IV-11-1947 ” (1 J UWBM, N. Vowles coll.); “Vancouver, B.C., 5 March, 1949, G.B. Rich” (1 J CNC); “Wash. Seattle, Feb. 15, 1954, B. Malkin” (1 ♀ FMNH, ex coll. B. Malkin).

Additional material examined. ARMENIA: Arzakanskaja dolina, Aghveran, [40.51, 44.56], 3.VII.1978, leg. R. Rous (1 ♀ MCZ); NE Dilizhan nr. Haghartsin Monastery, 40.8013, 44.8919, 1450 m, 21.V.2001, leg. Shaverdo & Schillhammer (4 JJ NMW); 18 km E Dilizhan, 3 km Gosh, 40.7405, 45.0191, 1850 m, 27.V.2001, leg. Shaverdo (1 J NMW); 21 km N Hrazdan, Ankavan, 40.6286, 44.5533, 1000 m, 22.V.2001, leg. Shaverdo & Schillhammer (1 J NMW); Teghut, Akhtala, 41.0901, 44.8120, 1009 m, 28.VI.2016, leg. J. Müller (1 J cSch); same locality, 41.0911, 44.8131, 1020m, 20.V.2016, leg.A. & J. Müller (2♀♀ ZMHB). AUSTRIA: Bisamberg, [48.33, 16.36], leg. Luze (1 ♀ NMW); S Steiermark, [46.91, 15.60] (1 J NMW); Tirol, Musau, [47.53, 10.67], litter, moss, and grass, 28.C.1995, leg. M.Uhlig (1 J ZMHB); Wolfpassing, [48.08, 15.06], VI.1965, leg. Wewalka (1 ♀ NMW). BULGARIA: Rila Monastery, [42.13, 23.34], 1150 m, 14.VII.1965, leg. P. Jørum (1 J NHMD). BOSNIA AND HERZEGOVINA: Zeleznica [Zeljeznica] n. Foinica, [43.89, 17.95], 500-700m, 13.IV.1969, leg. U. Heinz (1 J ZMHB). CANADA: BඋංඍංඌH Cඈඅඎආൻංൺ: Stanley Park, Vancouver, [49.30, -123.14], 9.VI.1954, leg. D. Lazorko (1 J 1 ♀ NMW). CZECH REPUBLIC: Brandeis, [50.18, 14.65], leg. Skalitzky (2 JJ NMW). DENMARK: Vigsø Bugt, 1 km E of Vigsø, 57.0950, 8.7394, pine forest, pitfall traps, 16.-18.VI.2014, leg.A.Hansen, M.Justesen,A.Solodovnikov & L.Kraemer (1 J NHMD) LඔඌØ: Laesø Klitplantage, 57.3082, 11.0081, mixed forest, sifting litter, 28.V.2013, leg.A. Solodovnikov & M.Justesen (1 NHMD). FAROE ISLANDS: Suderö [Suðuroy], [61.51, -6.83], 1907, leg. Cornu (1 J FMNH). FRANCE: Ain, Torcieu n. Amberieu, [45.91, 5.39] (1 J ZMHB); Bretagne, Locronan, [48.09, -4.21], VII.1983, leg. H.Korge (1 J ZMHB); Chigny n. Oise, [49.91, 3.76], leg. Gelchert (1 ♀ ZMHB); Morlaix, Bretagne, [48.57, -3.82], 1.VIII.1950, leg. A. Nilsson (2 JJ MZLU). GEORGIA: Borzhom Distr., Bakuriani, [41.75, 43.52], 27.VII.1947, leg. Bogachev (1 J 1 ♀ ZIN); Kartliskij Chreb. [Kartalinskiy Khrebet], Sabaduris Tye [Sabaduri Pass] [41.91, 44.91], 1800m, 8.VI.1987, leg. Wrase (1 J cSch); same locality, 25.VI.1988, leg. Wrase (1 J ZMHB); Trialetski, Khrebet Bakuriani, [41.74, 43.83], 1800-2200m, 4.