Quedius levicollis ( Brullé, 1832 )

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 : 252-254

Quedius levicollis ( Brullé, 1832)

( Figs 1 , 2B , 3 , 6D , 7D , 11B , 19 )

Staphylinus levicollis Brullé, 1832: 131 [Type locality: Morée: plaine de Modon]

Philonthus frontalis Nordmann, 1837: 76 [Type locality: Gallia; Symperopoli]

Quedius balthasari Mařan, 1935: 9 [Type locality: Knossos, Creta]

Staphylinus nigritarsis Ménétriés, 1832: 144 [Type locality: Caucase]

References. GඋൺඏൾඇHඈඋඌඍ (1802):34,(1847):231 (characters); SඍൾඉHൾඇඌ (1832): 215, (1839): 389 (characters); Fൺඅൽൾඋආൺඇඇ (1835): 123 (characters); Lൺർඈඋൽൺංඋൾ (1835): 376 (characters); Nඈඋൽආൺඇඇ (1837): 75 (distribution); Hඈඅආൾ (1837): 63 (distribution); Wൾඌඍඐඈඈൽ (1838): 167 (larva and pupa); EඋංർHඌඈඇ (1840): 537 (characters); Kඈඅൾඇൺඍං (1846): 22 (characters); Kංൾඌൾඇඐൾඍඍൾඋ (1848): 54 (characters); HඈർHHඎඍH (1849): 154, (1862): 46 (characters); RൾൽඍൾඇൻൺർHൾඋ (1849): 709, (1857): 201, (1874): 198 (characters); Fൺංඋආൺංඋൾ & Lൺൻඈඎඅൻජඇൾ (1856): 536 (characters); RඈඌൾඇHൺඎൾඋ (1856): 71 (distribution); Kඋൺൺඍඓ (1857): 502 (characters); Fൺංඋආൺංඋൾ & Cඈඊඎൾඋൾඅ (1860): 154 (distribution); Fൺඎඏൾඅ (1865): 293 (ecology); (1874): 289 (characters and distribution); (1878): 128, (1886): 58, (1902): 115 (distribution); Mඎඅඌൺඇඍ & Rൾඒ (1876): 688 (characters); Fඈඐඅൾඋ (1888): 237 (characters); GൺඇGඅൻൺඎൾඋ (1895): 403 (characters); Sൺංඇඍൾ- Cඅൺංඋൾ Dൾඏංඅඅൾ (1906): 102 (distribution); Pඈඋඍൺ (1907): 129 (characters); Rൾංඍඍൾඋ (1909): 111 (characters); JඈHൺඇඌൾඇ (1914): 363 (characters); Mඎඇඌඍൾඋ (1923): 196 (distribution); Gඋංൽൾඅඅං (1924):80 (characters and distribution); Pඈඋඍൾඏංඇ (1929):341 (characters); WඳඌඍHඈൿൿ (1938): fig. 22 (characters); TඈඍඍൾඇHൺආ (1949): 376 (characters); Hൺඋൽඒ (1851): 32 (distribution); Hൺඇඌൾඇ (1952): 142 (characters); SർHൾൾඋඉൾඅඍඓ (1956): 1102, (1965a): 407, (1965b): 511 (distribution); (1961): 133 (characters); Sආൾඍൺඇൺ (1958): 362 (characters and ecology); (1962): 134 (characters and distribution); (1963): 39, (1967): 558, (1970): 62, (1993):50 (distribution);(1978b):121 (distribution and ecology); Pൺඅආ (1963): 141 (characters and distribution); LඈHඌൾ (1964): 211 (characters); Fඈർൺඋංඅൾ (1964): 61 (distribution); Hඈඋංඈඇ (1965): 273 (distribution); Cඈංൿൿൺංඍ (1970):62 (distribution); (1978):192 (characters and distribution); Bඈඋൽඈඇං (1976a):93 (characters);(1982):140 (ecology); (1984):342 (distribution); Pඈඉൾ (1977): 31 (characters and distribution); Hංඇඍඈඇ (1981): 662 (characters of egg); TඬඍH (1984): 118 (distribution); Fඋൺඇඓ (1987): 70 (distribution); Gඎඌൺඋඈඏ (1993): 73 (synonymy); WൾඅർH (1993): 229 (ovary structure); Cංർൾඋඈඇං & Zൺඇൾඍඍං (1995): 33 (distribution); Nൾඐඍඈඇ (1995): 51 (requested to ICZN to place Q. levicollis on the Official List of Specific Names in Zoology); ICZN (1996): 215 ( Q. levicollis placed on Official List of Specific Names in Zoology); Bඅൺඇൽ & Sංඇർඅൺංඋ (1997): 204 (biology); MൺඋർHൺඅ & CH ൺඋൽඈඇඇൾඍ (2000): 72 (distribution); Oඐൾඇ (2000): 252 (ecology); AඌඌංඇG & Wඎඇൽൾඋඅൾ (2001): 37 (distribution); UHඅංG et al. (2006): 54 (distribution); Tඋඈඇඊඎൾඍ (2006): 102 (distribution); Aඇඅൺ෡ (2009): 229 (distribution); Sඍൺඇ (2009): 242 (distribution); Vඈඋඌඍ et al. (2009): 16 (distribution); Mൺඃඓඅൺඇ & BඈHගඹ (2012): 182 (distribution); Bൾൻൾඋආൺඇඌ et al. (2016): 135 (distribution); ÖඓGൾඇ et al. (2016): 620 (distribution); Rඎඍൺ et al. (2016): 37 (distribution); Sൺඅඇංඍඌĸൺ & Sඈඅඈൽඈඏඇංĸඈඏ (2019): 49 (distribution).

