Quedius molochinus ( Gravenhorst, 1806 )

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 : 259-261

Quedius molochinus ( Gravenhorst, 1806)

( Figs 1 , 4 , 9 , 8C,D , 12 , 13D , 20 )

Staphylinus molochinus Gravenhorst, 1806: 56 [Type locality:Brunsviga]

Quedius denudatus Stephens, 1832:216 [Type locality: London; Suffolk]

Quedius lathburii Stephens, 1832: 218 [Type locality: London; Suffolk, Orwell]

Quedius niger Sahlberg, 1876: 229 [Type locality: Muonioniska]

Quedius maculicornis Mulsant & Rey, 1876: 694 [Type locality: les montagnes du Lyonnais, à la Grande Chartreuse , dans les Pyrénées]

Raphirus picipennis Stephens, 1833: 243 [Type locality: London]

Staphylinus laevicollis Runde, 1835: 6 [Type locality: Halae]

Staphylinus lapponicus Zetterstedt, 1838: 61 [Type locality: Lapponia Umensis, Lycksele; Wilhelmina; Gaskelougt]

References. GඒඅඅൾඇHൺඅ (1810):302, (1827):477 (characters);SൺHඅൻൾඋG (1830): 314 (characters); (1876): 24 (distribution); MൺඇඇൾඋHൾංආ (1830): 25, (1831): 439 (distribution); (1843): 232 (characters); Rඎඇൽൾ (1835): 5 (characters); Lൺർඈඋൽൺංඋൾ (1835): 377 (characters); Cඎඋඍංඌ (1837): pl. 638 (characters); Nඈඋൽආൺඇඇ (1837): 76 (distribution); EඋංർHඌඈඇ (1839): 489, (1840): 535 (characters); Hൾൾඋ (1839): 276 (characters); SඍൾඉHൾඇඌ (1839):390 (characters);Lൺඉඈඋඍൾ (1840):177 (characters);Kංൾඌൾඇඐൾඍඍൾඋ (1848): 52 (characters); (1851): 419 (distribution); GඋൺඏൾඇHඈඋඌඍ (1847): 227 (characters); RൾൽඍൾඇൻൺർHൾඋ (1849): 709, (1857): 201, (1874): 198 (characters); Hൺඋൽඒ (1851): 32 (biology); Fൺංඋආൺංඋൾ & Lൺൻඈඎඅൻජඇൾ (1856): 537 (characters), Kඋൺൺඍඓ (1857): 500 (characters); WൺඍൾඋHඈඎඌൾ (1858): 23 (synonymy); THඈආඌඈඇ (1859): 25 (notes on types); (1860): 173 (synonymy); GൾආආංඇGൾඋ & Hൺඋඈඅൽ (1868): 570 (synonymy); Fൺඎඏൾඅ (1874): 290 (characters and distribution); Sൾංൽඅංඍඓ (1875): 267 (characters); Mඎඅඌൺඇඍ & Rൾඒ (1876): 692 (characters); Fඈඐඅൾඋ (1888): 237 (characters); Rൾංඍඍൾඋ (1909): 112 (characters); JඈHൺඇඌൾඇ (1914): 364 (characters); Gඋංൽൾඅඅං (1924): 81, (1962a): 320 (characters); Pඈඋඍൾඏංඇ (1929): 341 (characters); MඣඊඎංGඇඈඇ (1937): 3 (synonymy); Hൺඇඌൾඇ (1952): 142 (characters); Sආൾඍൺඇൺ (1958): 364 (characters and biology); (1962a): 136, (1965a): 38 (characters); (1962b): 134 (larval characters); (1964): 80, (1976a): 172, (1978a): 85, (1981): 638, (1990): 98, (1993): 50 (distribution); (1971): 133 (distribution; introduction to Nearctic); (1973): 1427 (introduction to Nearctic);KඈඋGൾ (1960b): 53 (characters); Cඈංൿൿൺංඍ (1961): 52, (1963): 423, (1978):194 (characters); Pൺඅආ (1963a):141 (characters); LඈHඌൾ (1964): 212 (characters); Dඏඈřák 1965: 89 (distribution); Pඈඍඈඍඌĸൺංൺ (1967): 82 (larval characters) Kൺඌඎඅൾ (1968a): 66 (biology); (1970): 61 (larval characters); Sඓඎඃൾർĸං (1968): 736 (distribution); Oඌൾඅ-අൺ & Zൺඇൾඍඍං (1975): 120 (biology); Bඈඋൽඈඇං (1976a): 95 (characters); SHංඅඈඏ (1975): 376 (distribution); Pඈඉൾ (1977): 31 (distribution); TඬඍH (1984):120 (characters);SൾGൾඋඌ(1987):182 (morphology);WൾඅർH (1993): 229 (morphology);Cංർൾඋඈඇං & Zൺඇൾඍඍං (1995):32 (distribution); Sඍൺඇංൾർ (1996):117 (pupal characters);Dඈඐඇංൾ &Aඋඇൾඍඍ (1996):391 (characters); Oඐൾඇ (2000): 252 (biology); AඌඌංඇG (2001): 75 (biology); UHඅංG et al. (2006): 55 (distribution); Mൺඃĸൺ (2007): 949 (distribution); Dൺඉĸඎඌ & Tൺආඎඍංඌ (2008): 33 (distribution); Cඎඉඉൾඇ et al. (2011): 15 (distribution); Sൾආංඈඇൾඇĸඈඏ et al. (2015): 335 (distribution).

