Revision of the Quedius fauna of Middle Asia (Coleoptera, Staphylinidae, Staphylininae)

Twenty eight species of the genus Quedius from Middle Asia comprising Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan and Uzbekistan, are revised. Quedius altaicus Korge, 1962, Q. capitalis Eppelsheim, 1892, Q. fusicornis Luze, 1904, Q. solskyi Luze, 1904 and Q. cohaesus Eppelsheim, 1888 are redescribed. The following new synonymies are established: Q. solskyi Luze, 1904 = Q. asiaticus Bernhauer, 1918, syn. n. ; Q. cohae-sus Eppelsheim, 1888 = Q. turkmenicus Coiffait, 1969, syn. n. , = Q. afghanicus Coiffait, 1977, syn. n. ; Q. hauseri Bernhauer, 1918 = Q. peneckei Bernhauer, 1918, syn. n. , = Q. ouzbekiscus Coiffait, 1969, syn. n. ; Q. imitator Luze, 1904 = Q. tschinganensis Coiffait, 1969, syn. n. ; Q. novus Eppelsheim, 1892 = Q. dzambulensis Coiffait, 1967, syn. n. , Q. pseudonigriceps Reitter, 1909 = Q. kirklarensis Korge, 1971, syn. n. Lectotypes are designated for Q. asiaticus Bernhauer, 1918, Q. fusicornis Luze, 1904, Q. hauseri Ber-nhauer, 1918, Q. imitator Luze, 1904, Q. novus Eppelsheim, 1892 and Q. solskyi Luze, 1904. For all revised species, taxonomy, distribution and bionomics are summarized. Quedius fuliginosus (Gravenhorst, 1802), Q. sundukovi Smetana, 2003 and Q. pseudon-igriceps Reitter, 1909 are recorded for Middle Asia for the first time. One species from the Q. coloratus -group, found to be new to science is not described due to shortage of material. Another possibly new species is tentatively identified as Q. fulvicollis Stephens, 1833 until the taxonomy of that widespread species is revised. An identification key to all species is provided.


Introduction
The rove beetle genus Quedius Stephens, 1829 is one of the largest in the family Staphylinidae.Even according to a recent phylogenetic study (Brunke et al. 2016) which restricted Quedius to a cluster of lineages confined mostly to the Holarctic region, it remains a very speciose taxon to deal with.The greatest diversity of Quedius in this restricted sense, ca.700 species, is confined to the humid areas of the Palaearctic region (Herman 2001;Schülke and Smetana 2015).A satisfactory alpha-taxonomic knowledge of the mega-diverse Palaearctic fauna of Quedius is crucial for implementing a badly needed phylogenetic mon (Kryzhanovsky 1965).However, one must bear in mind the poor correspondence of this large territory to biogeography.Due to certain patterns of geography, landscape mosaic and biogeographic history, various areas of Middle Asia may show stronger faunal connections with other respective neighboring regions than to each other.Nevertheless, we limit our paper by the formal political borders of the listed countries for practical reasons.As the former republics of the Soviet Union, they often were (and often still are) studied together.As a result, legacy taxonomic and faunistic literature considers Middle Asia largely within these borders.
Where necessary, we have considered literature or material from areas outside Middle Asia.However, species known only from outside this region were not included in this paper.One rather specialized and distinct group of species related to Quedius (Microsaurus) mutilatus, which comprises endemic Middle Asian species with narrow montane distributions, has been revised in a separate publication (Salnitska and Solodovnikov 2018).However, species of the Q. mutilatus group are here included in the identification key to all species of Quedius currently known from Middle Asia.We hope that this taxonomic revision and the first specialized key of Middle Asian Quedius will stimulate further investigations of the genus in this and adjacent poorly known areas of the Palaearctic region.

Depositories of material
Material for this paper is deposited in the public institutions and private collections abbreviated as follows: Private collection of M. Schülke, Berlin, Germany (to be deposited at the Museum of Natural History, Berlin, Germany)

Preparation, examination and illustration of specimens
Specimens were examined with Lomo MSP-2 ver. 2 and Leica M125 dissecting scopes.Habitus and genitalia photographs were obtained using a Nikon SMZ 1500 binocular microscope with a Nikon D700 digital SLR camera.Illustrations of the male genitalia were done from soft preparations of these structures in glycerin (after dissecting, maceration in 10% KOH, and rinsing in distilled water) using a drawing tube attached to a Nikon SMZ 1500 binocular microscope.All dissected aedeagi are kept in glycerin in genitalia microvials pinned under their respective specimens.

Measurements
Measurements were taken at X4.5 magnification using an ocular micrometer.They are abbreviated as follows: HL -head length (from base of labrum to neck constriction along the head midline); HW -head width (maximum, including eyes); PL -pronotum length (along midline); PW -pronotum width (maximum); EL -length of elytra (from humerus to the most distal part of the elytral posterior margin); EW -width of elytra (maximum, with elytra closed along suture).Overall body length was measured from apex of labrum to apex of abdomen.

Type material
Where possible type material was examined and supplied with our standard respective labels indicating the revised status or identity of the respective type specimens.All original labels of the type specimens are cited verbatim in the 'Material examined' sections and, where available, photographed.

Classification
We use conventional subdivision of the genus Quedius into subgenera as in e.g.Schülke and Smetana (2015).
Within the subgenera we list species so that those we presume to be closely related appear close to each other.Except the recently defined coloratus-group (Assing 2017) and mutilatus-group (Salnitska and Solodovnikov 2018), we cannot use any of the hitherto proposed species groups in Quedius.Species groups of Coiffait (1978) for the West Palaearctic fauna are very outdated, inconsistent and even lack any diagnoses.Among those of Smetana (1971Smetana ( , 1988Smetana ( , 1992Smetana ( , 1995bSmetana ( , 1996Smetana ( , 2001Smetana ( , 2015aSmetana ( , 2017)), species groups proposed for the fauna of China (Smetana 2017) are worth consideration, especially given that the large Xinjiang province of the north-western China borders with Middle Asia via Tajikistan, Kyrgyzstan and Kazakhstan.However, that large province of China seems to be one of the least explored areas there, what can be seen for example, from the lacking records for any wide-spread Middle Asian species from that province.Therefore, placing Middle Asian species in the species groups of Smetana (2017) was not possible, at least without extensive direct comparisons with the material from China.We can only propose that among the Middle Asian species, Q. hauseri and a species tentatively identified here as Q. fulvicollis may be related to the Chinese muscicola-group.Also, it should be noted that Smetana (2017) placed Q. koltzei in its own monotypic species group.We should also point to our disagreement with Smetana (2017) who considers Q. equus a member of the przewalskii-group, while we place it in the mutilatus-group (Salnitska and Solodovnikov 2018, and here).These disagreements are not essential for the taxonomic purposes of this paper and they once again call for a necessity of a large-scale phylogenetic study of Quedius.All species treated in this revision are also listed alphabetically in Table 1.

Distribution maps
All distributions were mapped using QGIS 2.12.0 and geographical coordinates indicated on the original locality labels of the specimens.In the case of older, non-georeferenced labels, we used approximate geographic coordinates that we were able to find for the respective toponyms with the aid of various printed maps or online systems (Google Maps, Google Earth, Global Gazetteer version 2.3 and others).Ambiguously indicated localities are cited verbatim in the 'Material examined' sections and taken in quotation marks.All our interpretations for such localities are given in square brackets.Those of which that are mapped are also given with their approximate coordinates in Table 2.