-7.VII.1986, leg.Wrase & Schülke (8JJ 4♀♀ ZMHB); Zchneti [Tskneti], [41.69, 44.69], 1200m, 26.VI.1986, leg. Wrase (1 J cSch). GERMANY: Bayern, Garmisch, [47.48, 11.08], 27.IV.1932, leg. Ihssen (1 J ZMHB); Bonn, [50.72, 7.09], X, leg. Verhoeff (1 ♀ ZMHB); Bayern, Eschenlohe, [47.58, 11.18], 3.VIII.1937, leg.Ihssen (1J ZMHB); Brandenburg, Strausberg, [52.52, 13.85], 17.IV.1987, leg. M. Schülke (1 J ZMHB); Dresden, Kötschenbroda,[51.11, 13.61], moss, 23.I.1920, leg. Dahl (1 J ZMHB); Düsseldrof Unterbach, [51.20, 6.89], 27.VII.1974, leg. H.Bremer (1 ♀ ZMHB); Marburg, [50.80, 8.76], 16.IV.1904, leg. Strand (1 ♀ ZMHB); Sarstadt, [52.24, 9.85], 12.IX.1960, leg. Kuntze (1 J ZMHB). ITALY: Aniene, Lazio, [41.95, 12.82], 1200 m, 18.III.1964, leg. A. Vigna-Taglianti (1 J cBor); Barni, Lombardia, [45.91, 9.26], 11.II.2001, leg. Diotti (1 J 1 ♀ NHMD); Moliterno, Basilicata, [40.24, 15.86], 30.IX.2000, leg. F. Angelini (1 ♀ NHMD); Pignola, Basilicata, [40.57, 15.78], 700 m, 19.IV.1998, leg. F. Angelini (1 J NHMD); Querciola, Toscana, [43.84, 11.32], 13.IV.2005, leg. Bordoni (1 J cBor). LATVIA: Curonia Libau [Liepāja], [56.49, 21.01], 4.IV.1910 (1 J 1 ♀ ZMHB). MOROCCO: Cirque du Jaafar, [32.55, -4.91], 4.III.1964 (1 J NMW) [potentially mislabelled or introduced specimen clearly belonging to this species]. MYANMAR: Mountains, Tenasserim, Siam Border, [11.79, 99.62], II.-V.1913, leg. K.G. Gairdner (1 J BMNH) [potentially mislabelled or introduced specimen clearly belonging to this species]. PO-LAND: Dunajec River n. Sromowce Nizne, [49.39, 20.39], 18.VII.1977, leg. F. Hieke (3 JJ 4 ♀♀ ZMHB); Wyspowy, Krzozonowka River n. Poim, [49.69, 20.24] 300m, 26.VIII.1980, leg. Hieke (1 J ZMHB). PORTUGAL: Aඓඈඋൾඌ: Santa Maria, 2 km ENE Almagreira, Miradoro dos Picos, [36.97, -25.07], 440 m, 19.III.1957, leg. Brinck & Dahl (1 J MZLU); São Miguel, 500 m E of Lagoa do Pau Pique, [37.83, -25.74], pond, 7.III.1957, leg. Brinck & Dahl (1 J MZLU); São Miguel, Pico de Vara, 37.8119, -25.2105, 1050m, boggy slope 24.VII.2013, leg.V.Assing (4 ♀♀ ZMHB). Mൺൽൾංඋൺ: Levada da Serra do Faial, [32.68, -16.85], creek, 820m, 11.IX.2014, leg. A. Kleeberg (1 J cSch). ROMANIA: Klaussenburg [Cluj-Napoca], [46.76, 23.62], leg.Verhoeff (1 J ZMHB); Hermannstadt, Siebenbürgen [Sibiu], [45.79, 24.15], leg. Skalitzky (1 J NMW); Piatra, Virful Tiganului, Sighetu Marmației, Maramureș, [47.99, 23.53], 1300 m, pasture, in rotten logs of beech, 14.VI.2006, leg. A. Grabant, Z. György, O. Merkl & A. Podlussany (1 J NHMD). RUSSIA: KൺඋൺർHൺඒ- CH ൾඋKൾඌඌංൺ: Teberda,[43.44, 41.74], VI.1912, leg.Roubal (1 ♀ SNMC). Mඈඌർඈඐ Oൻඅൺඌඍ: Bykovo village, VNIIKR institute territory, [55.61, 38.05], 16.X.2013, leg.S.A. Kurbatov (3 JJ ZIN); Odintsovsky Distr., near Zhavoronki vill., [55.64, 37.10], 6-13.VIII.2017, leg. E. M. Veselova & A. B. Ryvkin (1 J ZIN). NංඓHൾGඈඋඈൽඌKൺඃൺ Oൻඅൺඌඍ: near Gorodets town, [56.64, 43.47], mixed forest, 17.VIII.1989, leg. A. Klimenko (1J ZIN). NඈඏGඈඋඈൽඌKൺඃൺ Oൻඅൺඌඍ: Torbino, Novgorod, [58.58, 32.87], 10.IV.1914, leg. A. Filiniev (1 J cBor). NඈඋඍH Oඌඌൾඍංൺ Rൾඉ.: Caucasus, Skalisty Khrebet, Fiagdon River, [42.87, 44.32], 1700-2200 m, VI.1976, leg. R. Rous (3 JJ 1 ♀ MCZ). SLOVAKIA: Belá River n. Pribylina, [49.09, 19.81], 600m, 18.VIII.1981, leg. Hieke (1 J 3 ♀♀ ZMHB). SPAIN: Bera de Bidasoa, Navarra, [43.27, -1.68], 1.