Quedius fuliginosus ( Gravenhorst, 1802 )

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 : 248-252

Quedius fuliginosus ( Gravenhorst, 1802)

( Figs 1 , 2A,E , 3 , 6C , 7C , 11D , 12 , 18 )

Staphylinus fuliginosus Gravenhorst, 1802: 34 [Type locality:Brunsviga]. Note: See lectotype designation by Gඎඌൺඋඈඏ (1993)

Quedius granulipennis Motschulsky, 1858: 656 [Type locality: Autriche]. Note: See lectotype designation by Gඎඌൺඋඈඏ (1993)

Quedius impunctifrons Delahon, 1915: 395 . [Type locality: Mark Brandenburg]

Quedius jelineki Krása, 1904: 81 [Type locality: Vrané n. Vlt, Bohemia]

Quedius latus Hochhuth, 1851: 30 [Type locality: Kaukasien, Alagez- Gebirge], syn. nov.

Quedius molochinicolor Roubal, 1931: 1 [Type locality:RCS.:Harmanec]

Quedius picicornis Stephens, 1832: 215 [Type locality: London]

Quedius subfuliginosus Britten, 1944: 290 [Type locality: England: Cotterill Clough, Cheshire]

Quedius viduus Sawada, 1965: 17 [Type locality: Japan, Pref. Nagano, Mt. Jônen], syn. nov.

References. MൺඋඌHൺආ (1802): 504 (characters); Lൺඍඋൾංඅඅൾ (1804): 319 (characters); GඒඅඅൾඇHൺඅ (1810):301 (as tristis ; characters);MൺඇඇൾඋHൾංආ (1830): 25 (distribution); (1831): 439 (distribution); Nඈඋൽආൺඇඇ (1837): 78 (distribution); EඋංർHඌඈඇ (1839a): 490 (distribution); (1840): 537 (characters); Hൾൾඋ (1839): 276 (characters); Dඎൿඈඎඋ (1843): 33 (characters); Kඈඅൾඇൺඍං (1846): 22 (distribution); GඋൺඏൾඇHඈඋඌඍ (1847): 232 (characters); HඈർHHඎඍH (1849): 151 (characters); (1862): 46 (distribution); RൾൽඍൾඇൻൺർHൾඋ (1849): 710, (1857): 203, (1874): 200 (characters); Kංൾඌൾඇඐൾඍඍൾඋ (1851): 419 (distribution); Hൺඋൽඒ (1851): 33 (biology); Kඳඌඍൾඋ (1853): 62 (characters); Fൺංඋආൺංඋൾ & Lൺൻඈඎඅൻජඇൾ (1856): 539 (characters); Kඋൺൺඍඓ (1857): 503 (characters); (1858): 58 (distribution); Pൾඒඋඈඇ (1858): 428 (distribution); Fൺඎඏൾඅ (1865):293 (biology); (1874): 289 (characters); Sൾංൽඅංඍඓ (1875): 267 (characters); Mඎඅඌൺඇඍ & Rൾඒ (1876): 686 (characters); SൺHඅൻൾඋG (1876): 24 (distribution); Fඈඐඅൾඋ (1888): 237 (characters); GൺඇGඅൻൺඎൾඋ (1895): 403 (characters); Pඈඉඉංඎඌ (1905): 9 (distribution); Pඈඋඍൺ (1907): 128 (characters); Rൾංඍඍൾඋ (1909): 111 (characters); JඈHൺඇඌൾඇ (1914): 363 (characters); EංർHൾඅൻൺඎආ (1914): 339, (1915): 91 (morphology); Fංඈඋං (1915): 16 (distribution); Mඎඇඌඍൾඋ (1923): 195 (distribution); Gඋංൽൾඅඅං (1924): 79 (characters); Pඈඋඍൾඏංඇ (1929): 341 (characters); Wඳs ඍHඈൿൿ (1938): fig. 21 (morphology); Pൺඎඅංൺඇ (1941): 263 (larval characters); Hංඇඍඈඇ (1945): 70 (characters); FൺGൾඅ (1948): 196 (characters); Jൾൺඇඇൾඅ & JൺඋඋංGൾ (1949): 378 (biology); Hൺඇඌൾඇ (1952): 140 (characters); Sආൾඍൺඇൺ (1958): 361 (characters and biology); (1962): 133 (characters and distribution); (1964): 79 (distribution); (1976): 22 (distribution); (1978c): 85 (distribution); (1993): 50 (distribution); Pൺඅආ (1963): 139 (characters); LඈHඌൾ (1964):211 (characters); Hඈඋංඈඇ (1965): 272 (distribution); Pඈඍඈඍඌĸൺංൺ (1967): 84, 87 (larval characters); Kൺඌඎඅൾ (1968): 53 (biology); (1970): 63 (larval characters); KඈඋGൾ (1968): 52 (characters); Sඓඎඃൾർĸං (1968): 735 (distribution); Fඋൺඇĸ (1969): 266 (biology); (1982): 46 (list of parasites); Bඈඋൽඈඇං (1974): 12, (1976a): 89 (characters); (1976b): 238 (distribution); Oඌൾඅඅൺ & Zൺඇൾඍඍං (1975): 120 (biology); Pඈඉൾ (1977): 31 (distribution); Cඈංൿൿൺංඍ (1978): 190 (distribution); Tඈඉඉ (1979): 19 (development); Bඎඋൺĸඈඐඌĸං et al. (1980): 134 (distribution); TඬඍH (1984): 117 (characters); Nඈඐඈඌൺൽ (1990):143 (biology);Aඅඅൾඇ (1990): 4 (biology); WHංඍൾHൾൺൽ (1991): 6 (biology); Gඎඌൺඋඈඏ (1991):9 (identity); (1993): 72 (synonymic notes, lectotype designation); WൾඅർH (1993): 229 (morphology); HඈൽGൾ & Jඈඇൾඌ (1995): 42 (characters); Cංർൾඋඈඇං & Zൺඇൾඍඍං (1995):32 (distribution); Sඍൺඇංൾർ (1996):117 (pupal characters); (2010): 22 (distribution); Oඐൾඇ (1997): 149 (biology); Sඍൺඇංൾർ (1999): 52 (characters of pupa); Dൾඋඎඇĸඈඏ (2005): 279 (distribution); UHඅංG et al. (2006): 54 (distribution); Tඋඈඇඊඎൾඍ (2006): 102 (distribution); Mൺඃĸൺ & Sආൾඍൺඇൺ (2007): 428 (distribution); Mൺඓඎඋ et al. (2007): 29 (distribution);Sඍൺඇ (2009):242 (distribution);Sൾආൾඇඈඏ et al.(2015): 127 (distribution); Sൾආංඈඇൾඇĸඈඏ et al. (2015): 331 (distribution); Sൺඅඇංඍඌĸൺ & Sඈඅඈൽඈඏඇංĸඈඏ (2018a): 125, (2019): 48 (distribution); PඎർHĸඈඏ et al. (2020): 43 (distribution).