Borders and geography of Middle Asia
The term "Middle Asia" is somewhat fuzzy in the geographical or historical literature.For example, sometimes Kazakhstan is considered as a part of Middle Asia, sometimes an expression "Middle Asia and Kazakhstan" is used.Often the distinction between "Middle Asia" and "Central Asia" is not clear.English-language publications have used "Central Asia" to refer to areas of the former USSR, to areas of China and Mongolia and to areas that cross the former Soviet/Chinese border.To avoid this ambiguity we follow Cowan (2007) and use "Middle Asia" to refer to Kazakhstan, Turkmenistan, Uzbekistan, Tajikistan and Kyrgyzstan collectively.The geographic area covered by these five countries is a subject of this paper (Fig. 1).In the west, Middle Asia is bordered by the Caspian Sea and the state border between Russia and Kazakhstan, nearly coinciding with the Volga River.In the north, Middle Asia is outlined by the long administrative border between Kazakhstan and Russia.In the east, Middle Asia borders with north-western China through the eastern administrative borders of Kazakhstan, Kyrgyzstan and Tajikistan.In the south, it is outlined by the northern borders of Afghanistan and Iran.While large areas of Kazakhstan and Turkmenistan are covered by more or less flat, desert landscape, eastern and south-eastern Kazakhstan, as well as Kyrgyzstan and Tajikistan are mainly montane countries with complex relief and a diverse landscape mosaic.In north-eastern Kazakhstan, the Altai mountain chain stretches into Middle Asia from Russia.In eastern Kazakhstan, as well as in Kyrgyzstan and Tajikistan, the area is dominated by the vast mountain systems of Tien Shan and Pamir.Large lakes like the Aral Sea, Balkhash, Issyk-Kul, and rivers like Amu Darya or Syr Darya are significant elements in the geography of Middle Asia as well.
The next notable contribution to the study of Middle Asian Quedius was made in the papers by Coiffait (1954Coiffait ( , 1955Coiffait ( , 1963Coiffait ( , 1967Coiffait ( , 1969Coiffait ( , 1970Coiffait ( , 1975Coiffait ( , 1978) ) devoted to the Western Palearctic fauna.Henry Coiffait added aedeagus illustrations for many Middle Asian species and integrated them in his identifications keys for the Western Palearctic Quedius.He also described Q. (R.) dzambulensis Coiffait, 1967 Coiffait, 1969, and Q. (M.) tadjikiscus Coiffait, 1975, all from Middle Asia.Unfortunately, Coiffait's input was based on very limited material from Middle Asia and additionally suffered from inconsistent study of type material and omissions of the earlier literature.For example, three species described from Middle Asia by Luze (1904), Q. (M.) fusicornis, Q. (M.) rufilabris and Q. (M.) solskyi, were entirely overlooked in the influential monograph of Coiffait (1978) and have not been studied since their original description.The most unfortunate flaw of Coiffait's taxonomy was an artificial and over-splitting approach to species.As a result, all species of Quedius from Middle Asia he described as new, except Q. (M.) tadjikiscus, turned out to be synonyms here.
Finally, some bionomic and distributional data on Middle Asian Quedius were published by local authors stationed in that region (Kascheev, 1984(Kascheev, -2002;;Kadyrov et al., 2014a, b;Gabdullina, 2016).With the scattered, confusing and then poorly accessible taxonomic literature on Quedius, no surprise that their local faunistic papers were greatly infested by incorrect species identifications.Examination of the material collected by Kascheev (1984Kascheev ( -2002)), now deposited at ZIN, largely helped to reveal such misidentifications summed up in the Table 1 here.
Overall, due to a hitherto lacking targeted contemporary taxonomic investigation of the Middle Asian Quedius, identity of the majority of species described from, or recorded for, that region remained highly ambiguous.Most of the species described from Middle Asia needed broader comparisons and a revision of the type material.At the same time, a number of widespread species from Middle Asia were misidentified or overlooked.A large amount of Quedius material from Middle Asia remained undetermined and scattered in some institutional and private collections.The revision of Q. (M.) mutilatus species group by Salnitska and Solodovnikov (2018) was the only recent taxonomic work that touched upon Middle Asian Quedius.
According to the latest phylogenetic hypotheses (Solodovnikov, 2006;Chatzimanolis et al., 2010;Brunke et al., 2016) the genus Quedius as it stands now in the taxonomic literature (e.g., summaries in Herman, 2001 or Schülke and Smetana, 2015) is a polyphyletic assemblage of species belonging to several different subtribes of Staphylinini.Within the Palaearctic or Middle Asia, all species of Quedius are members of the subtribe Quediina in the restricted sense of Brunke et al. (2016).Because of the polyphyly, Quedius in the current composition lacks synapomorphies and clear diagnosis.However, genus descriptions and diagnostic combination of characters that can define any Palaearctic species as a member of the genus Quedius are available in Coiffait (1978), Smetana (1988), Assing and Schülke (2012) and other sources.The diagnosis of the genus Quedius and comparative notes we provide here are tuned for the fauna of Middle Asia.
Adults and larvae of Quedius seem to be predators hunting small invertebrates in various, sufficiently hu-mid ground-based debris, mostly in forest leaf litter.In a largely arid region like Middle Asia, Quedius are mainly confined to humid open or forested habitats along creeks or rivers in the lowland or forests, meadows, snowfield margins and talus in the mountains.Some members of the subgenus Microsaurus are specialized inhabitants of mammal burrows.Overall, bionomics of the genus in Middle Asia remain largely unstudied.
Diagnosis.Medium to large size (body length 3.5-24.0mm) rove beetles with glossy forebody, infraorbital ridges extended from neck to base of mandibles and pronotal hypomera strongly inflexed under pronotal disk (not visible in lateral view).First segment of antennae at most slightly longer than second and third segments together.Last segment of maxillary palps fusiform, not densely setose.Tarsal formula 5-5-5; anterior tarsi widened in both sexes, with pale adhesive setae ventrally, with pair of empodial setae.Males always with distinct apical emargination on abdominal sternite VIII.Aedeagus varies in shape, paramere mostly with sensory peg setae.

Q. ochripennis Ménetries, 1832 Microsaurus
Gridelli 1929, 21;Klimenko 1996, 121;Kascheev 2001, 102;Kadyrov et al. 2014a, 31;2014b Comments on taxonomy, distribution and bionomics.Bernhauer (1908) described Q. curtipennis as a variety of Q. fuliginosus without clear information on the type material.In addition to the morphological diagnosis of a new variety Bernhauer (1908) mentioned that it is common on the Faroe Islands and also occurs in "Vorarlberg, Buchara and Böhmen (Wrana.Moldau)".Interestingly, revision of the type material has never been published for this com-mon widespread species since its original description.We have examined three male syntypes of Q. curtipennis kept at the FMNH, two from Faroe Islands and one from 'Buchara' in Uzbekistan, all listed above.Our examination of the syntypes confirms that they are conspecific and matching current interpretation of this species (e.g. in Assing and Schülke 2012).Quedius curtipennis is a common species widely distributed in the forests and humid microhabitats of the open landscapes of the Western Palaearctic (Herman 2001;Assing and Schülke 2012).Because of the strong morphological similarity, Q. curtipennis can be easily confused with Q. fuliginosus.As a result, current broad distributions for both species as recently summarized in Assing and Schülke (2012), especially outside Europe, need revision.A male syntype of Q. curtipennis from "Buchara" (Uzbekistan) collected more than a century ago (see below) and overlooked in the subsequent literature is the only specimen of this species known from Middle Asia.Since the original description Q. curtipennis has not been recorded from any of the countries of Middle Asia.Comparative material on Quedius subunicolor.Type material: Paratypes: Sweden: 1 ♂, "Häggenäs s-n Jtl.T. Palm 4-8, 8 1945 [printed] Comments on taxonomy and type material.The original description of Quedius altaicus was based on two female specimens (a holotype and a paratype) from "Central-Altai" without precise record of the type locality (Korge, 1962).Such ambiguity was stressed by Korge who noted that the status of Q. altaicus, which externally appeared very similar to Q. unicolor and Q. subunicolor, should be confirmed by the examination of male genitalia.Toleutaev (2014) recorded Q. altaicus from Saur Mountains (Eastern Kazakhstan), but that record needs verification.