-7.I.2001, leg.A.Anichtchenko (1J NHMD); Caldetas n. Barcelona, [41.57, 2.52], leg. G. Heine (1 J ZMHB); La Molina, Pyrenees, [42.33, 1.93], 11.-16. VII.1963 (1 J MZLU); O Rial, Serra do Mirador, [42.74, -7.83], 450m, 3.VI.2003, leg. J. P. Valcárcel (1 J cSch); Picos de Europa, Camping El Redondo [Fuente Dé], [43.14, 4.81], forest, 1100 m, 14.-17.VII.1996, leg. Wrase (1 J cSch); Plaiaundi, Irun, Guipuzcoa, [43.34, -1.79], V.2006, A. Anichtchenko (1 J 1 ♀ NHMD); Pyrenees, Toses, 42.3502, 1.9955, 1660m, fir-spruce forest with moss, 9.X.1997, leg. Zerche (1 J SDEI); Sierra Ancares, [42.81, -6.86], 20.X.1984, leg. J. C. Otero (6 JJ 3 ♀♀ ZMHB); Sierra Caurel, [42.59, -7.18], 20.VII.1985, leg. J. C. Otero (5 JJ 4 ♀♀ ZMHB); Sierra de Gredos, Hoyos de Collado, [40.35, -5.21], 1600m, 14.IV.1991,leg.Heinz (2♀♀ ZMHB); Sierra Guadarrama, [40.85, -3.95], 1.VI.1980, leg. C. Otero (2 JJ ZMHB); Sierra de la Demanda, Val de Zcaray [Ezcaray], [42.32, -3.02], forest, 1650m, 23.07.1996, leg. Zaballos & Wrase (1 J cSch). SWEDEN: Brönnestad, [56.08, 13.70] (1 MZLU); Omberg, [58.33, 14.64] (1 MZLU); Strömholm, [59.52, 16.23] (1 MZLU); Visseltofta, [56.42, 13.85] (1 MZLU); Öඅൺඇൽ: Vickleby, [56.57, 16.46] (1 MZLU). SWITZERLAND: Wolfenschiessen, [46.91, 8.39], 5.VIII.1963, moss (1 J ZMHB). TURKEY: Abant n. Bolu, [40.61, 31.27], 30.V.1964, leg. H.Korge (1 J ZMHB); Akkus n. Ünye, [40.79, 37.01], 28.V.1964,leg. H.Korge (1J 2♀♀ ZMHB); Ilgaz Dağları, [41.11, 33.90], 17.-21.VI.1960, leg. F. Schubert (1 ♀ NMW); n. Ismetpasa & Cerkes, [40.75, 32.82], 05.IV.1979, leg. Heinz (1 J NMEG); Kulakkaya n. Giresun, [40.69, 38.33], 1450m, 27.VII.1963, leg. H. Korge (2 ♀♀ ZMHB). UKRAINE: Tausban Bazar, [44.97, 34.59], 20.VI.1907, leg. B. Grigoriev (1 J ZIN). UNITED KINGDOM: EඇGඅൺඇൽ: Barnstaple, Devon, [51.07, -4.06], 12.-31.V.1954, leg. Lindroth (1J MZLU); Devon, Bidefort [51.01, -4.21], leg. H. Korge (2 ♀♀ ZMHB); Kent, Knockholt, 51.3124, 0.1191, 2.IV.2011, leg. J.J. Shaw (1 J cJen); Shrops, Edgmond Harper Adams Uni. Campus, [52.77, -2.42], 6.III.2015, leg. J.J. Shaw (1 J cJen); Sർඈඍඅൺඇൽ: Loch Hope, [58.44, -4.62], 19.VII.1987, leg. G. Gillerfors (1 J MZLU). USA: Maine: Booth Bay, [43.85, -69.62], 15.VIII.1982, leg. Wewalka (1 ♀ NMW). UZBEKISTAN: Buchara, [39.77, 64.42], leg. Bang Haas (1 J FMNH).