Type material examined. Quedius fuliginosus : Lൾർඍඈඍඒඉൾ J (ZMHB), labelled: “coll. Hellwig / Paratypus fuliginosus Grav. ”.

Quedius granulipennis : Lൾർඍඈඍඒඉൾ J (ZMMU), labelled: “ Austria / Quedius granulipennis Motsch. Austria ”.

Quedius latus : Nൾඈඍඒඉൾ J (ZMHB), designated here, labelled: “AR-MENIA (AR-16-19) 35 km NW Sisian, 39°40’59”N 45°46’50”E, 2070 m, stream valley, litter beneath bushes near stream sifited 3.VII.2016, leg. M.Schülke / Quedius (Quedius) fuliginosus (Gravenhrost) det. M. Schülke 2019 / Museum für Naturkunde Berlin Sammlung M. Schülke / Neotype Quedius latus Hochhuth, 1851 A.K. Hansen & A. Solodovnikov des. 2021”.

Quedius molochinicolor : Sඒඇඍඒඉൾ ♀ (SNMC), labelled: “ Slovensko Harmanec Roubal 26.IX.1930 / f. molochinicolor mi [my] type / [red label] / fuliginosus f. molochinicolor cotypus”.

Additional material examined. ARMENIA: N Yerevan, NW Hrazdan, 40.6350 44.4602, grassy W slope with scattered Salix , litter and roots of grass sifted, 2500m, 26.VI.2016, leg. M. Schülke (1 ♀ cSch); same locality, 40.6944, 44.4878, stream valley, mixed decidious forest, litter and grass roots sifted, 2110m, 28.VI.2016, leg. M. Schülke (1 ♀ cSch). AUS-TRIA: Arbesbach; [48.49, 14.95], 1.-13.VII.1951, leg. Schubert (1 ♀ NMW); Brunnlust, Moosbrunn, Schwechat, [48.01, 16.45], 12.VI.1997, leg.Schillhammer (1J NMW);Faak, Kärnten,[46.56, 13.91], 5.VIII.1968, leg. Wewalka (1 ♀ NMW); Immerkrems, [46.96, 13.72], subalpine, 1400 - 2000 m, VI.1954, leg. Lindroth (1 ♀ MZLU); Plesch-Kogel n. Rein, [47.13, 15.28], 590m, sifting forest litter, 20.VI.1995, leg. Zerche & Behne (1 J SDEI); Rekawinkel, [48.17, 16.02], leg. Briet (1 J NMW). BULGARIA: Sofia, Tschernia [Cherni Vrah], [42.56, 23.27], 600m, 22.VI.1988,leg.Behne (1J SDEI); Sredna Gora,S Koprivschtiza, 42.8777, 25.0544, moss under Picea, 1000m , 29.VI.1997, leg. Zerche & Behne (1 ♀ SDEI); Stara Plania, 5km S Ribaritza, 42.75, 24.38, 750m, 7.VI.1997, Beech forest, sifting, leg. Zerche & Behne (1 J SDEI); SW-Pirin, Kurort Popina Laka, 16 km NO Sandanski, [41.56, 23.27], ca. 1350m, Pinus and Picea , under snow remains, sifted, 8.IV.2005 leg. Zerche & Behne (1 J SDEI); W Rhodopen, S Velingrad, Tschernowroh, [41.68, 24.05], 1600m 10.VI.1987, leg. Zerche & Behne (1 ♀ SDEI). CZECH REPUBLIC: Podyjí Nat. Park, forest at river Dyje valley ca. 2.5 km NW of Havraníky, 48.8300, 15.9804, h 240m, sifted flood debris, 8.VI.2016, leg.A. Solodovnikov, J. Jenkins Shaw,A. Hansen (1 ♀ NHMD). DENMARK: Amager, Kalvebod Faelled, Store Høj Søen, 55.6159, 12.5606, sifting leaf litter, debris near lake, 17.III.2019, leg.A. K. Hansen (1 J 1 ♀ NHMD); Lisjerg Forest, 8 km N of Aarhus, 56.2335, 10.1688, h 85m, edges of forest lakes, hand collected in leaf litter and moss, 17.IV.2019, leg.A. K.Hansen (4JJ 4 ♀♀ NHMD); Nors Sø, S of Nors Sø, 57.0197, 8.6197, pitfall trap, dry pine forest, 16-18.VI.2014, leg.Solodovnikov, Hansen, Kraemer, & Justesen (1 J NHMD). GEORGIA: N of Sviri, [41.73, 42.99], pitfall traps, 14.IV.