Quedionuchus glaber

Brunke, A. J., Salnitska, M., Hansen, A. K., Zmudzinska, A., Smetana, A., Buffam, J. & Solodovnikov, A., 2020, Are subcortical rove beetles truly Holarctic? An integrative taxonomic revision of north temperate Quedionuchus (Coleoptera: Staphylinidae: Staphylininae), Organisms Diversity & Evolution (New York, N. Y.) 20 (1), pp. 77-116 : 96-100

Quedionuchus glaber : Schülke and Smetana 2015 (as syn. of Q. plagiatus )

Type locality: ‘Denmark’ (unknown locality), most likely what is currently Norway.

Type material. Type material was not examined for this common Palaearctic species as only one species of Quedionuchus occurs in Europe. O. Müller (1776) described species from both Norway and Denmark , which were at the time unified under a broader Danish border. As there are no historical or current records of the genus in Denmark, the type was most likely collected from an unknown locality in Norway .

Quedionuchus planatus Sharp, 1888 syn nov.

Type material. Lectotype (male, BMNH) (here designated): Quedionuchus planatus Type D.S./Nikko, 12.8.1881, Lewis [handwriting on card]/Type [circle label with orange border]/ Japan, G. Lewis [printed label]/Sharp Coll. 1905-313 [printed label]/AJB0001085 [printed label]/ Quedionuchus planatus LECTOTYPE des. Brunke 2019 [printed red label]/ Quedionuchus glaber O. Müller det Brunke 2019.

Paralectotype (female, BMNH): same as above except: no date and no circular type label. AJB0001086/ PARALECTOTYPE det. Brunke 2019 [printed yellow label]

Sharp (1888) described this species from four specimens collected by G. Lewis in Nikko, Japan. Two specimens (one male, one female) were available for study and are morphologically identical to the specimens of Q. glaber from the East Palaearctic mainland. Sharp (1888) remarked that specimens of his Q. planatus were similar to but smaller than Q. glaber (as ‘European Q. plagiatus ’ in the very broad sense). While most Japanese specimens at hand are indeed smaller than the available specimens of Q. glaber , several are as large as some of the largest Q. glaber . Due to the possibility of an entirely dark morph of Q. glaber in the East Palaearctic, and therefore confusion with sympatric Q. samuraicus , the male specimen is designated as a lectotype.

Other material

West Palaearctic

ALBANIA: Pashtrik (Alban. Exped.), 29.v.–4.vi.1918 (1, FMNH) .

AUSTRIA: Carinthia: Lassen (1, FMNH); Lienz: Ausservillgraten , Mooshofalm , 1535 m, 18.VII.2013, Koehler, F. (barcode voucher GBOL_Col_FK_7060) (1 SNSB) .

CZECH REPUBLIC: Bohemia: Prachatice (1, FMNH) ; Moravia: Praded (Altvater), J. Spurny (1, FMNH) ; Praděd, Th. V. Wanka (1, FMNH, 1, CNC); Beskydy Mts. , P. Nohel, 11.VII.1962 (1, CNC) ; Travny, Beskid Mts. , P. Nohel, 28.VIII.1961 (1, CNC) .

ESTONIA: Ida-Viru: 'Shmecke' [part of Narva-Joesuu], Gersenshtein, 1890 (1, ZIN).

FINLAND: Lapland: Rovaniemi, Pisavaara , 66.309 25.127, 200 m, M. Pentinsaari, 27.VII.2010 (1, ZMOU) .

FRANCE: Haute Loire: Pradelles , 1000–1100 m, under pine bark, M.I. Russell, 25.VIII.1977 (1, BMNH) .

GERMANY: Bavaria: Bayerischer Wald, Freyung- Grafenau, Spiegelau , Schwarzachstrasse , 48.946 13.362, 800 m, Koehler, F. & M. Koehler, 9.VII.2011 (1, SNSB) ; Thuringia: Thueringer Wald, Gehlberg, Grosser Beerberg , 50.656 10.741, 960 m, intercept trap, conifer forest, 5.VII.2014 .

ITALY: Aosta: La Thuile, G.C.C. (1, BMNH).

LITHUANIA: Slavinskas, G, 9.VII.1986 (1, SEMC) .

NORWAY: Oppland: Stueflaaten [=Lesjaskog], G.C.C. (1, BMNH).