Redescription. Measurements JJ (n = 6): HW = 1.89– 1.96 (1.93); HL = 1.53–1.71 (1.61); HL/HW 0.81–0.88 (0.84); PW = 2.36–2.56 (2.49); PL = 2.16–2.36 (2.29); PL/PW 0.87–0.94 (0.92); EW = 2.51–2.67 (2.61); EL = 2.36–2.49 (2.44); EL/EW 0.92–0.96 (0.93); EL/PL 1.04– 1.10 (1.06); PW/HW 1.31–1.60 (1.49); forebody length 6.04–6.56 (6.34). J ♀ (n = 4): HW = 1.78–2.00 (1.89); HL = 1.51–1.64 (1.58); HL/HW 0.82–0.85 (0.84); PW = 2.22–2.56 (2.41); PL = 2.07–2.40 (2.18); PL/PW 0.86–0.94 (0.91); EW = 2.44–2.62 (2.52); EL = 2.20–2.49 (2.33); EL/EW 0.90–0.95 (0.92); EL/PL 1.03–1.15 (1.07); PW/ HW 1.47–1.55 (1.52); forebody length 5.78–6.53 (6.10).

Very large species; body black ( Fig. 7B ).

Head black, slightly transverse; eyes large (EyL/TL = 3.00–4.10 (3.55)) not protruding; microsculpture of transverse waves, four punctures between anterior frontal punctures ( Fig. 6B ); antennae and palpi pale reddish, antennae long, all antennomeres clearly elongate.

Thorax: pronotum black, slightly wider than long, clearly wider than head, microsculpture of transverse waves, three punctures in dorsal row and two in sublateral row with posteriormost puncture reaching level of middle puncture of dorsal row; scutellum smooth and glabrous; elytra black, uniformly pubescent, wider than long, of equal length as pronotum; legs dark brown with inner face of tibia darkened and tarsi paler.