-11.VI.2018, leg. D. Formynikh (1 J cSch); Tusheti, n. Omalo, Abanos Pass, [42.27, 45.51], river valley, 1700m, 14.-17.VII.2016, leg. Heinz (1 J NMEG). GERMANY: Bayern, Eschenlohe, [47.58, 11.18], 3.VIII.1937,leg.Ihssen (1J 1♀ SDEI);Berlin-Blankefelde,[52.62,13.39], 8.V.1994, leg. D. Wrase (1 J SDEI); Biesenthal, [52.76, 13.61], 500m, field sandish-clayish ruderal, 1.IV.2002, leg. D. Wrase (1 ♀ SDEI); Brandenburg Kiesfläche 1km S Oderberg, [52.84, 14.04], 13.X.1990, leg. F. Hieke (1 J SDEI); Brandenburg, Oranienburg, Briesetal, [52.71, 13.31], 12.IV.1992, leg.J. Ziegler (1J SDEI); Dars, Zingst, Freesenbruch, [54.43, 12.68], 20.VII.1987, leg. D. Wrase (1♀ SDEI); Erzgebirge,Prinzgehöhle, [50.63, 12.67], 29.IX.1921, leg. Uhmann (1 ♀ SDEI); Erzgebirge, W Teil, SW Kammweg & Morgenröthe, [50.43, 12.51], 4.V.1990 leg. O. Sorge (1 J SDEI); Finkenkrug n. Berlin, [52.56, 13.03], 12.IV.1898, leg. Dahl (1 J SDEI); Kehlheim Dürnbacher Forst, [48.74, 11.75], (1 ♀ SDEI); Liepnitzsee, n.Wandlitz,[52.74, 13.51], 01.X.1989, leg. M. Uhlig (1J SDEI); Marburg i Hess, [50.81, 8.77], 16.IV.1904, leg. Strand (1 J 1 ♀ SDEI); n. Märkische Buchholz, Königs Wusterhausen, [52.11, 13.76], 12.VI.1980, leg. H.Wendt (1 ♀ SDEI); Martinfeld, Krs. Heiligenstadt, [51.28, 10.18], 21.-29.VI.1990, leg. M. Uhlig (1 J SDEI); München-Grünwald, [48.04, 11.53], 3.VII.1931, leg. Ihssen (1 ♀ SDEI); Nauen, [52.61, 12.87], 19.III.1910, leg.M.Ude (1J SDEI);Sächsischen Schweiz, Rathen,[50.95, 14.08], 1.V.1977, leg.K. H.Mohr (1♀ SDEI); Schwäbisch Gmünd,Röben, [48.79, 9.81], 4.V.1953 (1 J SDEI); Seilershof, Gransee, [53.06, 13.18], 22.V.1988, leg. M. Schülke (1 ♀ SDEI); Suekow, [53.07, 11.79], XI.1974, leg. M. Moritz (3 JJ SDEI); Wandlitz, Liepnitzsee, [52.74, 13.51], 6.I.1981, leg. M. Uhlig (1 J 2 ♀♀ SDEI); Wildeschütz n. Deuben Silbersee Schilfgürtel, [51.12, 12.07], 11.X.1987, leg. M. Uhlig (1 ♀ SDEI). ITALY: Alpie Orobie,Bergamo Castione &Passo delia Presolana, 45.9202, 10.1001, 1170m, 18.-21.VI.2006, leg.P. Schnitter (1 J NMEG); Basilicata, Pognola Ris. L. Pignola, [40.58, 15.74], 700 m, 23.XII.1996, leg. F. Angelini (1 cBor); Trentino, Lago di Tenno,N Riva, [45.93, 10.81], 560m, 2.VII.1994, leg. L. Zerche (1 ♀ SDEI). KAZAKHSTAN: Altai, Bukhtarma riv., Chingistai, [49.19, 85.88], 30.VIII.2010, leg. V.A. Kastcheev (1 J ZIN);Akshatau Mt., NW Ayaguz, Semipalat, [48.25, 79.64], forest leaf litter, 17.VII.1962, leg. L. V.Arnoldi (1 J ZIN); Ivanovsky Mt. Ridge, 32 km S Leninogorsk, [50.04, 83.51], 1300 m, 14.VIII.1986, leg. I. I. Kabak (1 J ZIN). RUSSIA: Aඅඍൺං Kඋൺං: Talmenskij Distr., E of vill. Litvinovka 53.6918, 83.7309, 200m, mixed forest, sifted leaf litter, 24.VI.2019, leg. A. Solodovnikov, A. K. Hansen & A. Tokareva (1 NHMD); Chemalskij Distr., SE of vill. Edigan 51.0546, 86.3791, 