Type material examined. Staphylinus nigritarsis : Lൾർඍඈඍඒඉൾ: designated by Gඎඌൺඋඈඏ (1993) ( ZIN), labelled: “[golden square] / Caucas. / nigritarsis Menet. Cauc ”. Pൺඋൺඅൾർඍඈඍඒඉൾ: labelled: “[golden square] / Caucas.” (1 J ZIN).

Additional material examined. ALBANIA: 13km SE Elbasan, Mali i Shpatit, Mt. Bukanikut, 41.0431, 20.2172, brook bank in beech forest, 1500m, 22.V.2010, leg. D.W.Wrase (1J cSch). BOSNIAAND HERZE-GOVINA: Maklen-Pass,[43.85,17.65], 1902, leg.O.Leonhard (1 SDEI); Prozor, [43.82, 17.61], 1902, leg. O. Leonhard (1 SDEI). BULGARIA: Arkutino, S of Burgas, [42.41, 27.65], 28.V-9.VI.1971, leg. Wallis (1 J SDEI). CROATIA: Capela [Kapela], [44.89, 15.52], leg. Heyden (1 SDEI); Velebit Mountains, [44.52, 15.22], wine traps, V.2005 (1 J cJen). CYPRUS: Agios Nikolaios, [34.86, 32.75], 16.IV.1995, leg. H. Schmid (1 J NMW); Troodos Mountain, Stavros tis Psokas, [35.02, 32.63], 800- 950m, 12.-18.IV.2010, leg. I. Wolf (1 J cSch). DENMARK: Ovestrup Hede, 56.2412, 8.9297, pitfall trap, heathland grazed part, 7-22.VI.2017, leg. E. Kristensen, D. Byriel, & M. Justesen (1 J NHMD). FRANCE: Avignon, [43.94, 4.81], 22.V.1949, leg. R. Skovgaard (1 J NHMD); Chigny n. Oise, [49.91, 3.76], 19.X.1918, leg. Salchert (1 ♀ ZMHB); Paris, [48.83, 2.41], 19.V.1949, leg. R. Skovgaard (1 J NHMD). Cඈඋඌൾ: Bocognano,[42.08,9.06],1905, leg.O.Leonhard (1 SDEI);Col de Bavella, [41.79, 9.23], 1240m, East slope, 25.VII.1994 leg. L. Zerche (1 J SDEI); Foret de Valdo-Niello, Evisa, [42.28, 8.91], 1000-1200 m, 27.VII.1967, leg. Langemark-Lomholdt (1 J NHMD). GERMANY: Berlin-Marzahn, Hellersdorfer Berg, [52.53, 13.58], 8.VIII.1997, leg. Hieke (1 ♀ ZMHB); Berlin, Prenzlauerberg Hof, [52.53, 13.46], 29.V.1985, leg. I. Uhlig (1 J ZMHB); Grunewald n. Zehlendorf, [52.48, 13.22], leg.Wagner (1 SDEI). GREECE: Attica, [38.05, 23.84], 17.IV.1922, leg. W. Liebmann (1 SDEI); Paleokastro, Halkidiki, [40.43, 23.42], 800m, 11.VII.1983, leg.A. Casale (1 ♀ cBor); 2 km W Panagia, Trikala, [39.79, 21.33], 15.VI.1982, leg. R Danielsson (1 J MZLU); Voria Pindos, Morfi, [40.21, 21.21], dry slope, 26.VI.1997, leg. Winkelmann (1 ♀ cSch); Cඈඋൿඎ: Pandokratoras, 39.7492, 19.8701,N-slope with scattered pine,decidious trees,and bushes, litter and roots between rocks sifted, 800 m, 4.VI.2017, leg. Schülke (1 J cSch);Pantokrator,[39.74,19.87], V.1964, leg.Palm (1♀ MZLU). Cඋൾඍൾ: Georgioupouli, [35.36, 24.26], 24.