Abdomen black, tergites uniformly punctured, slightly iridescent.

Male. Aedeagus ( Fig. 11C ): paramere broad, rather parallel-sided, reaching apex of median lobe, with sensory peg setae forming two single rows following parameral lateral margins for 2/3 of their length; median lobe with apical part dorsoventrally flattened to form a plate-like structure; internal sac with two pairs of sclerites.

Differential diagnosis. Within the fuliginosus- group Q. curtipennis is very close to Q. fuliginosus . Both species are very frequently encountered, often co-occurring, nearly non-distinguishable from each other based on external morphological characters alone. Our study of hundreds of specimens across their entire distribution ranges revealed that Q. curtipennis always have pale antennomeres 1–3, whereas Q. fuliginosus have them at least slightly darkened. A negligible fraction (ca. 2 %) of Q. fuliginosus with pale basal antennomeres are mostly teneral. Antennae are generally more elongate in Q. curtipennis , as opposed to being stouter in Q. fuliginosus ( Fig. 7 ). Also Q. curtipennis has an almost smoothly rounded head with the eyes protruding while Q. fuliginosus has eyes clearly protruding and thus making its head seem more quadrate ( Fig. 6 ). Only some rare specimens, maybe hybrids, faded and very old museum specimens, or actual rare intraspecific variants, do not fit these trends and their identity can be ascertained only through examination of the aedeagi. In Q. curtipennis , contrary to Q. fuliginosus , the paramere lacks clear medial attenuation, the apical part of median lobe dorso-ventrally flattened, and internal sac with 2 pairs of clear sclerites.

The examined material for both species clearly shows that Q. curtipennis is much more common in the south and west and becomes increasingly rare towards the north and east of its range, while Q. fuliginosus shows an opposite trend. Quedius curtipennis seems to be more common in warmer and drier areas with more sandy soils, where Q. fuliginosus seems to prefer wetter, colder and more humus rich soils.

Quedius curtipennis is also very similar to Q. afrofuliginosus , from which it is easily recognized by having no additional punctures between anterior and posterior frontal punctures. Besides, distributions of Q. curtipennis and Q. afrofuliginosus , as far as known, are allopatric.

From Q. levicollis , Q. curtipennis is easily recognized by unpunctured and glabrous scutellum.

Lectotype designation. BൾඋඇHൺඎൾඋ (1908) described Q. curtipennis as a variety of Q. fuliginosus without giving any exact data on the type material. He had stressed that it is very common at Faroe Islands and mentioned its occurrence in Central Europe as well as in ‘Romania’ and even ‘Buchara’. Here (see above), we have listed specimens from Faroe Islands and ‘Buchara’ at the FMNH which qualify as syntypes of Q. curtipennis and which we recorded long ago but were unable to re-examine in detail recently. There is no doubt about the identity of the syntypes from Faroe Islands. However, a syntype from ‘Buchara’ must be carefully checked. Given, additionally, some ambiguity about the geographic origin of that Middle Asian syntype (see Distribution), a syntype male from Faroe Islands is here designated as a lectotype for Q. curtipennis .

Synonymic notes. Quedius parallelus was described by HൺඍർH (1957) in his work on beetles of the Pacific Northwest of North America without reference to European species. Based on the description of Q. parallelus , KඈඋGൾ (1962b) suspected it to be identical to the European Q. curtipennis . This was confirmed by Sආൾඍൺඇൺ (1971a) who studied the type material of Q. parallelus and found them conspecific with 2 male syntypes of Q. curtipennis from Faroe Islands at the FMNH. It is now firmly known that Quedius curtipennis occurs in the North American West (Sආൾඍൺඇൺ 1971a) and East Coasts (Sආൾඍൺඇൺ 1990, Bඋඎඇĸൾ & MൺඋඌHൺඅඅ 2011).