Quedius (s. str.) fuliginosus Gravenhorst, 1802
In spite of the ambiguous original description of Q. altaicus, new material from Altai including males examined here for the first time can be safely attributed to that species.This material perfectly matches Korge's original description, and the information together with high quality photos of the holotype available from the Field Museum online beetle type database (FMNH, 2018).Besides, there are no other species in the Altai region that could be misidentified as Q. altaicus.Quedus sundukovi, the only other similar species distributed from the Russian Far East to the South-Western Altai is distinctly different (for details see below).Comments on taxonomy, distribution and bionomics.Similarly to Q. curtipennis (see above), Q. fuliginosus is a widespread and common species in the Western Palaearctic, and subject of numerous publications.The most recent summary of its diagnostic characters, bionomics and distribution can be found in Assing and Schülke (2012).For the same reasons as Q. curtipennis above, the exact distribution of Q. fuliginosus needs careful revision.Limited material from Kazakhstan and Uzbekistan examined here represents the first records of this species from Middle Asia.Quedius altaicus Korge, 1962, 152 (original   The aedeagus of Q. altaicus (Fig. 5F, G) here examined for the first time is nearly identical with the aedeagus of the northern European Q. subunicolor (Fig. 5B,  C).Both species slightly differ from each other in the shape of a large sclerite in the internal sac (labeled as H in Fig. 5B, F) and the degree of development of the subapical teeth of the median lobe (less pronounced in Q. altaicus, compare Fig. 5B, F).Comparison of the external morphology of the multiple specimens of Q. altaicus to each other and with the available specimens of Q. subunicolor, including its paratypes, demonstrates that the external characters provided by Korge (1962) as unique for Q. altaicus (microstructure of the head, proportions of the pronotum, chaetotaxy of the head and pronotum) do not hold.Given a subtle morphological difference between both species and poorly sampled areas of Russia, there remains a possibility that Q. subunicolor may be a polytypic species continuously distributed from the Northern Europe to Altai.Or, Q. subunicolor and Q. altaicus may be a hitherto unrecorded case of the boreo-montane distribution.Both species should be subject to further sampling in the area which seems as a distribution gap between them.Also, a DNA-based phylogeographic investigation would be interesting.Below we provide a redescription of Q. altaicus.
Head with broadly rounded, but distinct hind angles with microsculpture consisting of transverse waves; eyes as a long as or slightly longer than tempora; posterior frontal puncture situated closer to posterior margin of head than to posteromedial margin of eye; two to four additional punctures present along medial margin of eye between anterior and posterior frontal punctures; temporal puncture situated close to posterior margin of eye at distance nearly equal to diameter of puncture.
Antennae moderately long, segment 3 somewhat longer than 2, segments 4-8 longer than wide, each gradually becoming shorter towards apex, segments 7-11 about as long as wide.
Pronotum wider than long PL/PW: 0.9-1.0(1.0), widest at posterior third, narrowed anteriad; hind angles broadly rounded, but distinct; dorsal rows each with three punctures; sublateral rows each with two to three punctures; waves of microsculpture transverse, similar to that on head.Scutellum finely punctured in its posterior half, with transverse or slightly isodiametric microsculpture.
Elytra parallel-sided, as long as pronotum, at base narrower than pronotum at widest point; shiny, punctation moderately dense and shallow, interspaces larger than diameter of punctures, pubescence yellowish-grey.
Abdomen with tergite VII (5th visible) with fine distinct whitish apical seam of palisade fringe; punctation dense and fine gradually becoming sparser towards apex of abdomen, surface between punctures with very superficial transverse irregularities, pubescence as on elytra.
Male: aedeagus: median lobe with acute apex and small teeth on its parameral side near apex (Fig. 5B, F); paramere distinctly protruding over apex of median lobe, with two pairs of setae apically and two pairs of longer setae laterally below apex, its underside with numerous sensory peg setae forming two subapical longitudinal rows connected near apex (Fig. 5C, G).Internal sac (examined in situ) with two pairs of strongly sclerotized microstructures positioned laterally and one characteristically shaped medial sclerite (Fig. 5H) with rounded apex.
Comparison.Based on the structure of the aedeagus, especially the characteristic armature of the internal sac with the large middle sclerite 'H' (Fig. 5B, F; fig 189 in Assing and Schülke, 2012), Q. altaicus can be placed in the group with Q. subunicolor, Q. balticus, Q. molochinus and Q. meridiocarpathicus.Quedius altaicus differs from Q. meridiocarpathicus in the unicolorus black coloration of the body (brown reddish with paler elytra in Q. meridiocarpathicus) and in the shape of the medial sclerite of the internal sac that has rounded apex.Some authors stressed a strong similarity of Q. subunicolor (from which Q. altaicus is hardly distinct) with Q. unicolor, and the latter mainly Central European montane species was incorrectly cited as Q. subunicolor in a number of the faunistic papers (e.g., Ciceroni and Zanetti 1995;Geiser et al. 2003;Boháč et al. 2004Boháč et al. , 2005;;Wojas 2006).Quedius subunicolor (and Q. altaicus), however, can be easily distinguished from Q. unicolor by transversal (not isodimetric) microsculpture of the frons and the structure of the aedeagus, especially by the internal sac with the obvious medial sclerite.From similar species that occur in Middle Asia Q. altaicus can be easily distinguished by the following characters: from Q. fuliginosus by the punctured (setose) scutellum and absence of additional punctures between anterior frontal punctures; from Q. sundukovi by normally developed elytra (very short in distinctly brachypterous in Q. sundukovi), presence of fine whitish apical seam of palisade fringe on VII tergite (5th visible), and distinctly larger body.
Distribution.Quedius altaicus is known from "central" (Korge, 1962) and southwestern Altai.Records from the southwestern Altai stretching across the border between Russia and Kazakhstan, provided here, are the first exact distributional data for this subspecies.We were not able to examine the material on which Toleutaev (2014) recorded this species from Saur Mountains, the latter records remains ambiguous.
Bionomics.All clearly georeferenced specimens of Q. altaicus have been collected at the elevations between 1200 and 2000 m.Comments on taxonomy, distribution and bionomics.Quedius sundukovi was known from the Russian Far East (Smetana, 2003) and from Irkutsk Province and Zabaikalsky Territory (Smetana and Shavrin 2018).From the newly examined material it has become clear that Q. sundukovi is distributed even wider: from the Russian Far East through southern Siberia to Altai Mountains in Northeastern Kazakhstan.In the material examined we here provide only new records for Middle Asia, because the detailed documentation of its entire distribution will be published elsewhere.

Quedius (s. str.) sundukovi Smetana, 2003
Detailed description and illustration of the species is available in Smetana (2003).Quedius sundukovi is one of the smallest species in the nominative subgenus Quedius s. str.and the smallest in this subgenus in the fauna of Middle Asia.Additionally, it stands out from all other Quedius s. str.species in Middle Asia as the only distinctly brachypterous species, with very short elytra and lacking whitish apical seam on abdominal tergite VII.
All hitherto known specimens of Q. sundukovi were collected by pitfall traps (Smetana 2003).Based on the newly examined material here, Q. sundukovi inhabits talus-associated debris.Also it is found in regular leaf litter and moss on the ground.