Type material examined. Staphylinus molochinus : Sඒඇඍඒඉൾ J ( ZMHB), labelled:“6230 / Braunsberg Dahl leg./ molochinus gr.gyll. / (Para)? typus Grav. / det. H. Korge Quedius molochinus Grav. Typus!”.

Quedius niger : Sඒඇඍඒඉൾ ♀ ( MZH), labelled:“Tjudi / J. Sahlbg./ 1617. / Mus.Zool.H:fors Spec.typ. No 157 Qued.molochinus var.niger J. Sbg. / Syntype / http://id.luomus.fi/GZ.83 / Photographed 2020 Pekka Malinen”. Additional material examined. AUSTRIA: Arbesbach, [48.49, 14.95], 13.VII.1957, leg. Schubert (1 ♀ NMW); Admont, [47.56, 14.45], leg. H. Franz (1 ♀ NMW); Prater, [48.21, 16.39], 19.IX.1964 (1 J NHMD). CZECH REPUBLIC: Iser [Jizera] River, n. Benešov u Semil, [50.60, 15.36], 350 m, 24.VII.1982, leg.Hieke (1♀ ZMHB);Pastviny n.Žamberk, [50.09, 16.56], 31.VII.-3.VIII.1982, leg. Hieke (1 ♀ ZMHB); Podyji Nat. Park,forest, at river Dyje valley ca. 2.5 km NW of Havraníky,48.83,15.98, 240 m, sifted flood debris, 08.VI.2016, leg. A. Solodovnikov, J. Jenkins Shaw,&A. Hansen (1J NHMD); Silesia Altvater [Praděd], [50.08, 17.23], leg. Letzner (1 SDEI); Teplice nad Metují, [50.59, 16.16], 3.-5.VIII.1992, leg.Hieke (1♀ ZMHB);Vysoká Hola, NW Karlov, Malá Morávka, [50.08, 17.23], sifting litter of Pinus mugo , 25.VII.1993, leg. Zerche (1 SDEI). DENMARK: Hjelm Hede, [56.49, 8.91], 20.VII.1951, leg. J. Petersen (2 ♀♀ NHMD). BඈඋඇHඈඅආ: Øle Å, 19.IV.1965, [55.05, 15.02], leg. F. Bangsholt (1 ♀ NHMD). Fඎඇൾඇ: Bjergsted, [55.66, 11.35], 1.VI.1981, leg. M.Hansen (1 J NHMD);Ristinge Klint, [54.82, 10.61], 14.VII.1978, leg. M. Hansen (2 JJ NHMD). Zൾൺඅൺඇൽ: Furesøstien, [55.81, 12.41], 27.IX.1954, leg. J. Petersen (3 ♀♀ NHMD); Holmegårds Mose, [55.29, 11.82], 26.VI.1975, leg. M. Hansen (1 J NHMD); Ormø, [55.21, 11.45], 13.IX.1997, leg. O. Martin (1 ♀ NHMD); Rude Skov , [55.83, 12.47], 19.VIII.1948, leg. J. Petersen (1 ♀ NHMD). FINLAND: Kilpisjärvi, [69.04,20.81], 21.-22.VII.1985, leg.Hieke (1♀ ZMHB);Kiuruvesi,[63.65, 26.62], 6.VII.1951, leg. Linnavuori (1 J 2 ♀♀ ZMUO); Mutenia, Sodankylä,[68.02, 27.41], 13.IX.2001, leg.J.Itämies, O. Nenonen (1♀ ZMUO); Rymättylä,[60.37, 21.94],leg.Linnavuori (1♀ ZMUO);Saana,Enontekiö, [69.04,20.85], 10.VII.2013, M.Pentinsaari (1J ZMUO). FRANCE: Chigny,Oise River, [49.91, 3.77],Selchert (1J ZMHB). GERMANY: Bayern, Schleissheim,[48.24, 11.59], 15.V.1931, leg. Ihssen (1J