-25.IV.1995, leg. B. Jaeger (1 ♀ cSch); Ida mts., ca. 6 km S Analipsis, 35.1752, 24.8391, Quercus forest, 1260m, 21.X.2006, leg. M. Schülke (1 J cSch); Mt.Ida, 28km SW Iraklio, [35.23, 24.77], 1600m, 7.IX.1991, leg. N. Esser (1 ♀ NHMD). KൺඋඉൺඍHඈඌ: Karpathos, 35.5097, 27.2189, 40 m, terracod grassy fallow, 01.I.2016, leg. V. Assing ( ZMHB). Lൾඌൻඈඌ: SW Agiassos, road to Olympos peak, [39.06, 26.36], 700 m, 10.VI.1996, leg. Jäch (1 J 1 ♀ NMW). Pൾඅඈඉඈඇඇൾඌ: Korinthia, Kyllini, Zoria, [37.94, 22.43], 2000 m, 3.VI.1995, leg. Giachino & Vailati (1 J 3 ♀♀ cBor); Kyllini Mountain, [37.97, 22.39], 2100m, North side, below summit over valley, 9.VI.1996, leg. P. Wunderle (1 ♀ cSch); Sparta, [37.06, 22.43], 3.V.1962, leg. Skovgaard (1 J NHMD); N Taygetos, 6 km NE Thouria, 37.1401, 22.1284, litter and soil under bushes, sifted, 584 m, 20.IV.2015, leg. M. Schülke (1 ♀ cSch). RHඈൽൾඌ: Prophitis Ilias, 36.2734, 27.9287, pine forest, under stones & sifted from litter & moss, 685m, 17.V.2015, leg. Wrase & Laser (1 J cSch). IRAN: Kohgiluyech-Va-Boyern-Ahmad province, 20 km SW Yasuk, 10 km SE Sepidor, [30.56, 51.51], 5.-6.V.2007, leg.A. Anistchschenko (1 ♀ SDEI). IRELAND: Inch Dingle Eire, [52.14, -9.97], IX.2012, leg. J.J. Shaw (1 J cJen); Pembury, Kent 51.1625, 0.3431, 24.VII.2011, leg. J.J. Shaw (2 JJ cJen). ISRAEL: Upper Galilee,Har Avivim,W Avivim, 33.0833, 35.4923, open stony grazing land limestone, 768m, 21.IV.2006, leg. D.W.Wrase (1 J cSch); Upper Galilee, Meron Mts. Har Meron, Kamin Rom, 32.9908, 35.4111, open stony grazing land, limestone, 1100m, 28.IV.2006,leg.D.W. Wrase (1J cSch). ITALY: CalabriaAspromonte,[38.21, 15.99],1905,leg. Pagametti (1 SDEI); Calabria, Roseto Capo Spulico, [39.98, 16.61], 20 m, 6.X.1998 leg.F.Angelini(1♀ NHMD); Dicomano, Toscana,[43.89,11.52], I.1971, leg Bordoni (1J cBor); Macerata, [43.29, 13.45], 1.X.1955, leg.D. Caccamicelo (1♀ cBor); Marche,Montemonaco, Isola San Biagio,[42.91, 13.31], fields, 900 m, 27.VII.-1.IX.2004, leg.M.Rismondo & R. Fabbri (2 JJ 2♀♀ NHMD); same locality, black hornbeam forest, 1.IX.-7.X.2004, leg. M. Rismondo & R. Fabbri (1 J NHMD); Modena, Prignano sulla Secchia, La Quercia, [44.45, 10.73], pitfall traps, 400 m, 5.-18.VII.2006, leg. R. Fabbri (1 J NHMD); Reggio di Calabria, river Sant Agata, [38.09, 15.66], 11.III.1959 (1 J cBor); Settecamini, Rome, Lazio, [41.93, 12.62], 23.Xi.1969 (1 J 1 ♀ cBor); Sovara, Anghiari, [43.53, 12.03], 19.V.1930, leg. A. Andreini (1 ♀ cBor). Sൺඋൽංඇංൺ: Aritzo Mountain, Gennargentu, [40.01, 9.29], leg. C.Kruger (2 SDEI); Ottana, [40.23, 9.04], 15.X.1974 (3 JJ 1♀ cBor);Oristano,[39.89,8.59], 3.V.1975,leg.S. Ripze (2♀♀ cBor); Sංർංඅඒ: Ficuzza, [37.85, 13.38], 1906, leg. O.Leonhard (1 SDEI);Messina, [38.18, 15.54], leg. Schiödte (4 ♀♀ NHMD). MOROCCO: Ain Leuh, [33.28, -5.33], 17.V.1925, leg. M. Escalera (1 J 4 ♀♀ MNCN); Ar Rif, Chefchaounen reg., env.Bad Bessen, 34.9913, -4.8555, 1124 m, 3.VI.2007, leg. P. Hlaváč (1 J cSch); Ar Rif, Chefchaounen region, rd. Bad Berret - Bad Bessen, 1392 m, 35.0169, -4.0035, 3.