Quedius unicolor Kiesenwetter, 1847

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 : 265-266

Quedius unicolor Kiesenwetter, 1847

( Figs 1 , 2G , 4 , 7E , 14A , 21 )

Quedius unicolor Kiesenwetter in Kංൾඌൾඇඐൾඍඍൾඋ & Mඟඋĸൾඅ (1847: 75). [Type locality: Riesengebirge]

References. Kංൾඌൾඇඐൾඍඍൾඋ (1848): 53 (characters); RൾൽඍൾඇൻൺർHൾඋ (1849): 825, (1857): 202, (1874): 199 (characters); Fൺංඋආൺංඋൾ & Lൺൻඈඎඅ-ൻජඇൾ (1856):536 (characters); Fൺඎඏൾඅ (1874):291 (characters);Mඎඅඌൺඇඍ & Rൾඒ (1876): 697 (characters); GൺඇGඅൻൺඎൾඋ (1895): 403 (characters); Pඈඋඍൺ (1907):132 (characters);Rൾංඍඍൾඋ (1909):112 (characters); Gඋංൽൾඅඅං (1924): 80 (characters); Pඈඋඍൾඏංඇ (1929): 341 (characters); WඳඌඍHඈൿൿ (1938): fig. 24 (characters); Sආൾඍൺඇൺ (1955): 143, (1960a): 259, (1964): 168, (1966): 331, (1993): 50 (distribution); (1958): 363 (characters and biology); (1962a): 134 (characters);Cඈංൿൿൺංඍ (1961): 49,(1978):193 (characters);LඈHඌൾ (1964):211 (characters); Hൺඏൾඅĸൺ (1964):93 (distribution); Hඈඋංඈඇ (1965): 274 (distribution); Bඈඋൽඈඇං (1976a): 94 (characters); ZൾඋർHൾ (1977): 9 (distribution), TඬඍH (1984): 119 (characters); LඎർHඍ (1987):109 (distribution);Jൺඇගĸ (1992):91 (distribution);Hൾඋආൺඇ (2001): 50 (application to ICZN), Tඋඈඇඊඎൾඍ (2006): 102 (distribution); Mൺඓඎඋ et al. (2007): 30 (distribution); MൺඍൾඅൾඌHĸඈ (2007): 184 (distribution); SർHൺඍඓ (2008): 388 (distribution); Zൺඇൾඍඍං (2015): 96 (distribution).

Material examined. AUSTRIA: Golling, [47.59, 13.16], leg. Skalitzky (1 ♀ NMW); Kerschbaumeralm, [46.75, 12.76], 24.VII.-7.VIII.1948, leg. F. Schubert (1 ♀ NMW); Koralpe, [46.88, 14.99], leg. A. Otto (3 JJ NMW); Lünersee, [47.05, 9.74], 14.VI.-3.VII.1952, leg. F. Schubert (1 ♀ NMW); Stuhleck, [47.56, 15.78], leg. Breit (1 ♀ NHMD); Stuhleck, Steiermark, [47.56, 15.78], leg.Wingelmüller (1 ♀ NMW); Tirol, [47.22, 11.51] (1 SDEI); Wechsel,[47.52, 15.91], leg. Scheerpeltz (3♀♀ NMW); Wolayersee, [46.61, 12.86], 3.-11.VIII.1949, leg. F. Schubert (2 JJ 1 ♀ NHMD); Zillertal nr. Mayrhofen, [47.16, 11.87], 4.VIII.1914, leg. H. Wagner (1 SDEI); Wolayersee, [46.61, 12.86], 3.11. VIII.1949, leg. F. Schubert (2 ♀♀ NMW). CZECH REPUBLIC: Krkonoše [50.71, 15.66], leg. Obenberger (1 J 2 ♀♀ NHMD); Spindelmühle [Špindlerův Mlýn], [50.73, 15.61], leg. Skalitzky (1 NHMD). GERMANY: Harz, Brocken, [51.79, 10.61], 9.V.1922, leg. Uhmann (1 J SDEI); Hercyn mont.[Harz Mountains], [51.74, 10.63], 14.VI.1914, leg. Beckenkamp (3 ♀♀ SDEI); Thüringer Wald Oberhof, [50.71, 10.73], 8.IX.1951, leg. W. Liebmann (1 ♀ SDEI). ITALY: Gressory [Gressoney] la Trinite, [45.87, 7.82], 15.VII.1901, leg. Künnemann (1 SDEI); Malga Preghena Alta, Trento, 46.4148, 10.8922, 2100 m, under stones along stream with moss, 26.VIII.2019, leg.A.Zanetti ( NHMD);Valle Brembana, Strada di Porcile, [46.06, 9.73], 1800-2000 m, 26.VI.1961, leg. V. Rosa (1 J NMW). PO-LAND: Silesia [Bad] Flinsberg [Świeradów-Zdrój], [50.92, 15.31], leg. Kraatz (1 SDEI). SPAIN: Cantabria, Alto Campoo, W Reinosa, [43.03, -4.37], 4.VI.1991 2000m leg. Zerche (1 SDEI). SLOVAKIA: Ružomberok, [49.04, 19.23], VII.1952, leg. Přívora (1 J NHMD); High Tatras, Mengušovská dolina, Vysoke Tatry, [49.16, 20.07], 1200 m, 17.V.1991, leg. J. Frisch (1 J ZMHB); Tatra, Kráľova hoľa, [48.88, 20.13], 1300- 1600 m, 12.VIII.1981, leg. Hieke (1 J ZMHB). SWITZERLAND: Aarau, [47.38, 8.05], leg. Stierlin (1 SDEI).