Quedius (s. str.) vicinus Ménétriés, 1832
Fig. 2C Quedius vicinus Ménétriés, 1832, 144 (original  Comments on taxonomy, distribution and bionomics.Gusarov (1993) clarified the identity and synonymy of Q. vicinus.A description and the illustrations of the aedeagus for this species can be found, for example, in Coiffait (1954Coiffait ( , 1978, as Q. libanicus), as Q. libanicus), Boháč (1988, as Q. libanicus), andHachikov (2003).By the brown body with reddish elytra, frons without additional setiferous punctures between anterior frontal punctures and by punctate scutellum, Q. vicinus can be distinguished from all other Middle Asian species of the nominal subgenus Quedius s. str.based on the external characters alone.The aedeagus of this species, with the parameral apex pointing outwards from the median lobe, is also very characteristic.
Based on literature data it is a common species in Western Asia.In Middle Asia it was known only from Turkmenistan (Boháč 1988, as Q. libanicus).Based on the newly examined material Q. vicinus is recorded for the first time from southern Kazakhstan (Karatau Mountains).The species prefers ground based debris and leaf litter, but also was found (Coiffait 1955, as Q. libanicus) in caves.Specimens from Middle Asia were collected along rivers, in ground traps and at light.

Comments on taxonomy, distribution and bionomics.
The diagnostic characters including illustrations of the aedeagus and the most recent summary of the bionomic and distribution data of this widespread and rather common Western Palaearctic species can be found in Assing and Schülke (2012).From similar Middle Asian species Q. capitalis and Q. fusicornis, Q. ochripennis is distinguished by the larger body and shape of the aedeagus.From the larger Q. solskyi it can be safely distinguished by the characters of aedeagus.
Quedius ochripennis is widely distributed in Europe and in the Mediterranean region.It is also known from Simla Hills in Himalaya, India (Smetana, 1988) and from Middle Asia where, based on earlier records (Table 1) and material examined here, it occurs in southwestern Turkmenistan, entire territory of Tajikistan, and southern Uzbekistan.
Quedius ochripennis inhabits various ground based debris, often associated with decaying wood, also in nests of mammals, ants and wasps (Assing and Schülke 2012).Based on the material examined here, in Middle Asia Q. ochripennis prefers humid plant debris usually near water bodies, also it was found in caves and in tree foliage in an aphid nest.In the mountains it was recorded at elevations up to 2000 m.Comments on taxonomy, distribution and bionomics.Quedius puncticollis is widely distributed in Northern, Central and Eastern Europe for which the latest summary of diagnostic characters, distribution and biology can be found in Assing and Schülke (2012).It is most similar to Q. ochripennis from which it can be easily distinguished by the apically not lanceolate paramere with a broad and shallow apical emargination.

Quedius (Microsaurus) puncticollis Thomson, 1867
Quedis puncticollis is commonly found in the burrows of small mammals, especially moles, also in bee and wasp nests (Assing and Schülke 2012).Based on literature data (Table 1) and material examined here, there are only few records of Q. puncticollis in Middle Asia: from southern Kazakhstan and southern Tajikistan.Comments on taxonomy and type material.In the original description, Eppelsheim (1892) mentioned morphological characters of both males and females and stated that the species was known to him from a few specimens from ''Tashkent''.As we learn from the introduction in his paper, specimens were collected by Hans Leder.Both male specimens from NMW labeled as "types" of Q. capitalis originally come from Eppelsheim's collection and their morphology and label information fit the original description; therefore they are syntypes.

Quedius (Microsaurus) capitalis Eppelsheim, 1892
We have examined aedeagi of both syntypes and confirm they are conspecific.Eppelsheim (1892) compared Q. capitalis with Q. ragusai Eppelsheim, 1889. Gridelli (1924), based on the examination of a syntype, provided additional morphological details for the species including verbal description of its aedeagus (but no illustration) and placed it near Q. ochripennis.Based on a syntype male, Coiffait (1978) again redescribed this species and provided its first and hitherto the only available illustration of the aedeagus.Our examination of the syntypes of Q. capitalis confirms the correct identification of this species by both Gridelli (1924) and Coiffait (1978).In Schülke and Smetana (2015) Q. capitalis was erroneously placed in the subgenus Raphirus.Here we redescribe this poorly known species and provide further data on its distribution.
Body black to dark brown, hind margins of abdominal tergites slightly paler; elytra reddish; palpi and other appendages slightly lighter; body glossy (Fig. 7A, B).
Head approximately as wide as long or slightly longer; eyes small, not convex; temples as long as longitudinal diameter of eye; posterior frontal puncture closer to posterior margin of head than to anterior frontal puncture; temporal puncture closer to posterior margin of head than to posterior margin of eye; two vertical punctures behind posterior frontal puncture arranged as slightly oblique line between posterior margin of eye and dorsal part of neck; microsculpture of head with transverse distinct wavelines.
Pronotum slightly wider than long, widest at about middle to posterior third; hind angles rounded but distinct; dorsal and sublateral rows each with three punctures; microsculpture with transverse waves as on posterior part of head.Scutellum impunctate with microsculpture slightly coarser than on pronotum.Elytra parallel-sided, slightly longer than wide, longer than pronotum, their punctation dense, interspaces shiny with distinct minute irregularities.
Abdomen: punctation fine and dense; interspaces with minute irregularities; posterior margin of tergite VII with palisade fringe.
Male: protarsi with tarsomeres 1-4 dilated stronger than in females.Sternite VIII with weak triangular medio-apical emargination; tergite X triangular with setae; sternite IX elongate, gradually narrowed apically, with moderately wide and long basal portion and obtusely rounded apical margin with numerous setae.Aedeagus (Fig. 7C, D): median lobe parallel-sided with broad and obtuse apex and tooth located near apex (Fig. 7 C).Paramere rhomboid sharply narrowing apicad; its apex almost reaching apex of median lobe, with two pairs of apical setae and two pairs of lateral setae below apex; paramere (underside) with ca.4-8 sensory peg setae in each of two sinuate lateral rows that extend basad over pairs of lateral setae (Fig. 7D).
Comparison.Quedius capitalis seems to be closely related to Q. fusicornis and Q. ochripennis from which it can be easily distinguished externally by smaller body size and proportions, and by the structure of paramere with two sinuate lateral rows of peg setae (ca.4-8 in each row) extending basad over pairs of lateral setae.
Distribution.Based on the literature data (Table 1) and newly examined material, Q. capitalis is known from several localities near Tashkent (Uzbekistan), Karatau Mountains (southwestern Kazakhstan) and Hazratisho Mountains (southern Tajikistan). Bionomics.Unknown.