1030m, larch-fur forest, sifted leaf litter, 27.VI.2019, leg.A. Solodovnikov & A. K. Hansen (1 NHMD); Chemalskij Distr., Verkhnij Berulu river, 51.3035, 86.1221, 790m, pine-birch forest, sifted leaf litter, 28.VI.2019, leg. A. Solodovnikov, A. K. Hansen & M. J. Justesen (1 NHMD). BൺඌHKඈඋඈඌඍൺඇ Rൾඉ.: Buzdyakskij Distr., nr vill. Arslanovo 54.6485, 54.3789, 229m, at creek bank, 11. VI.2019, leg.A. Solodovnikov & A. Tokareva (1 NHMD); Mechetlinskij Distr., Oka river nr vill. Bolshaya Oka, 56.1085, 58.1977, 224m, debris and ground at river bank, 12.VI.2019, leg.A. Solodovnikov & A. Tokareva (1 NHMD). IඋKඎඍඌK Oൻඅൺඌඍ: Baikal, Chamar-Daban, Solsan Valley, [51.36, 104.15], 17.-21.VI.1978, leg. Schilenkov (1 J cSch); Baikal, Baikalsk, Solsan River near mouth, [51.47, 104.37], 20.VII.1989, leg. Hieke (1 J 1 ♀ SDEI); Khamar-Daban Mts., valley of Snezhnaya River, [51.06, 103.99], 5.-8.VI.2007. leg.A. Shavrin (3 JJ 6 ♀♀ cSha); Podvoloshino, Valley of Nizhnyaya Tungska River, [58.25, 108.42], 4.-9. VIII.2008, leg.Shavrin & Enustschenko 18 km N Ust-Kut,Valley of Lena River, [56.95, 106.14], 26.-28.VII.2008, leg. Shavrin & Enustschenko (1 ♀ ZIN) Valley of Bodchakta River, [57.11, 106.61], 10.-11.VIII.2008, leg. Shavrin & Enustschenko (1 J ZIN). KൺඅඎGൺ Pඋඈඏ.: 4-5 km SE from Chernaya Grayz, [54.96, 36.86], V.1988, leg. I. Ushakovl (1 J cRyv); Kozelsk Reg., Berezich glass factory, Novaya Derevnya village, NR “Ugra”,[56.27, 40.25], broad leaved forest,leaf litter near stream (stagnant channel), 13.VII.2017, leg. M. Salnitska (1 ♀ ZIN). KൾආൾඋඈඏඌKൺඃൺ Oൻඅൺඌඍ: Tashtagolsky Distr.,Shorsky Nat.Park, 5 km SSE Tajmet [Verkhnij Tajmet] village, [52.48, 88.27], cedar forest, 8-18.VI. 2012, leg. L. A. Trilikauskas (1 J 1♀ ZIN). KඋൺඌඇඈඒൺඋඌKංඃ Kඋൺං: nr.Divnogorsk town, left tributary of Enissey river, [55.95, 92.38] cedar-birch forest, in leaf litter, 1.VI.1988, leg.A. B. Ryvkin (2 JJ ZIN); Enissey distr., environs of vill. Ust-Pit, [58.98, 91.76]‚ 02-10.VII.1995, leg. A. Rybalov (2 JJ 1 ♀ ZIN); Nature Preserve ‘Stolby’ [Kranoyarskie Stolby], river Mana near Berly hut, [54.98, 94.00], stream edge, in moss and litter, 20.VI.1990, leg. A. B. Ryvkin (1 J ZIN); Niznaya Lebedyanka river, [61.96, 89.45], 22.VI.1992, leg.V.S. (2JJ cRyv). LൾඇංඇGඋൺൽ Oൻඅൺඌඍ: Kirovsky Distr., Turishkino vill., Mga River, 59.684306, 31.2202, flood plain, meadow in forest, leaf litter, 02.VI.2018, leg. M. Salnitska, K. Fadeev (1 J ZIN); Leningrad [Sankt Petersborg], in litter, 29.VII.1976, leg. Schilow (3 JJ SDEI). Mඈඌർඈඐ Oൻඅൺඌඍ: Odintsovsky Distr., Zvenigorod Biological Station of MSU, [55.70, 36.72], soil sample, Betula forest, 7.VIII.1981, leg. K. G. Mikhailov (1 J cRyv). NඈඏGඈඋඈൽ Oൻඅൺඌඍ: Staraja Russa, [57.99, 31.35], leg. Kessler (1 J NMW). Pൾඋආ Kඋൺං: Perm town, ravine meadow on the right bank of Kama river, Motovilikha, [58.04, 56.30], 1989, leg. V. O. Kozminykh (1 J ZIN). Rൾඉ. ඈൿ AൽඒGൾൺ: