On the taxonomy and zoogeography of some West Palaearctic Quedius species, with a focus on the East Mediterranean and the species allied to Quedius umbrinus and Q. nivicola (Coleoptera: Staphylinidae: Staphylininae) Author Assing, Volker text Linzer biologische Beiträge 2018 2018-07-27 50 1 149 182 journal article 21205 10.5281/zenodo.3985368 efec1c9a-9bf5-4599-bfd4-b932b9d34026 0253-116X 3985368 Quedius ( Raphirus ) umbrinus ERICHSON , 1839 Quedius umbrinus ERICHSON, 1839: 491 f. [ type locality: "Neustadt-Eberswalde"]. Raphirus umbrinus var. maritimus J. SAHLBERG, 1876: 29 [first synonymy: Fauvel (1900) ]. Quedius ( Sauridus ) cyanescens MULSANT & REY, 1876: 727 ff.; nov.syn. Quedius umbrinus var. tetrastigma LEINBERG, 1900: 79 . Quedius dubius var. umbripennis BERNHAUER & SCHUBERT, 1916: 79; see comment below. Quedius bulgaricus SCHEERPELTZ, 1937: 219 ff.; nov.syn. Quedius cyprusensis LAST, 1955: 251 f.; nov.syn. Quedius freyi SCHEERPELTZ, 1956: 1102 ff.; nov.syn. Quedius pseudoumbrinus LOHSE, 1958: 59 ; synonymy by POPE (1977) and ASSING (1999) . Quedius maronitus COIFFAIT, 1963: 410 ; nov.syn. Quedius gueorguievi COIFFAIT, 1967: 399 f.; nov.syn. T y p e m a t e r i a l e x a m i n e d: Q. cyanescens : Lectotype ♂ , present designation [dissected prior to present study]: " ♂ 14 / Lectotype / D. Drugmand Rev. 1995 Quedius ( Sauridus ) cyanescens Muls. Rey / Lectotypus ♂ Quedius cyanescens Mulsant & Rey, desig. V. Assing 2017 / Quedius umbrinus Erichson , det. V. Assing 2017" ( MHNL ). Paralectotypes : 1♂ : "1 / Paralectotype / D. Drugmand Rev. 1995 Quedius ( Sauridus ) cyanescens Muls. Rey / Paralectotypus ♂ Quedius cyanescens Mulsant & Rey, desig. V. Assing 2017 / Quedius umbrinus Erichson , det. V. Assing 2017" ( MHNL ) ; 1♀ : " ♀ / Paralectotype / D. Drugmand Rev. 1995 Quedius ( Sauridus ) cyanescens Muls. Rey / Paralectotypus ♀ Quedius cyanescens Mulsant & Rey, desig. V. Assing 2017 / Quedius umbrinus Erichson , det. V. Assing 2017" ( MHNL ) . Q. bulgaricus : Lectotype ♂ , present designation [dissected prior to present study]: " ♂ / Rosalitopolje, Schipka-Balkan, ca. 1900 m / ex coll. Scheerpeltz / Typus Quedius bulgaricus O. Scheerpeltz / Lectotypus ♂ Quedius bulgaricus Scheerpeltz, desig. V. Assing 2017 / Quedius umbrinus Erichson , det. V. Assing 2017" ( NHMW ) . Paralectotypes : 1♀ : same labels as lectotype ( NHMW ) ; 1♂ : same data as lectotype, but "Cotypus" ( NHMW ) ; 1♀ : " ♀ / Jumruktschal, Schipka-Balkan, 1400-1800 m / ex coll. Scheerpeltz / Cotypus Quedius bulgaricus O. Scheerpeltz / Quedius umbrinus Erichson , det. V. Assing 2017" ( NHMW ) . A d d i t i o n a l m a t e r i a l e x a m i n e d: Numerous specimens from Germany , Poland , Italy , Austria , Slovakia , Romania , Slovenia , Serbia , Albania , Macedonia , Bulgaria , Greece , Turkey , Cyprus , Lebanon , Russia (West Caucasus), Georgia , and Armenia . C o m m e n t: Quedius umbrinus was described based on an unspecified number of syntypes from "Neustadt-Eberswalde", Germany (ERICHSON 1839). The original description of Q. cyanescens is based on an unspecified number of syntypes from "les environs d'Hyères" (MULSANT & REY 1876). The three syntypes in the Rey collection had been examined, partly dissected, and labelled as lectotype and paralectotypes , respectively, by Didier Drugmand. A designation, however, was never published. In order to establish a formal designation, the male in better condition is designated as the lectotype . The aedeagus of the lectotype is within the range of intraspecific variation of that of Q. umbrinus . Hence the synonymy proposed above. The first