Redescription. Measurements JJ (n = 5): HW = 1.47– 1.76 (1.58); HL = 1.31–1.44 (1.36); HL/HW 0.82–0.91 (0.86); PW = 1.96–2.33 (2.16); PL = 1.78–2.02 (1.91); PL/PW 0.86–0.92 (0.89); EW = 2.13–2.47 (2.32); EL = 2.18–2.47 (2.32); EL/EW 0.95–1.05 (1.00); EL/PL 1.15– 1.26 (1.22); PW/HW 1.47–1.65 (1.58); forebody length 5.36–5.93 (5.58). ♀♀ (n = 5): HW = 1.58–1.67 (1.61); HL = 1.31–1.42 (1.37); HL/HW 0.79–0.89 (0.85); PW = 2.00–2.18 (2.08); PL = 1.82–1.89 (1.87); PL/PW 0.87–0.92 (0.90); EW = 2.22–2.40 (2.31); EL = 2.20–2.40 (2.31); EL/EW 0.96–1.01 (0.99); EL/PL 1.19–1.27 (1.23); PW/ HW 1.45–1.57 (1.52); forebody length 5.36–5.69 (5.52).

Medium sized species; body black ( Fig. 7E ).

Head black, distinctly transverse, with eyes rather small (EyL/TL = 1.59–2.00 (1.82)); microsculpture of fine mesh, especially on frons, appears dull; no interocular punctures between anterior frontal punctures (cf. Fig. 6F ); antennae dark brown to black, all antennomeres slightly elongate; palpi dark brown to black.

Thorax: pronotum black, slightly wider than long, clearly wider than head, with microsculpture of fine transverse waves; three punctures in dorsal row and one to two in sublateral row with its posteriormost puncture reaching just beyond first puncture of dorsal row; scutellum punctured and pubescent; elytra black, uniformly pubescent, as long as wide, clearly longer than prontoum; legs dark, outer face of tibia and tarsi often paler.

Abdomen black, tergites uniformly punctured, without clear iridescence.

Male. Aedeagus ( Fig. 14A ): paramere lanceolate with slight medial attenuation and extending into a slight expansion broadest below apex, apex extending into small blunt knob away from the median lobe, apex slightly asymmetric, reaching just beyond apex of median lobe, with sensory peg setae forming two rows fusing together towards apex; median lobe broad with gentle constriction to a point at apex, on parameral side with two small ridges continuing basad, positioned at level near basal level of peg setae band of the paramere; internal sac without C-sclerite.