POLAND: Eastern Carpathians : Zatwarnica trail, Wetlinska-Polonia, Bieszczady, B. Malkin & E. Wilczynski, 18.VIII.1974 (1, FMNH) ; Zatwarnica, Bieszczady Mts. , B. & H. Malkin, 12.VII.1987 (1, FMNH) ; no locality, 15.VIII.1928 (1, SEMC) .

RUSSIA: Briansk Oblast: Briansk, V.N. Stark, 6.V.1927 (3, ZIN) ; Khanty-Mansi Autonomous District: Berezov [Beryozovo], Tobolsk. G., 63.94 65.05, V. Yu. Fridolin, 10.VI.1925 (1, ZIN) ; Beryozovo, downstream of the Ob River , Maksimov, 63.93 65.06, 20.VI.1928 (1, ZIN) ; Manya River basin, ' B. Ural' [Big Ural?], 64.34 61.01, Lyapin, Flerov, 15.– 23.VII.1923 (1, ZIN) ; Komi Republic: Letka village , 59.59 49.42, on mushrooms and under bark of Betula sp. , A.L. Lobanov, 14.V.1976 (1, ZIN) ; Leningrad Oblast: Kirillovskoye , 60.454 29.330, A.G. Kirejtshuk, 20.VII.1981 (2, ZIN) ; Komarovo village , 60.19 29.79, O.L. Kryzhanovsky, VII.1971 (1, ZIN) ; near station Vyritsa , 59.41 30.35, G. Zinovjev, 6.IX.1956 (3, ZIN) ; Petrograd [Saint-Petersburg], 59.90 30.35, 7.V.1922 (1, ZIN) ; Beloostrov village, Karelian Isthmus , 60.15 30.03, V.N. Prasolov, 2.VI.1981 (1, ZIN) ; Moscow Oblast: near Zvenigorod , 55.73 36.85, O.L. Kryzhanovsky, IX.1972 (1, ZIN) ; Balashikha district, Pehra- Pokrovsky vill., 55.84 37.94, V.V. Zolotikhin, 10.VI.1969 (1, ZIN) ; Bronnitsy village , 55.41 38.28, under Pinus sp. bark, A.P. Rasnitsyn, 22.IX.1957 (1, ZIN) ; Prioksko - Terrasny State NB, 54.88 37.67, in mushrooms, Mulgaupt, 16.VIII.1957 (1, ZIN) ; Sergiev Posad [Zagorsky] district, Abramtsevo village , 56.23 37.97, V.V. Stark (1, ZIN) ; Murmansk Oblast: basin of Vydyavr Lake, Khibin. c. Kolsk. [Khibiny Mts., Kola Peninsula], 67.65 33.67, N.M. Cheburova, 10–11.IX.1930 (4, ZIN) ; Imandra , north Karelia, 67.84 33.26, Stepanov, 9.VIII.1928 (2, ZIN) ; Varnichniy stream, Kola Bay , 68.96 33.08, Kapustin, 12.V.1924 ; Perm Oblast: near city Kungur , 57.36 57.17, forestry station "Preduralye", G. Zinovjev, 16.VII.1955 (1, ZIN) ; near city Gubakha , 58.84 57.62, G. Zinovjev, 7.VI.1956 (1, ZIN) ; Republic of Mordovia: Bolshebereznikovsky district, Simkino village , 54.26 46.17, Klimenko, 19.VI.1987 (1, ZIN) ; Vladimir Oblast: 1909-V / Vladimir gub., 56.16 40.4 (1, ZIN) .

SCOTLAND: Aviemore , 57.19 -3.82, VI.1938 (1, CNC) .

SLOVENIA: Triglav Nat. Park nr. Trenta, 46.37 13.74, A. Solodovnikov, 2.VI.2016 (1, NHMD)

SLOVAKIA: Low Tatra Mountains nr. Pavčina Lehota, 49.03 19.59, A.K.Hansen, 22.VII.2018 (5, 2 dark form, NHMD); Pol’ ana, 48.63 19.48, J. Háček, 3.VIII.1986 (1, NHMD)

SWEDEN: Lapland: Jokkmokk, 66.61 19.82, T. Palm, 3– 11.VII.1948 (1, CNC); Örebro: Kopparberg, F.W.C. (1, BMNH) .