and second editions of the Palaearctic Catalogue ( SMETANA 2004 , SCHÜLKE & SMETANA 2015) list Quedius umbripennis ROUBAL, 1913 as a junior synonym of Q. umbrinus . ROUBAL (1913) described Q. umbripennis as an aberration of Quedius dubius from the Caucasus, i.e., as an infrasubspecific and consequently unavailable name (see also HERMAN 2001 ). In treating the name as a variety of Q. dubius ( HEER, 1839 ), BERNHAUER & SCHUBERT (1916) made the name available and, according to the Code, they must be considered the authors of the taxon. In more recent articles ( COIFFAIT 1963 , 1968, KORGE 1964 , HERMAN 2001 ) it is treated as a distinct species. Based on the illustrations of the aedeagus provided by COIFFAIT (1978a) , however, it was correctly placed in synonymy with Q. umbrinus . LAST (1955) described Q. cyprusensis based on a male holotype and an unspecified number of paratypes from "Mount Troodus", Cyprus . Quedius freyi , which was described based on 16 type specimens from "Pedhoulas" only one year later ( SCHEERPELTZ 1956 ) was subsequently synonymized with Q. cyprusensis by COIFFAIT (1978a) . At present, Q. cyprusensis is known from Cyprus , Lebanon , and Turkey (ANLAŞ & ROSE 2009, ASSING & WUNDERLE 2001, BORDONI 1984 , COIFFAIT 1963 , 1970 , 1978a ). According to LAST (1955) and SCHEERPELTZ (1956) , the species is distinguished from Q. umbrinus by darker coloration of the legs, a larger and more robust body, differently shaped antennae and pronotum, and by the shape of the paramere. LOHSE (1958) already noted that in both external and the male sexual characters Q. cyprusensis much resembled Q. umbrinus . A comparison of specimens previously identified as Q. cyprusensis (see ASSING & WUNDERLE 2001) with material of Q. umbrinus from other regions yield- ed no discrete characters suggesting that the former should represent a distinct species. Neither is the aedeagus of significantly different shape (males of Q. cyprusensis from Cyprus and Lebanon examined), nor are the specimens significantly larger than material of Q. umbrinus seen, e.g., from Greece , Georgia and Armenia , even though they are at the upper end of the size range. Specimens examined from Cyprus and Lebanon belong to the macropterous morph (hence the long elytra). I have seen macropterous and similarly dark-coloured specimens also from South Greece , Armenia , and Georgia . In consequence, both Q. cyprusensis and its junior synonym Q. freyi are placed in synonymy with Q. umbrinus . The original description of Q. bulgaricus is based on " 2♂♂ ( 1 Typus , 1 Cotypus) und 2♀♀ ( 1 Typus , 1 Cotypus) vom Rosalito-polje, Schipka Balkan" and " 1♀ vom Jumruktchal, Zentral-Balkan... und 1♀ von ebendort... (Cotypen)" ( SCHEERPELTZ 1937 ). Since a holotype is not specified, all the type specimens have syntype status. The male labelled by Scheerpeltz as "Typus" is designated as the lectotype . According to the key provided by SCHEERPELTZ (1937) , Q. bulgaricus is mainly distinguished from Q. umbrinus by the shape of the head, shorter hind wings, and the absence of a palisade fringe at the posterior margin of the abdominal tergite VII. An examination of the type specimens revealed, however, that the shape of the head, the length of the hind wings, and the shape of the aedeagus are within the range of intraspecific variation of Q. umbrinus and that the palisade fringe at the posterior margin of tergite VII is indeed present. The median lobe of the aedeagus is of similar shape as that of material seen from Italy and Georgia . Hence the synonymy proposed above. Externally, the populations from the Stara