Caucasus, Adygeja, Guzeripl, Belaja River, [43.99, 40.13], 2000m, 23.VI.1999, leg. Putchkov (1 ♀ cSch). Rൾඉ. ඈൿ N. Oඌඌൾඍංൺ- Aඅൺඇංൺ: N Ossetia Nat. Res., mt. Kariuchoch, Kora pass. (Alansky), 42.8301, 44.2123, ~ 2200m, leaf litter Betula sp. , 24.V.2017, leg. M. Salnitska (1 J ZIN). SඏൾඋൽඅඈඏඌK Oൻඅൺඌඍ: Visimskij Nature Res., 57.3733, 59.7734, 570m, spruce dominated forest, pitfalls, 18.VI.2019, leg. A. Solodovnikov, A. K. Hansen & A. Tokareva (1 NHMD); W of vill. Elan’, 57.6406, 63.6220, 100m, mixed forest, sifted leaf litter, 20.VI.2019, leg.A. Solodovnikov,A. K. Hansen & A. Tokareva (1 NHMD). Tඒඎආൾඇ Oൻඅൺඌඍ: Uvatsky Distr., 8-14 km S from vill. Gornoslinkino, near Tobolsk station of IEE RAS,[58.68, 68.79], 11.VI.2004, leg. A. B. Ryvkin (2 JJ 1 ♀ cRyv); Tyumenskij Distr., Tura river nr vill. Konyashina, 57.2925, 65.0281, 80m, flood forest, sifted leaf litter, 21.VI.2019, leg. A. Solodovnikov, A. K. Hansen & A. Tokareva (1 NHMD); nr Zavodoukovsk, 56.5329, 66.5689, 116m, pine-birch grove, sifted leaf litter, 21.VI.2019, leg. A. Solodovnikov, A. K. Hansen & A. Tokareva (1 NHMD). SLOVAKIA: Belá River n. Vavrisovo n. Lipt. Hrádek,[49.06, 19.77], 18.VII.1980, leg. Hieke (12JJ 1♀ SDEI); Gombasek n. Rožňava, [48.57, 20.45], 6.VIII.1981, leg. Hieke (1 ♀ SDEI); Harmanec, [48.79, 19.07], 26.IX.1930, leg. Roubal (1 ♀ SNMC); Pieninsky National Park, [49.38, 20.51], 18.VII.1993, leg. Zerche (1 J SDEI); Rudohorie, Hnilec-Tal n. Švedlár, [48.83, 20.51], 8.VII.1981 leg. Hieke (1 ♀ SDEI). SLOVENIA: Triglav Nat. Park, river Soča valley, ca. 3 km SW Trenta; Obcina Bovec, 46.3667, 13.7255, sifting flood debris and under stones, 03.VI.2016, leg. A. Solodovnikov, A. Hansen. & M. Salnitska (1 NHMD). SWEDEN: Månsatorp, 5km SW of Vittsjö, Skåne; 56.3105, 13.5955, hand collected in/under moss, 21.I.2018, leg. A. K. Hansen (1 J 2 ♀♀ NHMD); Ödeshög V, Ekopark Omberg, Storpissan nature reserve, 58.3344, 14.6508 sifting leaf litter on stream edge; 30.V.2017, leg.D. Żyła, K. Koszela, A. Solodovnikov (1 J NHMD); Vegesjö, Skåne, 56.3031, 13.6473, banks of bog in wet sphagnum, 100 m, 22.IV.2018, leg. A. K. Hansen (1 ♀ NHMD). TURKEY: Harçbeli Pass, Ordu, [40.59, 37.64], 1900 m, 28.V.1989,leg. Schönmann & Schillhammer (1♀ NMW); Sümela, Altındere, [40.68, 39.65], 10.-11.VI.1969, leg. P. Brignoli (1 ♀ cBor). UNITED KINGDOM: EඇGඅൺඇൽ: Norwich, Norfolk, UEA Campus, [52.62, 1.23], 3.III.2013, leg. C. Billet (1 J cJen). UKRAINE: Kanev [Kaniv] near Kiew [49.75,31.46], 28.I.1988, leg. Zerche (2 ♀♀ SDEI); Ivano-Frankovo, ca. 40 km WNW of Lvov, Rostochye, State Reserve, [49.94,23.65] Quercus , Pinus etc.forest, 16-20.IX.1999, leg.S. Golovatch (1 J cRyv); N Yablunytsia vill., S Gorgany Mnts., valley Prut riv., [48.34, 24.45], 800m, pitfalls, 22-24.V.2018, leg.Panin R.(1J cGon). UZBEKIS-TAN: Tashkent, [41.29, 69.25], near railway station, plant residues, 24.V.1986, leg. S. A. Kurbatov (1 J 1 ♀ ZIN).