-6.VI.2007, leg. P. Hlaváč (1 ♀ cSch); Taza, [34.22, -4.00], leg. F. Schubert (1 ♀ NMW). PORTUGAL: Penedo da Meditação, Coimbra, [40.224371, -8.40], 24.IV.1961, leg.R.N. F. (1 J NMW). Mൺൽൾංඋൺ: Faja da Nogueira, [32.74, -16.90], laurisilva with big Ocoteas, 750 m, 20.IX.1980, leg. H. Enghoff & O. Martin (2 JJ 3 ♀♀ NHMD); Paul da Serra, [32.73, -17.05], 27.XII.1987, leg.Gillerfors (1 J MZLU). SPAIN: Castanos de Calabazas, 1 km S Castanar de Ibor, Extemadura, 39.6235, -5.4128, 860 m, sifted mossy rock scree litter, chestnut and oak forest /w rock screes and small stream, 10.V.2019, leg.A. K. Hansen, J. J. Shaw, & J. Kypke ( NHMD); Cavarrubias [Covarrubias], Burgos,[42.05, -3.51], 10.XI.1995, leg.A.Anichtchenko(1J cSha); Lloret de Mar, [41.69, 2.83], 17.IX.-6.X.1961, leg. Palm (1 J MZLU); Sierra de Ancares,A Degrada vill., [42.81, -6.87], 1100-1250 m, 8.-9.VII.1996, leg. Wrase (2 JJ cSch); Sierra de Guadarrama, Navacerrada,Arroyo de Peca [Peña] Cabrita, [40.77, -3.99], 1800 m, 5.IX.2001, leg.A.Anichtchenko (1 J NHMD); Sierra Nevada,rio Lanjaron,Tello,[36.92, -3.47], 1500 m, 29.- 31.III.2010, leg.Shavrin &Anichtchenko (1J NHMD);Toledo,El Pielago Navamorcruende, [40.15, -4.79], 4.VIII.2005 leg. Lencina & Andujar (1 ZMHB). SYRIA: Krak des Chevaliers, [34.75, 36.29], 600 m, 7.V.1982 (1 J 1 ♀ MCZ); MashtalHelu [Mashta Al Hilu], E of Safita, [34.87, 36.26], 30.IV.2008, leg.Skoupý (1 J cSch); Zainie, 45 km NE of Latakia, [35.78, 36.19], 4.V.1982, leg. M. Dvorák (3♀♀ MCZ). SWEDEN: Fൺඋද: Farön, [57.94, 19.14] (1 MZLU). GඈඍඌKൺ Sൺඇൽදඇ: Sandön, [58.36, 19.24] (1 MZLU). Öඅൺඇൽ: Stenåsa, [56.51, 16.59] (1 MZLU). TURKEY: Amanos [Nur Mountains], [36.74, 36.33], 24.V.1987, leg. Jäch (1 ♀ NMW); Bolu, [40.72, 31.61], leg. F. Schubert (1 ♀ NMW); Çıkrıkçı, Turgutlu, [38.47, 27.81], 15.II.2006, leg. S.Anlaş (1 J NHMD); Çıkrıkçı, Turgutlu, [38.47, 27.81], 17.II.2006, leg.S.Anlaş (1J 3♀♀ NHMD);Danin-Danin, Hakkari Prov, [37.44,42.86], 2200 m, 1.VI.1987, leg.Schönemann & Schillhammer (2 JJ 7♀♀ NMW); Izmir, Boz daglari, Boz dag köy env., [38.34, 27.75], 1500-1700 m, 30.V-3.VI.2003, leg. R. Lohaj (1 J 1 ♀ cSch); 50 km N Antalya, [37.33, 30.77], 18.IV.2015 leg. Snizek (1 ♀ NMEG); Sogukoluk, Belen, Hatay, [36.49,36.14], 24.V.1987, leg.Schönemann &Schillhammer (1 ♀ NMW). UNITED KINGDOM: Sർඈඍඅൺඇൽ: Bridge of Orchy, [56.51, -4.76], 23.VII.1987, leg. Gillerfors (1 J 1 ♀ MZLU); Hawock [55.42, -2.78], VI.1964, leg. M. Paoletti (1 ♀ NMW).