Differential diagnosis. Quedius unicolor is very similar to other species with dark elytra from the molochinus- group – Q. subunicolor , Q. sundukovi and the dark form of Q. molochinus . It can be distinguished from Q. subunicolor by the dull frons due to meshed microsculpture (in Q. subunicolor the microsculpture consists of fine transverse waves or only slightly meshed). Also, the genitalia and the distributions are clearly different in the two species. It usually differs from the dark form of Q. molochinus in the darker appendages and the finer microsculpture in addition to a clearly different aedagus, which is without a C-sclerite in Q. unicolor , but with such in Q. molochinus . Quedius unicolor has non-overlapping distribution with Quedius sundukovi and can be clearly distinguished from the latter species by the presence of palisade fringe on tergite VII.

Quedius unicolor has historically been confused with Q. subunicolor . For details see comments about the latter species.

Comments. Quedius unicolor was originally described from Riesengebirge [Krkonoše] (Kංൾඌൾඇඐൾඍඍൾඋ & Mඟඋĸൾඅ 1847), a mountain range at the border between Poland and Czech Republic. It was later discovered in other mountainous area, including the Alps, the Carpathians, the Pyrenees, Picos de Europa, the Harz Mountains, and possibly extending into Sierra Nevada and even central Anatolia, although the latter records are not confirmed here (Cඈංൿൿൺංඍ 1961, AൻൺർංGංඅ et. al 2009; Fig. 21 ). At these sites it seems to be restricted to a very specific habitat type, with most specimens being found under stones and in moss near mountain streams at higher elevations. COI barcodes of specimens from several sites, including the Austrian and Italian Alps, the Carpathians of Slovakia, and Picos de Europa in Spain displayed high genetic variability ( Fig. 4 ), with the sites being grouped into three OTUs (the Carpathians, Picos de Europa, and the Alps) ranging from 3.5 % to 4.5 % divergence, with a 0.2% divergence between the Austrian and Italian Alps ( Table 2 ). With such a high genetic divergence in the barcode region, we carefully checked for morphological differences. However, we found no obvious differences corresponding to the molecular clusters. Since the wings seem to be fully developed in this species, these genetic differences may be attributed to isolation by distance, the very specific habitat requirements of the species and its close association to a now fragmented habitat, which would limit gene flow among populations.

Bionomics. Quedius unicolor seems to be very habitat selective and is only found in alpine sites in sphagnum moss and under rocks near bogs, swamps and creeks ( Fig. 2G ). It is found at altitudes between 900 and 2300 m depending on the latitude of the locality, with higher elevations at more southern sites.

Distribution. Quedius unicolor is restricted to the montane and subalpine zone of Central and southern Europe ( Fig. 21 ). Specifically, it is known from sites in the Carpathians, the Alps, the Pyrenees, Picos de Europa, the Sudetes, the Harz and the Ore mountain ranges. Due to its very specific habitat requirement, it may be under-collected and may actually be present in mountains further south. Cඈංൿൿൺංඍ (1961) reported it from Sierra Nevada in Southern Spain, but this record may be in fact a misidentified Q. hispanicus . Another doubtful record of Q. unicolor comes from Koçere Stream in the Kazdağları Mountain of the North-western Anatolia by AൻൺർංGංඅ et. al (2009), which is a location remote from the main range of this species.

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. 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. 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. 14. Aedeagus of Quedius s. str. A – Q. unicolor Kiesenwetter, 1847; B – Q. levasseuri Coiffait, 1964 (syn. of Q. pallipes Lucas, 1846); C – Q. hispanicus Bernhauer, 1898; D – Q. pallipes Lucas, 1846; E – Q. laticollis (Gravenhorst, 1802); F – Q. strenuus Casey, 1915. 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. 21. Distribution of Quedius unicolor Kiesenwetter, 1847 (top),.Q. labradorensis Smetana, 1965 (middle), Quedius molochinoides-group (bottom). Barcoded specimens are denoted with +.

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).