Quedius (Microsaurus) fusicornis Luze, 1904
Fig. 8 Quedius fusicornis Luze, 1904, 28 ( Comments on taxonomy and type material.In the original description, Luze (1904) provided no information on the type material, but he indicated 7.7-8.5 mm body size range for the species.This suggests that he must have had more than one specimen to base a description on.He also indicated "Seravschan: Putschin-Pass, Boschara'' as a locality that his material was from.Finally, we know from the introduction in Luze's paper that the material he examined was collected by Glasunov.Therefore, a single male (NMW) and two females (ZIN) that we examined and that match the original description morphologically and in the label data, are syntypes.Luze (1904) compared Quedius fusicornis with his Q. solskyi and the widespread Q. cruentus Ol.Gridelli (1924) apparently based his short notes about this species exclusively on Luze's description, without seeing any material.Similarly to other species of Quedius described by Luze (1904), Q. fusicornis is missing in the monograph by Coiffait (1978) who apparently overlooked Luze's publication.
Here we provide a redescription and first illustrations of this poorly known species, including its aedeagus.
Head approximately as wide as long HL/HW: 0.9-1.1 (1.0); eyes small, not convex; temples slightly longer or as long as longitudinal diameter of eye; posterior frontal puncture closer to posterior margin of head than to anterior frontal puncture; temporal puncture closer to posterior margin of head than to posterior margin of eye; two vertical punctures behind posterior frontal puncture arranged as slightly oblique line between posterior margin of eye and dorsal part of neck; microsculpture of entire surface of head with transverse waves.
Pronotum slightly wider than long PL/PW: 0.8-0.9(0.9), widest at about posterior third, gradually narrowing anteriad; hind angles rounded but distinct; dorsal and sublateral rows each with three punctures; microsculpture with transverse waves similar to that on posterior part of head.Scutellum impunctate with microsculpture as on pronotum.Elytra parallel-sided, slightly longer than wide, as long as or slightly longer than pronotum and narrower than maximum width of pronotum; punctation dense; setation gray; interspaces shiny, with distinct minute irregularities.
Abdomen: punctation fine and moderately dense; interspaces with vaguely distinct minute irregularities; posterior margin of tergite VII with palisade fringe.
Male: protarsi with tarsomeres 1-4 dilated stronger than in females.Aedeagus (Fig. 8C-E: median lobe parallel-sided along most of its length with broad and obtuse apex and tooth located near apex (Fig. 8C).Paramere parallel-sided, narrowing only in rhomboid apical portion; its apex almost reaching apex of median lobe, with two pairs of apical setae and two pairs of lateral setae below apex, with 6 peg setae arranged in two regular longitudinal rows apically extending basad over pairs of lateral setae (Fig. 8D).
Comparison.Quedius fusicornis is similar to Q. capitalis.For comparison, see the latter species above.From other similar species such as Q. solskyi, Q. cruentus and Q. ochripennis, it can be easily distinguished by the structure of the apical part of the paramere with two medially situated short rows of peg setae (3 in each row) extending basad the pairs of lateral setae.
Distribution.We were not able to locate the type locality "Putchin Pass" situated somewhere along Zeravchan River that is extended from eastern Uzbekistan to western Tajikistan.Additional material was studied from eastern Uzbekistan (near Aman-Kutan) and north-west- ern Kyrgyzstan (Kyrgyz-Alatoo).Finally, one specimen was from 'Tangi-Gharuh', a toponym in Afghanistan that we could not locate.
Bionomics.Unknown.Comments on taxonomy, lectotype designation and new synonymy.In the original description of Q. solskyi, Luze (1904) did not specify the number of specimens he studied, but provided characters for both sexes and the locality "Jagnob: Kol, Schach-Sara'' [Tajikistan, Yaghnob river, Sughd Distr.].Therefore, a male from NMW with the locality label ''Trkst.Jagnob Schach-Sara'' is considered a syntype.We could not locate other syntypes.Gridelli (1924), similarly to the case with Q. fusicornis, based his notes about Q. solskyi only on Luze's description, without checking type material.And as with other species of Quedius described by Luze (1904), Q. solskyi is missing in the monograph of Coiffait (1978).Under the circumstances of uncertain identity of other syntypes, we designate the only available male syntype as a lectotype to unambigiously fix the identity of Q. solskyi.Bernhauer (1918) described Q. asiaticus from ''Ost.Buchara'' and compared it with Q. abietum distributed in southern Europe.Bernhauer (1918) did not even mention Luze's Q. solskyi, even though his description matches the latter species.Both examined syntypes of Q. asiaticus are clearly conspecific in morphology.In order to fix the identity of the species, we designate here one better  preserved male syntype (Fig. 10A, B) with the locality label "Ost.Buchara" exactly matching the data from the original description and the identification label "asiaticus Bernh.Typus" hand written by Bernhauer as a lectotype.

Quedius (Microsaurus) solskyi Luze, 1904
Our examination of the mentioned types of both Q. solskyi and Q. asiaticus undoubtedly reveal they are conspecific.Thus we place Q. asiaticus Bernhauer, 1918 in synonymy with Q. solskyi Luze, 1904 and provide a redescription with the first illustration of the aedeagus of this poorly known species.
Head wider than long HL/HW: 0.7-0.8(0.8), eyes very small, not convex; temples more than two times as long as longitudinal diameter of eye; posterior frontal puncture in the middle between anterior puncture and posterior margin of head; temporal puncture closer to posterior margin of head than to posterior margin of eye; two vertical punctures arranged in almost straight line between posterior frontal puncture and neck; microsculpture with transverse waves.Antennae long: antennal segments: 3rd longer than 2nd; 4th-10th slightly widening towards apex of antenna.
Pronotum slightly wider than long PL/PW: 0.8-0.9(0.9), widest at its middle, slightly narrowing anteriad; hind angles rounded, barely distinct; dorsal and sublateral rows each with three punctures; microsculpture with transverse waves similar to that on posterior part of head.Scutellum impunctate, with microsculpture as on pronotum.Elytra parallel-sided, as long as or longer than wide, narrower and longer than pronotum; punctation dense, setation brownish, interspaces shiny and with distinct minute irregularities.
Abdomen: punctation fine and moderately dense; interspaces with vaguely distinct minute irregularities; posterior margin of tergite VII with palisade fringe.
Male: head wider than long, larger than in females and with longer temples (Luze 1904).Aedeagus (Figs 9B-D,  10C, D, E, F): Median lobe (in parameral view) parallel-sided along most of its length with obtusely pointed apex, with tooth located near apex (Figs 9B , 10C, E).Paramere parallel-sided, its apex almost reaching apex of median lobe; with two pairs of apical setae and two pairs of lateral setae below apex; underside with pair of peg setae close to apical margin on each side of medial emargination (Figs 9C, 10D, F).
Comparison.Quedius solskyi is similar to Q. fusicornis and Q. ochripennis, but it can be externally distinguished from both by the larger body size, distinctly elongated elytra and smaller eyes with their diameter two times as short as tempora.In the structure of the aedeagus Q. solskyi is more similar to Q. fusicornis but differs from the latter by the paramere with incised apex and two pairs of sensory peg setae.The aedeagi of Q. solskyi and Q. ochripennis differ in many ways.
Distribution.Vaguely recorded type localities for Q. solskyi and Q. asiaticus are located somewhere in northern Tajikistan and in eastern Uzbekistan or western Tajikistan.The only additional and better georeferenced specimen examined here comes from western Tajikistan: Ramid, Kafirnigan River. Bionomics.Unknown.