Redescription. Measurements JJ (n = 6): HW = 1.78– 1.87 (1.81); HL = 1.47–1.56 (1.51); HL/HW 0.81–0.85 (0.83); PW = 2.27–2.36 (2.31); PL = 2.22–2.24 (2.23); PL/PW 0.94–0.98 (0.96); EW = 2.60–2.69 (2.63); EL = 2.49–2.60 (2.54); EL/EW 0.95–0.98 (0.96); EL/PL 1.12– 1.16 (1.14); PW/HW 1.49–1.58 (1.53); forebody length 6.20–6.33 (6.28). ♀♀ (n = 4): HW = 1.80–1.87 (1.84); HL = 1.49–1.56 (1.52); HL/HW 0.81–0.86 (0.83); PW = 2.38–2.51 (2.44); PL = 2.27–2.36 (2.31); PL/PW 0.94–0.95 (0.95); EW = 2.73–2.89 (2.82); EL = 2.58–2.69 (2.64); EL/EW 0.92–0.97 (0.94); EL/PL 1.09–1.17 (1.14); PW/ HW 1.53–1.69 (1.61); forebody length 6.42–6.56 (6.47).

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

Head black, slightly transverse, with eyes large (EyL/TL = 3.64–4.78 (4.22)) following contour of head; microsculpture of transverse waves, 4 to 6 punctures between anterior frontal punctures on frons ( Fig. 6D ); antennae pale reddish, long, all antennomeres clearly elongate; palpi pale reddish.

Thorax: pronotum black, almost quadrate, as wide as long, clearly wider than head, microsculpture of transverse waves, three punctures in dorsal row and three in sublateral row with the most posterior one reaching the level of third puncture of dorsal row; scutellum punctured and setiferous, elytra black, densely punctured and pubescent, slightly wider than long, slightly longer than pronotum; legs dark brown with inner face of tibia darkened and tarsi paler.

Abdomen black, tergites uniformly punctured, slightly iridescent.

Male. Aedeagus ( Fig. 11B ): Paramere slender, lanceolate, reaching apex of median lobe, sensory peg setae forming two clear rows; median lobe with protrusion at apices and two small but distinct teeth near lower level of parameres peg setae; internal sac with two small distal medial sclerites.