Planina are characterized by rather extensive yellowish coloration of the lateral and posterior margins of the elytra. Quedius maronitus was described from a unique male collected in " Liban , Aïn Batara" ( COIFFAIT 1963 ). The holotype is deposited in the Coiffait collection and consequently unavailable for taxonomic revision (see introduction). However, based on the descriptions and illustrations of the aedeagus provided by COIFFAIT (1963 , 1978a ), as well as on additional material from Lebanon , the holotype is undoubtedly a macropterous specimen of Q. umbrinus . Quedius gueorguievi was described based on a male holotype and two paratypes from "Grotte Porojnata Dupka, Bulgarie " ( COIFFAIT 1967 ). The type material is currently inaccessible for scientific study (see introduction). However, neither the illustrations of the aedeagus nor the description provided by COIFFAIT (1967) provide any evidence suggesting that Q. gueorguievi should be distinct from Q. umbrinus . The shape of the median lobe of the aedeagus is identical to that of Q. umbrinus and the shape and chaetotaxy of the paramere (figure 2L in COIFFAIT 1967 ) are within the range of its intraspecific variation. Hence the synonymy proposed above. I n t r a - a n d i n t e r s p e c i f i c v a r i a t i o n: Quedius umbrinus is subject to enormous variation not only of external characters such as size, coloration, and the length of the hind wings and the elytra, but also of the shape of the aedeagus. The macropterous morph, which is usually of more robust habitus, darker coloration, and larger size, was observed in populations from South Greece , Cyprus , the Middle East, and the Caucasus region ( Georgia , Armenia ). In Armenia , it was repeatedly collected together with the brachypterous morph. The coloration of the body is mostly dark-brown to blackish-brown, but ranges from nearly uniformly reddish (some specimens from North Germany seen) to partly or completely black. The elytra may be bicoloured (dark with more or less extensively paler margins and suture) or unicolorous, and the colour of the legs ranges from yellowish-red (usually with the meso- and metatibiae more or less distinctly infuscate) to blackish. The position of the subapical tooth of the median lobe of the aedeagus may be very close to the apex or more distant (with all transitions). The paramere varies in breadth and, to a lesser extent, also in length; it may be basally constricted or not. The apex of the paramere may be smoothly rounded or obtusely pointed; the sensory peg setae may be arranged in two distinct lateral rows or occupy nearly all of the apical portion of the paramere. Some of the aedeagal variation is illustrated by ASSING (1999) . Quedius sigwalti COIFFAIT, 1972, an endemic of Crete and probably the closest relative of Q. umbrinus , resembles Q. umbrinus both in external and in sexual characters, but is distinguished by a much broader paramere. Since transitional conditions have not been observed, this difference is interpreted as interspecific variation. It seems likely that Crete was colonized by the ancestors of Q. sigwalti millions of years ago and that the Cretan populations have been isolated ever since. Quedius umbrinus is widespread and common in practically all of Europe (eastwards to the Caucasus region), Turkey , and the Middle East (material from France , Germany , Slovakia , Austria , Italy , Slovenia , Romania , Serbia , Albania , Bulgaria , Greece , Cyprus , Turkey , Lebanon , Russia (West Caucasus), and Armenia examined). The species is an epigeic and eurytopic inhabitant of moist habitats.