Redescription. Measurements JJ (n = 5): HW = 1.78– 1.89 (1.81); HL = 1.40–1.51 (1.46); HL/HW 0.79–0.83 (0.80); PW = 2.22–2.42 (2.31); PL = 2.04–2.20 (2.13); PL/PW 0.90–0.98 (0.92); EW = 2.44–2.51 (2.48); EL = 2.29–2.38 (2.32); EL/EW 0.91–0.96 (0.93); EL/PL 1.05– 1.14 (1.09); PW/HW 1.54–1.60 (1.58); forebody length 5.78–6.09 (5.91). ♀♀ (n = 5): HW = 1.80–1.96 (1.84); HL = 1.42–1.51 (1.46); HL/HW 0.77–0.81 (0.80); PW = 2.22–2.42 (2.32); PL = 2.09–2.13 (2.10); PL/PW 0.88–0.94 (0.91); EW = 2.40–2.60 (2.50); EL = 2.29–2.44 (2.34); EL/EW 0.91–0.97 (0.94); EL/PL 1.08–1.15 (1.11); PW/ HW 1.54–1.66 (1.59); forebody length 5.82–6.09 (5.90).

Large species; body black ( Fig. 7C ).

Head black, clearly transverse, with eyes very large (EyL/TL = 3.00–3.90 (3.54)) and clearly protruding, microsculpture of transverse waves, four punctures between anterior frontal punctures on frons ( Fig. 6C ); antennae and palpi darkened, except the joints, which are slightly paler, antenna with all antennomeres elongate. Thorax: pronotum black, slightly wider than long, wider than head, microsculpture of transverse waves, three punctures in dorsal row and two in sublateral row with the posteriormost puncture not reaching level of second puncture of dorsal row; scutellum smooth and glabrous, elytra black, uniformly pubescent, slightly wider than long, slightly longer than pronotum; legs dark brown to black with tarsi slightly paler.