Quedius (Microsaurus) koltzei Eppelsheim, 1887
Fig. 2D Quedius koltzei Eppelsheim, 1887, 420 (original description); Bernhauer and Schubert1916, 425 (calalog); Gridelli 1924, 24 (characters, new  Comments on taxonomy, type material and distribution.Quedius koltzei was described by Eppelsheim (1887) from "Chabarovka" [Khabarovsk, Far East, Russia] based on a single female specimen.Gridelli (1924) basically repeated the original description.Coiffait (1978) interpreted a few males as that species from Terskey-Alatoo, a mountain range in Kazakhstan very far from the type locality of Q. koltzei.Based on that material, he redescribed Q. koltzei again and provided the illustration of the aedeagus for the first time.Smetana (1998) also redescribed Q. koltzei, but based on the holotype.Later, Smetana (2015b) determined one male and one female from Heilongjiang province of China as Q. koltzei and illustrated their genital structures.Smetana's comparison of the Chinese specimens with the type material and geographic proximity of Heilongjiang province to the type locality of Q. koltzei corroborate his identification.Our examination of the male specimens from Terskey-Alatoo from Henry Coiffait's collection that he identified as Q. koltzei revealed that they match as far as we can observe, with the illustrations of Q. koltzei from China in Smetana (2015b).But since Smetana (2015b) did not illustrate the lateral view of the aedeagus, only the re-examination of Chinese and, preferably, additional material may help to clarify the status of Middle Asian specimens from Terskey-Alatoo.In the absence of neg-ative evidence, we consider Q. koltzei as a potentially widespread Asian species.It is also possible that Q. rufilabris, whose identity currently remains ambiguous, is conspecific with Q. koltzei (for details see the former species below).
Quedius koltzei differs from other similar Middle Asian Microsaurus as follows: from Q. fusicornis, Q. capitalis and Q. solskyi in peg setae on paramere arranged in irregular lines or groups; from Q. ochripennis, Q. puncticollis and Q. tadjikiscus in median lobe (in lateral view) narrowing into a blunt, but clear apex and peg setae on paramere arranged in four irregular groups.From Q. bucharensis, a species whose identity remains ambiguous (for details see that species below) Q. koltzei differs in the chaetotaxy of head (posterior frontal puncture situated closer to nuchal ridge than to posterior margin of eye) and pronotum (two punctures in dorsal row and sublateral group always situated before or at most at the same level as large lateral puncture).
Based on the material examined here, we have additional records for Q. koltzei from Kazakhstan.Bionomics remains unknown.
Comments on taxonomy.Luze (1904) Coiffait (1978) overlooked this species in his monograph.Unfortunately, we were unable to find the holotype of Q. rufilabris, but based on its original description all diagnostic characters, especially chaetotaxy of the head and pronotum, match Q. koltzei.Since the presumed type locality of Q. rufilabris is rather remote from the distribution of Q. koltzei, if the latter even occurs in Middle Asia (see above), we treat the former species as different from Q. koltzei, at least until more material from relevant geographic areas will be studied.
Comments on taxonomy.We could not locate and examine the type material of Q. tadjikiscus described from "Tadjikabad, Daran-Nazaran" in Tajikistan, and did not come across any material that could be identified as that species.The description and the illustrations of the aedeagus of Q. tadjikiscus available from Coiffait (1975Coiffait ( , 1978) ) suggest that this may be a species very similar to Q. koltzei.But Quedius tadjikiscus differs from Q. koltzei in the presence of three punctures in the dorsal row of pronotum and the absence of apical groups of peg setae on the paramere.
We have examined one male and one female from the FMNH which are clearly syntypes of Q. bucharensis.Of them, a male specimen was earlier dissected and its aedeagus must have been glued on the card point beside the specimen, but was obviously lost.Since there were no publications with the structure of Q. bucharensis aedeagus, the identity of this species remains ambiguous.An additional two females from NHMW with the same locality labels as in the original description but without Bernhauer's handwritten type labels, seem conspecific with both mentioned syntypes even though they are somewhat smaller than the latter.Their earlier identifications as Q. solskyi, evident from the labels, are wrong because of the following characters: chaetotaxy of head with posterior frontal puncture closer to posterior margin of eye than to nuchal ridge, larger eyes, head longer than wide and as long and as wide as elytra.
The material used by Bernhauer (1918) in the original description of Q. bucharensis comes from localities rather remote from each other.Given that and the body size variation among the examined specimens from different localities, it is possible that they are not conspecific.On the other hand, significant intraspecific variability in body size and proportions is usual in some Microsaurus species.More extensive material including males is needed to clarify the case.

Quedius (Microsaurus) mutilatus-group
Fig. 2E Comments.We have proposed the mutilatus-group for several Middle Asian species in Salnitska and Solodovnikov (2018), where we revised all available material.Thus only brief information for each of these species is provided with reference to the revision for details.
Diagnosis.The mutilatus-group is characterized by the following: brown to dark brown dorso-ventrally flattened body, notably small eyes, short elytra, absence of palisade fringe on abdominal tergite VII; aedeagus robust, with apical portion of median lobe slightly curved towards paramere with characteristic tooth near apex (in lateral view), with paramere widest shortly before apex (in parameral view) having four distinct groups of sensory peg setae on the underside: two apical and two lateral.
Distribution and biology.The mutilatus-group is restricted to the Tien-Shan Mountains where all species of the group are confined to high elevations, up to 3600 m.Based on the morphology and limited bionomic data, all species of the group are hypogean and are mostly found under stones or deep in leaf litter.Diagnosis.Quedius mutilatus is most similar to Q. kungeicus from which it can be distinguished by the rhomboid shape of the paramere with slight apical incision; by the less curved apical portion of the median lobe (lateral view) with more stronger ventral sub-apical tooth.From Q. kalabi and Q. equus it differs by the not so deeply incised apex of paramere and distinctly larger number of sensory peg setae in lateral groups on the paramere.

Quedius (Microsaurus) mutilatus
Distribution.Quedius mutilatus is restricted to the central part of Terskey-Alatoo Mountains south from Issyk-Kul lake in Kyrgyzstan.Smetana, 1995 figs 1, 3, 4C-D in Salnitska andSolodovnikov 2018 Diagnosis.Quedius kalabi differs from all other species of the mutilatus-group by its narrower and somewhat curved apical portion of the median lobe of the aedeagus with relatively short blade of its subapical tooth (aedeagus in lateral view).In shape of the apical portion of the paramere and degree of its incision Q. kalabi displays a transition between Q. mutilatus having lesser incised paramere with more peg setae in lateral groups, and Q. equus having deeper incised paramere with lesser peg setae in lateral groups.

Quedius (Microsaurus) kalabi
Distribution.Quedius kalabi replaces Q. mutilatus in the eastern part of Terskey-Alatoo Mountains in Kyrgyzstan.Diagnosis.Among all species of the group, Quedius kungeicus can be distinguished by the ovoid apical part of the paramere without a distinct apical incision (in parameral view) and by the distinctly curved apical portion of the median lobe (in lateral view) with longer tip and without distinct sub-apical tooth.

Quedius
Distribution.Quedius kungeicus is known only from the holotype collected in the Kungey-Alatoo Mountains of Kazakhstan.

Quedius (Raphirus) limbatus Heer, 1839
Fig. 3D Herman 2001, 3187 (summary of literature); Kascheev 2001, 102;2002, 181  Comments on taxonomy, distributon and bionomics.The latest summary of diagnostic characters, bionomics and distribution of Q. limbatus, a common Western Palearctic species can be found in Assing and Schülke (2012).Based on earlier records (Table 1) and newly examined material in Middle Asia it is known from southern Kazakhstan and Turkmenistan.
Among all Middle Asian species Q. limbatus is more similar to Q. cohaesus from which it can be easily distinguished by the structure of aedeagus with a sharper apex of the median lobe (in lateral view) and sensory peg setae of the paramere (underside) arranged in short regular rows, slightly diverging from each other basally.
Usually this species occurs in lowlands up to the subalpine zone, but is mostly confined to forests and humid ground-based debris, often near streams (Assing and Schülke 2012).In Middle Asia Q. limbatus was collected at elevations up to 1750 m near rivers in forested landscapes.Quedius kirklarensis Korge, 1971 pseudonigriceps from South Europe and Western Asia except for the absence of palisade fringe on abdominal tergite VII in the former.Therefore he suggested that Q. kirklarensis may be a brachypterous form of Q. pseudonigriceps.A new synonymy could not be established back then because of the limited material and also due to the similar species Quedius cohaesus and Quedius turkmenicus from Middle Asia.With more material available here for all relevant taxa we can undoubtedly place Quedius kirklarensis Korge, 1971 in synonymy to Q. pseudonigriceps Reitter, 1909.For details on Quedius cohaesus and Quedius turkmenicus, see below.