ZMHB); Berlin--Marzahn, Hellersdorfer Berg, [52.53,13.58], 8.VIII.1997, leg. Hieke (1J ZMHB); Brunn, Dosse [52.91, 12.51], 16.III.1980 leg. E. Rössner (2 JJ SDEI); Engolling,Auerbach, [48.79,13.10], 5.IX.1978, leg.K.Koufrauen (1 J 1 ♀ ZMHB); Erzgebirge, Flohtelberg, [50.42, 12.95], 2.VI.1925, leg Uhmann (1 J SDEI); Finkenkrug nr. Berlin, [52.56, 13.03], 12.IV.1898, leg. Dahl (1 ♀ ZMHB); Flessenow, Schwerin, [53.75, 11.49], 4.VII.1968, leg.Uhlig (1 ZMHB); Hainleite, SW Günseroda and Seega,[51.32, 11.02], 10.VIII.1993, leg. F. Hieke (1 J ZMHB); Marburg in Hess, [50.81, 8.77], 16.IV.1904, leg. Strand (1 J ZMHB); Neuendorf, Teupitz, [52.11, 13.58], 30.IV.1992, leg. Renner (1 J ZMHB); Sarnow n.Anklam, [53.75, 13.63], 29.IX.1964, leg. F. Hieke (1 J ZMHB); Strausberg, 23.VII.1981, [52.56, 13.88], leg. H. Wendt (1 ♀ ZMHB); Thüringer Wald nr. Scheibe, [50.48, 11.05], under stones, 8.I.1957, leg. Dieckmann (1 SDEI); Willersdorf, [51.02, 8.85], VII (1 ♀ ZMHB). ITALY: Gello, [43.96, 10.45], VI.1922, leg. A. Strand (1 J NHMD); Zeccone n. Pavia, [45.26, 9.19], 17.X.1977, leg. P. Kanaar (1 J ZMHB). NORWAY: Hemnesberget, [66.22, 13.61], 8.-14.VII.1903, leg. E. Strand (2 JJ ZMHB); Kongsvinger Eis, [60.19, 12.01], 27.X.1980, leg. F. Midtgaard (1 J NHMD); Pasvik Lake, Sydvaranger, [69.34, 29.53], VI.1994, leg. G. Nietsch (1 J NMW); Utgardsjön, Rønningen,Austmarka, [60.01, 12.37], 28.VI.-10.VII.1975, leg.P.Kanaar (1 ♀ ZMHB). POLAND: Misdroy, [53.91, 14.44] (1 J ZMHB); Pölitz [Police], Pommern,[53.55, 14.57], 13.VI.1920,leg.E. Hanau (1♀ ZMHB); Riesengebirge [Krkonoše], [50.79, 15.57], 750m, moss, 16.X.1902, leg. Dahl (1 J ZMHB); Tamsel, [52.62, 14.70], 20.V.1920, leg. Kuntzen (1 ♀ ZMHB). RUSSIA: CH ංඍൺ Oൻඅൺඌඍ: Malkhanskiy Khrebet, pass to Maleta, 20 km S Maloarkhangel’sk, 50.2352 108.916, 19.-25.VIII.1998, leg. A. Shavrin ( NHMD). KHൺඇඍඒ- Mൺඇඌං Aඎඍඈඇඈආඈඎඌ Dංඌඍඋංർඍ: Sovetsky region, Malaya Sosva Nature Reserve, near Cordon Hangokurt, road to Belaya Gora, 61.9504, 64.2489, 81 m, nr a roadside puddle with Polytrichum sp. on clay, 28.VI.2017, leg. A.B. Ryvkin (1 J cRyv). Kඈආං Rൾඉ: Ukhta surrounding,[63.57, 53.71], 29.VI.1974, leg.V.Shulov (6JJ 6 ♀♀ ZMHB). KඋൺඌඇඈඒൺඋඌK Kඋൺං: Niznaya Lebedyanka river, [61.96, 89.45], 23.VI.1992, leg. V.S. (1 J cRyv); Turukhanskiy Distr., Mirnoye, [62.28, 89.03] 6.