Abdomen black, tergites uniformly punctured, clearly iridescent.

Male. Aedeagus ( Fig. 11D ): paramere broadly lanceolate, with clear medial attenuation, sensory peg setae forming two subapical, longitudinal rows along paramere edge apically but slightly turning inwards at half their length. Median lobe with two small but distinct teeth in front of basal part of parameral peg setae. Internal sac without clear sclerites.

Differential diagnosis. Even though Quedius fuliginosus differs from all other members of the fuliginosus -group by having the basal antennomeres at least slightly darkened, this is not always obvious in teneral or old museum specimens as coloration is altered. It is easily distinguished from Q. levicollis by the glabrous scutellum. It can be safely distinguished from Q. curtipennis by the aedeagi. In Q. fuliginosus , contrary to Q. curtipennis , the paramere has clear medial attenuation, a median lobe with the apical part being robustly pyramid-shaped and not laterally flattened, and an internal sac with no clear sclerotization. Subtle external difference between Q. fuliginosus and Q. curtipennis includes slightly stouter antennae, more transverse head, larger eyes clearly protruding thus making the head shape more quadrate, as well as slightly longer elytra in Q. fuliginosus . Also, Q. fuliginosus is very similar to Q. afrofuliginosus , but is easily recognized from the latter by having no additional punctures between anterior and posterior frontal punctures. It is also unclear if the two species have a distributional overlap.

Synonymic notes and neotype designation. Quedius fuliginosus was described from environs of Braunschweig, North Germany (GඋൺඏൾඇHඈඋඌඍ 1802). Two syntypes from the collection of Johann Christian Ludwig Hellwig (1743–1831) were revised by Gඎඌൺඋඈඏ (1993), who designated a lectotype to represent our current concept of Q. fuliginosus , while the paralectotype was identified as Q. curtipennis . Gඎඌൺඋඈඏ (1993) also studied the type of Quedius granulipennis ; he found it to be a junior subjective synonym of Q. fuliginosus .

Quedius latus was described from “Alagez-Gebirge” (HඈർHHඎඍH 1851), the Aragatz mountain area in Armenia and has never been recorded since the original description. Based on the rather extensive faunistic knowledge of the Caucasus region, Q. latus could either be conspecific to Q. fuliginosus or Q. curtipennis , both common in the area (AඌඌංඇG & SർHඳඅĸൾ 2019) and both fitting the original description of Q. latus . Since no type or any other authentic material of Q. latus was found in Hochhuth’s collection in Kyiv, Ukraine (S. Glotov, pers. comm.), apparently it is lost. To secure current and future stability of the name Quedius latus Hochhuth, 1851 , in accordance with the ICZN, Article 75, we designate its neotype. For this, a male specimen of Q. fuliginosus collected in the vicinity of the original type locality (for details see Type Material section above) has been selected. The neotype fits the original description of Q. latus and thus allows us to consider the so far dubious name Quedius latus Hochhuth, 1851 syn. nov. as a junior subjective synonym of Quedius fuliginosus Gravenhorst, 1802 .

The specimen we studied from the syntype series of Quedius molochinicolor Roubal, 1931 represents Quedius fuliginosus with slightly brownish elytra, thus confirming the currently accepted synonymy.

Bඋංඍඍൾඇ (1944) recognized that there were two closely related species of Quedius s.str. in Britain. Based on his descriptions and illustration, it is clear that Q. fuliginosus sensu Britten is Q. curtipennis while the new species he described, Q. subfuliginosus , is identical to Q. fuliginosus . This is in agreement with the currently accepted synonymy.

We were not able to check the types of Quedius impunctifrons Delahon, 1915 and Quedius jelineki Krása, 1904 from Germany and Czech Republic, respectively. Both names are currently in synonymy with Q. fuliginosus . Since the descriptions state that they lack the interocular punctures on frons characteristic of the Q. fuliginosus -group, they either represent variability in chaetotaxy – or more likely, they are dark variants of Q. molochinus , the latter possibility calling for a revised synonymy.