Quedius gracilis Stephens, 1832 was described from London, UK. Later, SඍൾඉHൾඇඌ (1837) had it as a variety of Q. tristis (now Q. levicollis ). In more recent works it was cited as a synonym of Q. fuliginosus . Based on examination of its two syntypes by Roger Booth (BMNH, pers. comm.) on our request, it was established that these specimens are conspecific with Q. curtipennis . As the name is older than Q. curtipennis we here establish the prevailing use of Q. curtipennis as opposed to the rule of priority (ICZN Article 23.9). Quedius curtipennis has been used at least 26 times in the last 50 years by more than ten authors (Bඈඋൽඈඇං 1974b, 1976b; Pඈඉൾ 1977; Cඈංൿൿൺංඍ 1978; Oඎඍൾඋൾඅඈ 1978; Bඎඋൺĸඈඐඌĸං et al. 1980; TඬඍH 1984; DඋඎGආൺඇൽ 1987; Nඈඐඈඌൺൽ 1990; BඈඋGൾඌ 1990; Cංർൾඋඈඇං & Zൺඇൾඍඍං 1995; Oඐൾඇ 1997; Oඎඍൾඋൾඅඈ et al. 1998; AඌඌංඇG 2001; UHඅංG et al. 2006; Tඋඈඇඊඎൾඍ 2006; Mൺඃĸൺ & Sආൾඍൺඇൺ 2007; ÖඓGൾඇ 2011; Bඋඎඇĸൾ & MൺඋඌHൺඅඅ 2011; Wൾൻඌඍൾඋ et al. 2012; MൺGඎඋൺ et al. 2013; TඬඍHආඣඋඣඌඓ et al. 2014; Sൾආൾඇඈඏ et al. 2015; Sൾආංඈඇൾඇĸඈඏ et al. 2015; Sൺඅඇංඍඌĸൺ & Sඈඅඈൽඈඏඇංĸඈඏ 2018a, 2019). On the other hand, Q. gracilis has only been used once after its description as a valid species which was more than 150 years ago (Cඎඋඍංඌ 1837). This deems the name Quedius gracilis Stephens, 1832 a nomen oblitum in relation to Quedius curtipennis Bernhauer 1908 (nomen protectum). Quedius gracilis Stephens, 1832 syn. rev. is thus moved from synonymy with Quedius fuliginosus ( Gravenhorst, 1802) to synonymy with Quedius curtipennis Bernhauer 1908 .

Bionomics. Based on published data as well as our observations, Q. curtipenni s is a polytopic, somewhat thermophilous species found in a large range of habitats and various ground-based microhabitats with a tendency to occur in open woodlands. For example, in Central and Eastern Europe it was found under stones in dry pastures in Poland (Dඏඈෞගĸ 1965), in xerothermous grassland in Germany (AඌඌංඇG 2001), and in closed oak forest with extensive ground vegetation and shrub cover in Hungary (TඬඍHආඣඋඣඌඓ et. al 2014). We have collected this species in a mixed forest on the sandy dry island of Laesø in Denmark. There are also records from mole ( Talpa europaea ) nests built from grasses and reeds on wet meadow in Poland (Nඈඐඈඌൺൽ 1990). In south-western Europe it was recorded from under rock in pine forest at an altitude of 1550 m in Central Spain (Oඎඍൾඋൾඅඈ 1978) and from a cave in Northern Spain (Oඎඍൾඋൾඅඈ et al. 1998). On the Azores (Sආൾඍൺඇൺ 1970) it was found under rocks at the shore of a freshwater lake at an elevation of 400– 500 m. Presumably, the nest or cave-based subterranean records are accidental occurrences, as there is no evidence of the species preferring these habitats. The species is found within the wide range of elevations from sea level in the northern part of its range up to about 2.000 m the southern part.

As an introduced species in North America, it was found especially around human settlements in various debris, under rocks, and in greenhouses. Some specimens were also collected by sifting barnyard litter on a pasture in Oregon (Sආൾඍൺඇൺ 1978a) and in natural habitats away from settlements (in moss, in leaf litter, etc.).