-17.VII.1990, leg. L.B. Rybalov (1 J 1 ♀ cRyv). Lൾඇංඇ-G උൺൽ Oൻඅൺඌඍ: Saint-Petersburg, Sestroretsk, Rzhavaya kanava, [60.12, 29.96], 16.VIII.2009, leg.V.N.Prasolov (1J ZIN); Stary Petergof, [59.87, 29.87], forest, 7.VI.1997, in leaf litter, leg.A. Solodovnikov (1J NHMD) Yaschera, [58.92, 29.98], 18.V.1996, leg.L.Anisutkin (1J NHMD). Mඈ-ඌർඈඐ Oൻඅൺඌඍ: Moscow,Izmajlovskij Park,[55.75,37.62], 7.VII.1983, leg. A.Isaev (1J NHMD); Odintsovsky Distr., Zvenigorod Biological Station of MSU, [55.70, 36.72], mixed forest, 22.VIII.1984, leg. K.G. Mikhailov (2 JJ cRyv). MඎඋආൺඇඌK Oൻඅൺඌඍ: Kyndomys [Kindo] Peninsula,[66.65, 33.22], forest floor, 16.VII.1992, leg. Wegner (1 J ZMHB). Pൾඋආ Kඋൺං: Perm city, [58.01, 56.22], pitfall, 29.VIII.1978, leg. S.L. Esunin (1 J NHMD). Rൾඉ. ඈൿ Bඎඋඒൺඍංൺ: Barguzinsky Distr., Barguzin valley, 20 km E Ust-Barguzin, Gusikha, [53.42, 109.42], aspen forest, pitfalls, leg. V. Shilenkov (1 J NHMD); Vitimsky Nat. Preserve, lake Oron, mouth or Labaznyj creek, Oron hut, [54.82, 112.26], 27-30.VII.2000, leg. A. Shavrin (2 JJ NHMD). Rඒൺඓൺඇ Oൻඅൺඌඍ: Ryazan Prov., Spassky Distr., Oka nature reserve, [54.71, 40.85], leaf litter, 26.VI.1981, leg. Kormylutsyn & Eskov (1J 1♀ cRyv). SඏൾඋൽඅඈඏඌK Oൻඅൺඌඍ: Ural,Visim Nature Reserve, [57.37, 59.77], 25.I.1985, leg L.P. Titova (2 JJ cRyv). Tඒඎආൾඇ Oൻඅൺඌඍ: Surgutsky Distr., Yugansky Nature Reserve, Ai-Magromsy River Basin, near ‘Medvezhii Ugol’, [59.59, 74.71], 22.IX.2002, leg.A.B. Ryvkin (1 J cRyv). UඅඒൺඇඈඏඌKൺඃൺ Oൻඅൺඌඍ: Inzenskij Distr., Argash village, [53.96, 46.25]‚ birch forest, 9.VII.1995, leg.A.Isaev (2JJ NHMD). SLOVAKIA: Inovec, [48.77, 18.04], IX.1931, leg. Čepelák (1 J ZMHB). SWEDEN: Dagsås-trakten, Hall, [57.06, 12.48], 26.X.1953, leg. J.T. Skovgaard (2 JJ 3 ♀♀ NHMD); Södermanand, Ornö, [59.05, 18.41], 15.V.1946, leg. J.T. Skovgaard (1 J 1 ♀ NHMD); Stockholm, Djurgården 23.III.1945, leg. J.T. Skovgaard (1 ♀ NHMD); Vitamölle, [55.69, 14.21], 1.VIII.1926, leg.Stephensen (1J NHMD). Gඈඍඅൺඇൽ: Lau,[57.28, 18.61], 25.VI.1950 leg. J.T. Skovgaard (1 J NHMD). SWITZERLAND: Reussdelta, Seedorf, [46.89, 8.61], 435 m, VIII.1998, leg. Herger & Rezb. (1 J ZMHB). UNITED KINGDOM: EඇGඅൺඇൽ: Stanford le Hope, Essex,[51.51, 0.42], 7.IX.1969,leg.Wewalka (1♀ NMW). Sർඈඍඅൺඇൽ: Bridge of Orchy,[56.51, -4.76], 23.VII.1987, leg.Gillerfors (1♀ MZLU);S Braemer,[56.99,-3.39], 15.VII.1987, leg. Gillerfors (1 ♀ MZLU).