Sൺඐൺൽൺ (1965) described Q. viduus as being very similar to Q. fuliginosus , but different in the broader head, larger and more convex eyes, pronotum with lateral sides less convergent anteriorly, broader elytra, and rougher punctation of abdomen. This original description based on the single female specimen is the only record of Quedius s. str. from Japan. In spite of the thorough searches in the collections of National Museum of Nature and Science, Tokyo (Shuhei Nomura), Osaka Museum of Natural History, Osaka (Shigehiko Shiyake), Nakane Collection at Hokkaido University (Masahiro Ohara), and Sawada personal collection (Yasuhiko Hayashi), all performed by respective curators upon our request, the holotype of Q. viduus could not be located. Based on the original description and lack of any other species of Quedius s. str. in Japan, we suspect this is either a mislabeled specimen not from Japan, or an introduced specimen of Q. fuliginosus , which was not established or recollected in Japan. Thus, Q. viduus Sawada, 1965 syn. nov. is considered as a junior subjective synonym of Q. fuliginosus Gravenhorst, 1802 .

Bionomics. Quedius fuliginosus is widespread and eurytopic species found in a variety of forests (both deciduous and coniferous) and in the open habitats too. Usually, it occurs in moss and leaf litter or in various other kinds of ground debris (Sඈඅඈൽඈඏඇංĸඈඏ 2012). We have commonly collected this species in moss near the edge of a lake in Denmark ( Fig. 2A ), in various forests often with dense fern or grassy undergrowth, across larger parts of Western Siberia ( Fig. 2E ), in flood debris of a forested mountain stream in Triglav, Slovenia and along river forested on either side in the southern Czech Republic. It seems to prefer mossy patches and moist litter, and is thus to some degree often associated with water bodies, although it is not found at the very edge of these. It has also been reported from mole ( Talpa europaea ) nests (Nඈඐඈඌൺൽ 1990), from bracket fungi ( Polyporus squamosus ) (Sൾආൾඇඈඏ et al. 2015), and from ant nests (Gඈඋൾඌඅൺඏൾඍඌ 2010, 2016), all of which are presumably accidental occurrences, as there is no indication of the species preferring these habitats.

Larvae were described by Pൺඎඅංൺඇ (1941), Pඈඍඈඍඌĸൺඒൺ (1967) and Kൺඌඎඅൾ (1970). Pupae of the species are described in Sඍൺඇංൾർ (1999); Sඓඎඃൾർĸං (1966) described the egg. Adult females from Wytham, UK were observed containing eggs between May and June (Fඋൺඇĸ 1969).

Distribution. Quedius fuliginosus can be found across a large part of the Palearctic region ( Fig. 18 ). In Europe it is common in the North-East, but becomes rarer south--westwards and is probably absent from the southern parts of the Iberian Peninsula and Italy. It is found across most of Siberia from the Urals to at least the Baikal region. In the southern part of the range the species is confined to the forested zone in mountains. Here it is found along the Pyrenees, the Alps, the North Anatolian mountains, the Caucasus, and Transcaucasia. Eastwards it extends to the western forested regions of the Tian Shan Mountain Range in central Asia. The northern distributional border of the species seems to be determined by the forested boreal zone, as it does not continue into the tundra.

Records of Q. fuliginosus from Tunisia and Algeria (Fൺඎඏൾඅ 1902) most likely refer to either Q. levicollis or Q. afrofuliginosus , as we did not come across any specimens of this species from northern Africa or nearby regions. Moreover, Q. fuliginosus is apparently absent in the south Mediterreanean coastal areas, which makes its presence in North Africa even more unlikely.

Introduced to eastern North America in an area of Nova Scotia where the first specimens were found in 1996 (Mൺඃ- ĸൺ & Sආൾඍൺඇൺ 2007) ( Fig. 12 ).

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. 2. Habitats of Quedius s. str. A – Q. fuliginosus (Gravenhorst, 1802) in mossy habitat in a forest on Zealand, Denmark; B – Oak forest habitat of Q. levicollis Brullé, 1832 found in litter in Extremadura, Spain; C – Saline lake habitat of Q. meridiocarpathicus Smetana, 1958 found in bank litter on Peloponese, Greece; D – River bank habitat of Q. hispanicus Bernhauer, 1898 found in flood debris in Extremadura, Spain; E – Wet mixed forest habitat of Q. fuliginosus found in litter in Southern Ural, Russia; F – Alpine near tree line habitat of Q. subunicolor Korge, 1961 found under stones at 1700 m in Altai Mountains, Russia; G – Alpine mossy creek habitat of Q. unicolor Kiesenwetter, 1847 found in moss in Italian Alps (Photo:A. Zanetti).

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).