SWITZERLAND: Aigle: Solalex , 46.28 7.13, under bark, pine logs, 1400–1500 m, P.M. Hammond, 1.VI.1975 (1, BMNH) ; Sierre: Montana, Cameron, 20.VIII.1925 (1, BMNH) ; Vaud: Col du Marchairuz , 46.55 6.25, 1450 m, B. & H. Malkin, 13.VI.1976 (1, FMNH) .

UKRAINE: Mt. Menchul nr. Rakhov town, 48.02 24.25, A. Gontarenko, 21.V.2002 (1, NHMD); Svidovec, 48.26 24.13, Dr. Klicka, VII.1930 (1, NHMD)

East Palaearctic

CHINA: Gansu: [likely Lanzhou, along Yellow River ( Royal Geographical Society 1887)], G. Potanin, 1885 (1, CNC) .

JAPAN: Hokkaido: Sapporo, Usubetsu, Muine Bridge , 42.90 141.12, 600 m, under bark of Abies, A. Smetana , 7.VIII.1991; Honshu: Yamanashi Pref., Hacchoudaira, Sudama-cho, 36.76 139.65, T. Ueno, 22.VII.2001 (2, cSch); Yamanashi Pref., Kitazawatoge Pass, Ashiyasu-mura, 35.74 138.138.21, T. Ueno, 2001 (1, Sch) .

KAZAKHSTAN: East Kazakhstan Region: Altai, Bukhtarma, Archaty, 49.30 86.53, V.A. Kastcheev, 25.VII.1989 (2, ZIN) ; Altai, Ul'ba riv., Cheremshanka River , 50.26 83.05, V.A. Kascheev, 5.VII.1988 (1, ZIN) ; Altai, Rakhmanovskie Kluchi, Barel River , 49.52 86.50, V.A. Kascheev, 11.VI.1988 (11, ZIN) .

RUSSIA: Altai Territory/ Republic of Altai: Altai / Staudinger. /813 (1, ZIN) ; Altai Territory: Soloneshensky District, Butachikha Mt. between Elinovo and Telezhino villages, 51.47 84.36, 1700–2000 m, A.G. Kirejtshuk, 13.VI.1988 (2, ZIN) ; Amur Oblast: Zeya Nature Reserve , 'kordon' [hut] 52 km, 54.08 126.88, V.V. Belov, 2.VII.1978 (1, ZIN) ; ' Erahta' [Khingan Nature Reserve, near Erahta cordon, forest № 1], 49.155 130.720, Abies sp. with bark beetles, between brown phloem and xylem, V.A. Potockaya, 21.V.1976 (1, ZMMU) ; Buryatia Republic: Barguzinsky Nature Reserve, upper reaches of the river Davsha , 54.35 109.50, V.G. Shilenkov, 12–14.VII.1989 (1, ZIN) ; East Siberia, Baikalsky Nature Reserve, middle flow of Pereemnaya River , "Shum" winternut, 51.49, 105.23, Y. Sundakov, 25–28.VII.2009 (1, ZIN) ; East Siberia, Khamar-Daban Mts., Snezhnaya River valley, 51.37 104.65, under bark of old poplar log, A.V. Shavrin, 19.V.1997 (1, ZIN) ; East Siberia, Khamar-Daban Mts. , upper Slyudyanka River , 51.61 103.64, A.V. Shavrin, 15–16.VI.2008 (2, ZIN) ; Irkutsk Oblast: Khamar-Daban Mts., valley of Podkomarnaya River, 51.62 103.65, A.V. Shavrin, 15.VII.2008; Nizhneudinsky District , 7 km NW Trud village, 56.01 98.02, A.V. Shavrin, 28.VI.1999 (1, ZIN) ; North part of Baikal Range, upstream of Kunerma River , 15 km ESE Kunerma station, 55.65 108.60, 1200 m, A.Yu. et. R.Yu. Dudko, and D.E. Lomakin, 9.VIII.1995 (1, ZIN) ; ' Sludanka River , railway station Sludanka, ca. 150 km from Irkutsk, 51.63 103.67, S. Rodionoff, 10.VII.1914 –1915 (1, ZIN) ; Khabarovsk Territory: Verkhnebureinsky District, the Bureinsky Nature Reserve , basin of Right Bureya River , upper from Medvezhye Zimovye Hut , 52.11 134.31, 800 m, on Leccinum mushroom in floodplain forest, A.B. Ryvkin, 27.VII.2007 (1, cRyv); Badshalski-Gebrige, Khabarski-Krai, Mundg. des Omot- Makit, 50.58 134.34, 850 m, Y. Sundukov, 12.VII.1997 (2, cSch); Budshalski-Gebrige , Khabaraski Krai, Omot-See, 50.54 134.24, 1165 m, Y. Sundukov, 13.VII.1997 (6, cSch); Primorsk Territory: Gorelaja Sopka Mt. , Sikhote-Alin Mts ., Primorsky Krai, Gorelaja Sobka, 43.51 134.10, 1300–1400 m, Y. Sundukov, 17.VI.1999 (1, cSch); Podnebesnyi pass, 70 km NW Plastun, 45.13 135.71, 600–800 m, A. Plutenko, 18– 28.VII.1996 (1, cSch); Sikhote-Alin Nature Reserve , Jasnaja-Munding, 45.24 136.51, J. Sundukov, 26.VI– 4.VII.1998 (1, cSch); Suputinski zapov. [Ussurisky Nature Reserve], 43.68 132.55, O.L. Kryzhanovsky, 2.IX.1969 (1, ZIN) ; River Valley of Jelanovskovo , Partisanski Krai, Lazovsky district, 43.24 133, J. Sundukov, 21.VI.1994 (1, AnPut); Sikhote-Alinsky preserve, 4 km above from cordon Maisa , 45.32 136.25, A.G. Kirejtshuk, 24.V.2001 (2, ZIN) ; North Sikhote Alin (1, ZIN) ; Sakha Republic: East Yakutiya, Suntar-Khayata Mts., Tyry valley, Nezhdanskoe river, Kidyrki River , near Khalya River , 800–950, S. Alexeev, 14–18.VIII.1991 (1, ZIN) ; Tuva Republic: Buren River valley, Khorumnug-Taiga , 51.42 95.52, 1000 m, 15.VI.1999 (4, cHAY); Zabaikalsky Territory: Stanovoe nagorje, upstream of Chara River , lake Bolshoe Leprindo , 56.61 117.48, 1000 m, A.Yu. et. R.Yu. Dudko, and D.E .