Quedius (Raphirus) pseudonigriceps Reitter, 1909
In Middle Asia Q. pseudonigriceps usually inhabits moist leaf litter in deciduous and mixed forests and wet ground debris near streams in the mountains at the altitudes up to 2800 m.

Quedius (Raphirus) cohaesus Eppelsheim, 1888
Fig. 12 Q.afghanicus Coiffait, 1977, syn.n. (Fig. 13) Q. turkmenicus Coiffait, 1969, syn Coiffait, 1977, 139 (original description).Quedius turkmenicus Coiffait, 1969, 49 (original description); Coiffait 1978, 245 (characters, notes).Comments on taxonomy and new synonymy.Coiffait (1969Coiffait ( , 1977) ) described Q. turkmenicus and Q. afghanicus from Turkmenistan and Afghanistan, respectively.We were able to study the type material for Q. afghanicus only (Fig. 13), which turns out to be conspecific with Q. cohaesus and therefore is placed here into synonymy with the latter.Unfortunately, we were unable to examine the type material of Q. turkmenicus which, according to Coiffait ( 1969) is deposited in the collection of the Paul Sabatier University at Toulouse, France.Nevertheless, because it is obvious from the original descriptions and illustrations that Q. turkmenicus is conspecific with Q. cohaesus, the former is also placed into synonymy with the latter.These new synonymies are consistent with the earlier revealed synonymy of Q. cohaesus with Q. meurguesae Coiffait, 1977from Iran (Solodovnikov 2004).Below we redescribe this insufficiently known widespread species and provide data on its distribution and bionomics.
Body light to dark brownish; head black, pronotum dark brown to brown; elytra brownish with hind angles paler; abdomen dark brown with posterior margins distinctly lighter; hind legs yellowish, antennae, maxillary and labial palps darker, body glossy (Figs 12A; 13A).
Head slightly wider than long HL/HW: 0.9-1.1 (1.0), eyes large and convex; temples distinctly shorter than eyes (ratio 0.2-0.3(0.3); with shallow, but dense transverse microsculpture; punctation: one puncture at anterior margin near antennal pit, anterior frontal puncture at posterior margin of antennal pit, posterior frontal and temporal punctures closer to posterior margin of eye than to posterior margin of head; vertical punctures (ca.[1][2] closer to neck than to posterior margin of eye.
Pronotum slightly wider than long or transverse PL/ PW: 0.9-1.1 (1.0), widest at its posterior half, vaguely narrowing anteriad, wider and longer than head; hind angles rounded barely distinct; dorsal rows each with three punctures; sublateral rows each with two punctures; microsculpture with shallow hardly visible transverse waves.
Scutellum punctate with microsculpture distinctly denser as on pronotum.
Abdomen: punctation fine and dense; interspaces with minute irregularities; posterior margin of tergite VII with palisade fringe.
Male: protarsi with tarsomers 1-4 dilated stronger than in females.Comparison.Among other Raphirus that occurs in Middle Asia, Q. cohaesus is most similar to Q. pseudonigriceps from which it can be easily distinguished by the presence of an apical seam of palisade fringe VII and normally developed elytra, as well as by the characters of the aedeagus.
Distribution.Quedius cohaesus was described from "Turcmenia" which is not necessarily Turkmenistan in the modern sense, but certainly some locality in Middle Asia (Eppelsheim, 1888).Based on the literature (Table 1) and material examined here, Q. cohaesus is known from Iran (material not recorded here), Turkmenistan, Tajikistan and Afghanistan (most of the material not recorded here).

Quedius (Raphirus) imitator Luze, 1904
Comments on the lectotype designation.In the original description of Q. imitator, Luze (1904) did not specify the number of syntypes but provided geographical data that indicated multiple syntypes collected in the localities "Seravschan, Putchin-Pass, Darch, Obburden, Urmitan, Kumar; Jagnob, Varsaut; Iskander-Kul, Iskander-Darja" (approximate coordinates as we interpret these localities are given in the Table .2).Also, the syntype series must have included both sexes because male characters were specified separately in the description.In the ZIN collection we found 11 specimens from several localities along Zeravchan and Iskander Darya Rivers matching those in the original description (for details see 'Material examined' above).Based on that and additional information from the specimen labels, there is no doubt that they are syntypes.Earlier they were identified by Boháč (1988) as Q. imitator without recognizing them as syntypes.In order to fix the identity of the species, we designate here one male syntype with more preciselocality "Seravschan Darch Glasunov 1892" (Darg, Sughd Distr.) as the lectotype.Bernhauer (1905) considered Q. imitator as a "rough form" of Q. oblitteratus (now synonym of Q. humeralis Stephens, 1832).Gridelli (1924) not seeing types or any other material of Q. imitator was not sure about the status of this species.Based on the non-type material, Coiffait (1967Coiffait ( , 1978) ) illustrated its aedeagus for the first time that here is shown to be the correct species interpretation.Boháč (1988) provided new records for the species from Uzbekistan and Turkmenistan that are reliable because he examined syntypes.
Comments on the new synonym.Coiffait (1969) described Quedius tschinganensis (Fig. 14) from Uzbekistan and separated it from Q. imitator by darker body coloration, antennal segment 3 longer than 2, presence of 'lateral' puncture on pronotum, denser punctation of the elytra and more elongated median lobe.Additionally Coiffait (1977) described Q. tschinganensis gracilicornis, a variety of Q. tschinganensis from Tajikistan based on some differences in coloration of the body and the proportions of antennae.Later (1977), he replaced the preoccupied name gracilicornis by the new name debilicornis.Both are unavailable names due to ICZN Article 15.2 as already noted in Herman (2001).
Our examination of the material from Middle Asia, including types, showed continuous variability in the external morphology and aedeagus that connects the states of Q. imitator and Q. tschinganensis.The shape of the paramere varies from the state with narrow and sharp apex with lesser number of sensory peg setae arranged in regular rows away from the apex, to the state with obtuse apex and with more sensory peg setae arranged denser and closer to the apex (Figs 14C,G;15).Shape of the median lobe is more stable and varies only slightly in length and degree of sharpness of its apex (Figs 14B,F;15).Mapping of this variability across the species distribution does not show any geographical patterns.Externally all specimens including females, also show no traits that would correspond to variants different in the shape of the paramere.Thus we place Q. tschinganensis Coiffait, 1969 in synonymy with Q. imitator Luze, 1904.Our study of the type specimen of Q. tschinganensis debilicornis also shows it to be conspecifc with Q. imitator.
Comments on taxonomy, distribution and bionomics.Quedius imitator can be diagnosed by the following character combination: body dark brown with darker head and abdomen; elytra with slightly yellowish anterior angles; antennae usually pale; scutellum without setiferous punctation; aedeagus with ventral tooth of median lobe located remotely from its apex, with median lobe and paramere very narrow, apex of paramere obtusely sharpened and sensory peg setae arranged in two regular rows convergent to each other.Among other Raphirus that occur in Middle Asia, Q. imitator is most similar to Q. cohaesus from which it can be easily distinguished by the mentioned diagnostic characters of the aedeagus.
Based on the examined material and literature (Table 1), Q. imitator is widely distributed in all countries of Middle Asia (Fig. 15).According to the label data of the examined material, Q. imitator inhabits ground based debris and leaf litter of mainly deciduous forests along rivers and streams at various elevations, up to 3000 m.Also it can be found in dung or under stones.