Redescription. Measurements JJ (n = 5): HW = 1.60– 1.78 (1.68); HL = 1.36–1.53 (1.44); HL/HW 0.83–0.88 (0.85); PW = 2.09–2.24 (2.20); PL = 1.93–2.11 (2.04); PL/PW 0.91–0.94 (0.93); EW = 2.00–2.18 (2.11); EL = 1.73–2.07 (1.92); EL/EW 0.87–0.95 (0.91); EL/PL 0.90– 1.02 (0.94); PW/HW 1.32–1.57 (1.47); forebody length 5.02–5.58 (5.40). ♀♀ (n = 5): HW = 1.60–1.69 (1.64); HL = 1.38–1.51 (1.45); HL/HW 0.86–0.93 (0.89); PW = 2.07–2.22 (2.13); PL = 1.91–2.09 (2.00); PL/PW 0.90–0.98 (0.94); EW = 2.04–2.18 (2.11); EL = 1.84–1.93 (1.91); EL/EW 0.88–0.95 (0.91); EL/PL 0.91–1.01 (0.96); PW/ HW 1.40–1.53 (1.47); forebody length 5.22–5.51 (5.35).

Medium sized, robust species; body dark brown to black ( Figs 8C,D ).

Head black, distinctly transverse, with eyes medium sized (EyL/TL = 1.67–2.13 (1.76)); microsculpture of transverse waves; no interocular punctures between anterior frontal punctures (cf. Fig. 6F ); antennae pale reddish with antennomeres 3–5 slightly darkened, all antennomeres elongate; palpi pale reddish.

Thorax: pronotum dark brown to black, slightly wider than long, 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 punctured and pubescent; elytra most often reddish brown rarely fully darkened, uniformly pubescent, slightly wider than long, roughly same length as pronotum; legs reddish brown with inner face of tibia and inner femur darkened.

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

Male. Aedeagus ( Fig. 13D ): paramere lanceolate with slight medial attenuation and extending into a slight expansion broadest below slightly asymmetric apex, reaching just beyond apex of median lobe, with sensory peg setae forming two long rows fusing together towards apex; median lobe broad with gentle constriction to a point at apex, on parameral side with two small teeth pointing slightly basad, positioned at level near basal level of peg setae band of paramere; C-shaped sclerite of internal sac with spine-like basal extension.