Lomakin, 23.VII.1995 (1, ZIN) .

Diagnosis. Quedionuchus glaber is the only species of the Glaber group in the Palaearctic region as far as known. The morphological variation of male and female genitalia, and external features overlaps with that of Nearctic Q. longipennis but these allopatric species can be separated by their DNA barcodes.

Redescription. Measurements ♂ (n = 5): HW/HL 1.31–1.44; PW/PL 1.06–1.20; EW/EL 1.07–1.17; ESut/PL 0.77–0.84; PW/HW 1.09–1.16; forebody length 3.4– 4.4 mm.

Measurements ♀ (n = 5): HW/HL 1.31–1.39; PW/PL 1.09–1.17; EW/EL 1.08–1.12; ESut/PL 0.75–0.83; PW/HW 1.10–1.18; forebody length 3.5–4.2 mm.

Head, pronotum and abdomen brownish, pronotum sometimes slightly paler, reddish-brown, scutellum dark, elytra either entirely reddish, entirely black, or yellowish brown with darker spot on disc, spot often occupying most of disc with only suture, apical margin and humeral area paler, legs reddish brown to reddish-yellow, with tibiae darkened on inner face, maxillary and labial palpi entirely light reddish brown, antennomeres 1–3 light reddish, often darkened apically, 4– 11 reddish to brownish, apices of abdominal segments varying to paler to markedly paler (usually corresponding with spotted elytra morphotype).

Head distinctly transverse, males with the most extreme values, disc with moderately coarse microsculpture of transverse waves, eyes large, convex, about 1.5–1.75 times the length of temples; antennomeres 1–3 elongate, 4 weakly transverse to slightly elongate, 5–10 increasingly transverse; pronotum slightly to moderately transverse, disc with moderately coarse microsculpture of transverse waves, waves whorled at middle of basal third, between the dorsal rows, with usually 3 punctures in the dorsal row, frequently one or two punctures missing at one side; punctation of abdominal tergites moderately dense and moderately to strongly asperate, most punctures separated by about 1.0–1.5 puncture diameter, punctation most dense and asperate in spotted elytra morphotype, with median impunctate area usually only first visible abdominal tergite but occasionally on second, abdominal segments with fine, dense microsculpture of transverse waves, separated by approximately their widths.