Pupae of the species are described in Oඎඍൾඋൾඅඈ (1978). Adult females from Wytham, UK were observed to contain eggs between October and January (Fඋൺඇĸ 1969).

Distribution. Quedius curtipennis is distributed from the Atlantic coast of Europe to Asia Minor, east to the Ural Mountains and Middle Asia ( Fig. 18 ). It is also present on the islands of Madeira, the Azores, and the Faroe Islands. One specimen, which we studied was found in the central Morrocan mountains, and one old specimen from the type series is from ‘Buchara’, Uzbekistan. These are the only specimens we know from North Africa and Middle Asia, respectively. The species’ occurrence in these areas needs confirmation to rule out possible mislabelling. Most previous records of Q. curtipennis (and Q. fuliginosus ) from North Africa could be assigned to Q. afrofuliginosus , which is externally very similar. As Q. curtipennis does not seem to occur in southern Spain, Corse and Sardinia, it is very likely absent from North Africa too.

Quedius curtipennis is also known as introduced to at least four regions of North America ( Fig. 12 ). The presence of Q. curtipennis in the Nearctic was first recognized by Sආൾඍൺඇൺ (1971a) when he reviewed specimens of Q. parallelus , a newly described species from the Nearctic. Quedius parallelus was found to be identical to the Palearctic Q. curtipennis , something already hinted by KඈඋGൾ (1962b). One specimen from the type series of Q. parallelus was collected in 1934 near the city of Seattle in Washington, which is currently the oldest known record of the species from North America. In 1971, the species was already found in a relatively large area along the Pacific coast of Oregon, Washington, and British Columbia (Sආൾ-ඍൺඇൺ 1971a), suggesting that it may have been established in North America even prior to 1934. Currently, Q. curtipennis in western North America is still known only from these states and provinces, where it is found as far East as Cranbrook, British Columbia in the Canadian Rockies (from web www.boldsystems.org) (teal in Fig. 12 ). One to three additional introductions of Q. curtipenni s were later reported from the East Coast, in New Hampshire, USA, as well as Ontario and Nova Scotia, Canada. Specimens from near Guelph in Ontario were collected in 1976 (Bඋඎඇĸൾ & MൺඋඌHൺඅඅ 2011), from New Hampshire in 1983 (Sආൾඍൺඇൺ 1990) and from Nova Scotia in 1997 (Mൺඃĸൺ & Kඅංආൺ-ඌඓൾඐඌĸං 2008).

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. 3. Phylogeny of Quedius s. str. based on Bayesian Inference (BI) and Maximum Likelihood (ML) analyses of COI barcodes (part 1). 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.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).

Fig. 7. Habitus of Quedius s. str. A – Q. afrofuliginosus Gusarov, 1991; B – Q. curtipennis Bernhauer, 1908; C – Q. fuliginosus (Gravenhorst, 1802); D – Q. levicollis Brullé, 1832; E – Q. unicolor Kiesenwetter, 1847; F – Q. subunicolor Korge, 1961.

Fig. 11. Aedeagus ofQuedius s. str. A – Q. afrofuliginosus Gusarov, 1991; B – Q. levicollis Brullé, 1832; C – Q. curtipennis Bernhauer, 1908; D – Q. fuliginosus (Gravenhorst, 1802). 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. 12. Distribution of Eurasian species introduced to North America: Quedius curtipennis Bernhauer, 1908 (teal), Q. fuliginosus (Gravenhorst, 1802) (green), and Q. molochinus (Gravenhorst, 1806) (orange).

Fig. 18. Distribution of Quedius afrofuliginosus Gusarov, 1991 (top), Q. curtipennis Bernhauer, 1908 (middle) and Q. fuliginosus (Gravenhorst, 1802) (bottom).