Differential diagnosis. Quedius molochinus is very similar to the other species with red elytra in the molochinus -group: Q. meridiocarpathicus , Q. balticus , and Q. vicinus . It can be distinguished from Q. balticus by the pale base of the basal three antennomeres. In most cases it has darker middle antennomeres and is generally slightly darker with abdomen black to dark brown (dark to light brown in Q. meridiocarpathicus and Q. vicinus ). For confident discrimination of these species genitalia should be examined: aedeagus of Q. molochinus has an extension of the C-sclerite in the internal sac. Quedius molochinus with black elytra can be confused with Q. subunicolor where their ranges overlap. Usually, the lighter colored appendages and the coarser microsculpture of Q. molochinus are sufficient to distinguish it from Q. subunicolor . Quedius molochinus can be confused with Q. picipes from the subgenus Raphirus , which may occur in similar environments. It can be readily distinguished from the latter by more parallel-sided abdomen and entire labrum. In North America the species can be confused with Q. laticollis , from which it can be distinguished by stouter antennae with slightly darker middle antennomeres. Also, in Q. molochinus elytra are more often brick red or almost black, whereas in Q. laticollis they are most often dark brown.

Synonymic notes. Quedius picipennis Stephens, 1832 from London, Norfolk, Suffolk, Devonshire, and Somerset and Raphirus picipennis Stephens, 1833 from London have recently been mixed up as the same species in both Hൾඋආൺඇ (2001) and Nൾඐඍඈඇ (2020). Based on the original descriptions, Raphirus picipennis clearly belongs in the Quedius subgenus Raphirus , whereas Quedius picipennis fits Q. molochinus . Original descriptions of Staphylinus laevicollis Runde, 1935 described from Halle in Central Germany and Staphylinus lapponicus Zetterstedt, 1838 described from Finland fit Q. molochinus , consistently with the currently accepted synonymy.

Bionomics. Quedius molochinus is found in meadows, heaths and grasslands, as well as in mainly coniferous forests. Most records are from under rocks or in moss. We have encountered it in flood debris near a river forested on either side in Czech Republic, in Sphagnum moss of a bog in Southern Sweden, and from pitfall traps set in a heathland in Denmark. Records from Russia (Sൾආංඈඇൾඇ-ĸ ඈඏ et al. 2015), Switzerland (UHඅංG et al. 2006) and the Netherlands (Cඎඉඉൾඇ et al. 2011) are from meadows. In North America it seems to be in more ruderal habitats as well, having been found in e.g., potato fields on Prince Edward Island (Mൺඃĸൺ 2007).

Distribution. Among the closest congeners, Quedius molochinus is a more northern species widely distributed from France and British Isles through Central and Northern Europe, across European Russia and western Siberia to Transbaicalia ( Fig. 20 ). In the southern areas of its distribution, it gradually moves to higher elevations, for example in the Alps extending into the Apennines. More thorough sampling in Russia may widen its distribution significantly. Quedius molochinus is introduced in North America, where it is currently found in three separate areas in Canada: around the Avalon Peninsula of Newfoundland (Sආൾ-ඍൺඇൺ 1981), in the Canadian Maritime Provinces (Nova Scotia, New Brunswick and Prince Edward Island, Mൺඃĸൺ 2007) and around Quebec City (Sආൾඍൺඇൺ 1981) ( 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. 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. 9. Habitus of Quedius s. str. A – Q. hispanicus Bernhauer, 1898; B – Q. hispanicus (pale sutures variant); C – Q. pallipes Lucas, 1846; D – Q. laticollis (Gravenhorst, 1802) (black elytra variant); E – Q. laticollis (red elytra variant); F – Q. strenuus Casey, 1915.

Fig. 8. Habitus of Quedius s. str. A – Q. vicinus (Ménétriés, 1832); B – Q. meridiocarpathicus Smetana, 1958; C – Q. molochinus (Gravenhorst, 1806) (red elytra variant); D – Q. molochinus (black elytra variant); E – Q. balticus Korge, 1960; F – Q. sundukovi Smetana, 2003.

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. 13. Aedeagus of Quedius s. str. A – Q. vicinus (Ménétriés, 1832); B – Q. sundukovi Smetana, 2003; C – Q. subunicolor Korge, 1961; D – Q. molochinus (Gravenhorst, 1806); E – Q. balticus Korge, 1960; F – Q. meridiocarpathicus Smetana, 1958. 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 in parameral view.

Fig. 20. Distribution of Quedius molochinus (Gravenhorst, 1806), Q. meridiocarpathicus Smetana, 1958, and Q. vicinus (Ménétriés, 1832) (top), Q. subunicolor Korge, 1961 (middle), and Q. sundukovi Smetana, 2003 (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).