Male. Sternite VII varying from not appreciably emarginate to slightly and shallowly emarginate, sternite VIII with distinct, moderately deep and rounded emargination, tergite X elongate triangular, evenly converging to apex, sternite IX moderately elongate, with broadly rounded apex varying from entire to slightly emarginate. Median lobe in ventral view varying from moderately short to elongate and acute apex, evenly converging to apex, median lobe in lateral view always with distinct and moderately large tooth forming triangular apical part ( Fig. 6a–b ), paramere elongate ( Fig. 6d ) but variable, subparallel-sided to broad basally and gradually converging to apex, with peg setae in two short longitudinal fields of 2–3 peg setae wide. Hind femora with spines twice as long as basalmost spines, posterior edge usually sinuate.

Female. Tergite X with thin, elongate central lobe (as in Fig. 7c ). Female hind femora with spines in apical half not twice as long as basalmost spines, posterior edge not sinuate.

Distribution. Broadly distributed in the Palaearctic ( Fig. 8a ), including Europe, non-European Russia, Kazakhstan, far northern China (Gansu) and Japan. It has a boreomontane distribution in Europe. Records are known from northern Europe, east through the forest belt of Siberia, to the Ural Mountains in the north, and in the south, concentrated in the mountains of Central and southern Europe as well as southern and eastern Siberia, Altai and Japan. The species is newly reported from Lithuania. It may occur in North and South Korea but has not yet been recorded. The single record from China is based on an old specimen (1885, CNC) collected by G. Potanin during an 1884-1887 expedition and likely comes from near the city of Lanzhou ( Royal Geographical Society 1887). This record may indicate a continuous distribution from Japan and the Russian Far East to north-westernmost China and Mongolia. Its presence in Mongolia, reported by Smetana (1967) and Schülke and Smetana (2015) could not be confirmed here.

Bionomics. This species is collected in forests, primarily under the bark of dead trees including Pinus and Betula in the Western Palaearctic, and Abies and Populus in the Eastern Palaearctic. Larvae and pupae are also found under bark. Unlike Nearctic Q. longipennis , Q. glaber is not known from rich hardwood forests. This species was found to prefer shaded to sun-exposed logs ( Johansson et al. 2007) and mainly predate on bark beetle larvae (including Ips typographus (L.)) ( Weslien 1992). Several specimens were collected on mushrooms (including Leccinum ) but these may be incidental findings. One specimen in Amur was collected in association with bark beetles (Scotylinae?). Specimens have been collected in floodplain forests to montane spruce forests at an elevational range of 170–2000 m.

Comments. Staphylinus glaber O. Müller, 1776 is the oldest available name for the Palaearctic species of the Glaber lineage. The ICZN (2004) ruled that Staphylinus plagiatus Mannerheim, 1843 should have precedence over both Staphylinus glaber O. Müller, 1776 and Staphylinus flavopterus Geoffrey, 1785 but the first two taxa are here no longer considered conspecific. Coiffait (1978) recognized correctly that Q. glaber (as Quedius laevigatus ) was colour polymorphic and treated Q. ater Sahlberg, 1880 and Q. resplendens Thomson, 1860 as all-dark and spotted colour varieties, respectively, of the former. The all-dark form of Q. glaber is uncommon, apparently restricted to the West Palaearctic and only a few specimens from the Tatra Mountains in Slovakia were available for study.

Figures

Fig. 6 Quedionuchus glaber (O. Müller) (a, West Palaearctic, b, d, East Palaearctic), Q. longipennis (Mannerheim), typical shape (c), Q. plagiatus (Mannerheim) (e–g), Q. gilaensis Brunke (i–k) and Q. armipes Sharp (l–o). Aedeagus in ventral view (l), median lobe in

Fig. 7 Female tergite X of: Quedionuchus samuraicus (Bernhauer and Schubert) (a); Q. reitterianus (Bernhauer) (b); Q. longipennis (Mannerheim) (c); Q. plagiatus (Mannerheim) (d); Q. gilaensis Brunke (E); Q. armipes Sharp (F). Scale bars = 0.5 mm

Fig. 8 Distribution of: Quedionuchus glaber (O. Müller) (black circles), Q. caucasicus Brunke (blue square), Q. reitterianus (Bernhauer) (yellow triangle) and Q. yunnanensis Brunke (pink star) (a); Q. deceptor Brunke (black squares) and Q. samuraicus (Bernhauer and Schubert) (pink circles) (b)