Differential diagnosis. Quedius levicollis is easily distinguished from other members of the fuliginosus -group by the punctured and setiferous scutellum and three punctures in the sublateral row of pronotum with the posteriormost one reaching the level of the third (basalmost) puncture of the dorsal row. It can be additionally distinguished from Q. fuliginosus by the pale antennae, and from Q. curtipennis by the slender paramere and median lobe with apical protrusion. Quedius levicollis is mainly found in drier habitats and rarely co-occurs with any of the other species in the species group.

Synonymic notes. The name Quedius tristis ( Gravenhorst, 1802) has long been applied to this species. GඋൺඏൾඇHඈඋඌඍ (1802) did not describe Staphylinus tristis as a new species but attributed it to Fabricius. The Fabrician species is now in Ocypus (Pseudocypus) . Nൾඐඍඈඇ (1995) made an appeal to the International Commission of Zoological Nomenclature requesting that the species takes the name Quedius levicollis , as this was the next available name associated with this species. As a result, Quedius levicollis ( Brullé, 1832) was accepted on the Official List of Specific Names in Zoology ( ICZN 1996: 215).

BൾඋඇHൺඎൾඋ & SർHඎൻൾඋඍ (1916) placed Staphylinus nigritarsis Ménétriés, 1832 in synonymy to Philonthus splendens . Gඎඌൺඋඈඏ (1993) revised the type series of S. nigritarsis and designated a lectotype. He found S. nigritarsis to be conspecific with Q. levicollis ( Brullé, 1832) based on both external characters and the shape of the aedeagus. We confirm Gusarov’s identification through examination of photos of the type from ZIN provided by M. Salnitska (see supplementary Fig. S1 ).

We were not able to examine the types of Philonthus frontalis Nordmann, 1837 and Quedius balthasari Mařan, 1935 , both currently accepted synonyms of Q. levicollis . Based on the original descriptions, both synonymies appear to be correct. Philonthus frontalis was described from Italy and Greece and fits the description of Q. levicollis , with interorbital punctures and punctate scutellum. Quedius balthasari Mařan, 1935 , described from central Crete as a species different from Q. levicollis in sparser punctation and reddish margins of elytra is likely a color aberration because Q. levicollis is also known from Crete.

Bionomics. Quedius levicollis is a clearly thermophilous eurytopic species, which prefers sandy soils and is found in both open habitats and dry forests. It occurs in various ground debris and under rocks, e.g., it has been found in forest litter at different elevations (200–1900 m) in Cyprus (AඌඌංඇG & Wඎඇൽൾඋඅൾ 2001), under rocks in Corse (MൺඋർHൺඅ & CH ൺඋൽඈඇඇൾඍ 2000), and from sifted wet leaf-litter under oleander trees, at 500–1100 m in Israel (Sආൾඍൺඇൺ 1978b). We have encountered this species under rocks in open oak and chestnut forest in Spain ( Fig. 2B ), and between the roots of Erica in a heathland in Denmark. It has been recorded from caves in Sardinia (Bඈඋൽඈඇං 1982), although this most likely represents an accidental occurrence.

WൾඅඌH (1993) found this species to contain six ovaries with a fasciculata type of structure.

Distribution. Quedius levicollis is widely distributed across the Western Palearctic from the Atlantic coast, including British Islands through the Mediterranean, including North Africa, in the west to the Russian Black Sea coast and the Middle East ( Fig. 19 ). In the Middle East the species extends into the Levant region, but not into the Nile delta. The south-easternmost record of Q. levicollis is from South Iran. The species is found on all major Mediterranean islands, including Cyprus, Crete, Corse and Sardinia, as well on the island of Madeira in the Atlantic Ocean. It is most common in the southern part of its distribution range gradually becoming rarer towards the north-east where it occurs at xerothermic sites reaching its northernmost limit on the Swedish island of Gotland in the Baltic Sea.

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. 19. Distribution of Quedius levicollis Brullé, 1832 (top), Q. hispanicus Bernhauer, 1898 and Q. pallipes Lucas, 1846 (middle), and Q.balticus Korge, 1960 (bottom).