Figs 3E, 17
Quedius dzambulensis Coiffait, 1967, syn.n. (Fig. 16) Quedius novus Eppelsheim, 1892, 331 (original description); Gridelli 1925, 125;Wüsthoff 1938    "Taschkent Leder.Reitter" indicating that they are likely to be syntypes as well.Examination of all syntypes confirms that previous authors correctly interpreted this species.In order to fix its identity, here we designate one male syntype from NMW as the lectotype.Due to the intraspecific variability (Figs 16,17) and resulting new synonymy Q. novus = Q.dzhambulensis (see below), we chose a syntype for lectotypification which has a more narrow longitudinal row of sensory peg setae on the paramere, best matching Coiffait's (1967) illustration for Q. novus.
Comments on the new synonym.The aedeagus of Q. novus was first illustrated by Wüsthoff (1938) based on non-type material.Coiffait (1963Coiffait ( , 1970Coiffait ( , 1978) ) redescribed the species, also illustrated the aedeagus and provided more records for Q. novus from Uzbekistan.Our examination of syntypes proved both Wüsthoff's and Coiffait's interpretation of this species was correct.Also Coiffait (1967) described Q. dzambulensis from Dzambul (Kazakhstan) (Fig. 16), a species which seemed to be very similar to Q. novus even from the illustrations of the aedeagi for both species.
Later, Boháč (1988) examined material from the ZIN collection and provided new records from Uzbekistan, Tajikistan, Kyrgyzstan of Q. dzambulensis and only one record from Uzbekistan for Q. novus.He also stated that Q. novus is very closely related to Q. dzambulensis with which it can be easily confused.We checked all material from ZIN studied by Boháč (1988) and found that, without knowing it, the only specimens he identified as Q. no- Comments on taxonomy and lectotype designation.In the original description of Q. novus, Eppelsheim (1892) stated that he had examined numerous specimens from Tashkent and one from Margelan [Margilan in Uzbekistan].He also stated in the introduction of that study that he received material from 'Turkestan' from multiple collections of Hauser, Staudinger and Reitter.In particular, he mentioned that the material from Tashkent from Reitter's collection was collected by Leder.In NMW altogether we found 11 conspecific specimens, all originally from Eppelsheim's collection (with printed label "c.Epplsh.Steind."),whose morphology and label data match with the original description.We consider all of them to be syntypes.Of them, 8 specimens (on 5 pins) were earlier supplied with the curatorial printed red labels "types"; only two specimens have what we consider Eppelsheim's hand written labels "novus Epp.Taschkent Leder." and one specimen having "novus Epp.Deutsch.ent.Zeit.1892.P. 331" label in a different handwriting probably attached by somebody later, after the species description was published.Also in the ZIN collection we found two more specimens conspecific with the syntypes at NMW and with the label vus were the syntypes of that species.All other specimens he identified as Q. dzambulensis.
Our examination of a broader sample from Middle Asia, including types of both species, showed continuous variability in the structure of the aedeagus connecting the state of Q. novus with the state of Q. dzambulensis.Sensory peg setae on the paramere vary in arrangement, from denser (as in Coiffait's illustration for Q. novus) to sparser (as in Coiffait's illustration for Q. dzambulenisis) witin a longitudinal group (Fig. 17).The mentioned variability has no geographic pattern.Therefore, we consider Q. dzambulensis Coiffait, 1967 to be a junior synonym of Q. novus Epp.
Diagnosis.Body dark brown; elytra with lighter colored humeri and shallow micropunctation between punctures; antennae slightly paler; scutellum without setiferous punctation.(Figs 3E, 16A) Aedeagus (Figs 16B, C; 17): ventral tooth of median lobe located remotely from its apex; median lobe and paramere very broad (Figs 16B,17); apex of paramere obtusely pointed and sensory peg setae arranged in long wide band in the middle of paramere (Figs 16C,17).Quedius novus can be easily distinguished from the similar Middle Asian species Quedius umbrinus by the coloration and micropunctation of elytra and also by the mentioned above aedeagal characters.
Distribution.Based on the literature data (Table 1) that proved to be reliable for this species and the material examined here, Q. novus is widely distributed in Middle Asia and appears the most common in southern Kazakhstan, eastern Uzbekistan, western Kyrgyzstan and northeastern Tajikistan (Fig. 17).
Bionomics.Quedius novus prefers various wet ground based plant debris or moss usually near water bodies.It seems to occur both in forested and open habitats, up to 2700 m.Occasionally it was also found under stones and in dung.

Quedius (Raphirus) umbrinus Erichson, 1839
Fig. 4B Quedius umbrinus: Herman 2001, 3287 ( Comments on taxonomy, distribution and bionomics.Among all Middle Asian Raphirus, Q. umbrinus is most similar to Q. novus from which it can be distinguished by the structure of aedeagus: median lobe with distinct ventral tooth near its apex and apical portion slightly curved dorso-ventrally (in lateral view); paramere (underside) with sensory peg setae arranged in wide lateral rows merging at parameral anterior margin.
Comments on taxonomy, distribution and bionomics.Externally and by the structure of the aedeagus, a single male specimen from Kyrgyzstan (Fig. 18) examined here seems to be a new species from the coloratus-group, recently revised by Assing (2017).Quedius coloratus and allied species forming that group are regional Mediterranean endemics with allopatric distributions, altogether extending from Greece, through Turkey to Jordan.Our specimen differs from all known species of the coloratus-group in the structure of aedeagus (sharp apex of median lobe, subapical tooth located much further away form the apex, peg setae of the paramere less distinctly arranged in longitudinal rows and situated more medially (Fig. 18B, C).
This specimen from the high elevations of Kyrgyz Alatoo, far from the Mediterranean region, is a noteworthy finding for the coloratus-group.More material is needed for a clearer understanding of its identity and formal description.Comments on taxonomy, lectotype designation and new synonymy.In the original description of Q. hauseri, Bernhauer (1918) did not specify the type material but he mentioned localities "Baldschuan [Baljuvon], 924 m, Sary-pul, 1482 m" [Tadjikistan: Karateghin Mts.] (Fig. 19E) and "Ost-Buchara: Tschitschantan, Karatag und Repetek, vor." [Tadjikistan: Vorukh jamoat, according to Frisch 2015] where his material came from.Also it is clear from the original description that he studied both sexes.All this suggests multiple syntypes.Bernhauer (1918) compared Q. hauseri with Q. boops and Q. acuminatus.Wüsthoff (1938) illustrated the structure of the aedeagus for Q. hauseri for the first time based on some material "aus Buchara" [from Buchara].Next, the aedeagus for Q. hauseri was illustrated by Coiffait (1978), also based on some non-type material.

Quedius (Raphirus) hauseri Bernhauer, 1918
We were able to study a male specimen from the FMNH (for details see above) which is clearly a syntype and which we designate as the lectotype to fix the identity of that species.Our examination of the type specimen of Q. hauseri confirms the correct identification of this species by both Wüsthoff (1938) and Coiffait (1978).
In the same paper, Bernhauer (1918) described Quedius peneckei as a brachypterous variation of Q. hauseri from 'Tien-Shan, Przewalsk, Karakoltal' [now Karakol, Issyk-Kul region, Kyrgyzstan], also not specifying either a number or sex of the material he studied.He only stated that Q. peneckei was similar to Q. fulvicollis from which it could be distinguished by the elongate pronotum and more densely punctured abdomen.Gridelli (1924) and Coiffait (1978) also considered Q. peneckei as a variation of Q. hauseri.Of them, Gridelli (1924) stated that he had studied the type material but without details on sex or number of specimens.In catalogs Q. peneckei is given as a variation (Scheerpeltz, 1933;Hermann, 2001) or synonym (Schülke & Smetana, 2015) of Q. hauseri.There was not a single illustration of Q. peneckei ever published.We were able to study one female specimen from the FMNH which is clearly a syntype of Q. peneckei.It is conspecific with Q. hauseri and does not look to be distinctly brachypterous.Based on that and the fact that there is only one species of this type in Middle Asia, Diagnosis.Head and abdomen usually black, pronotum, elytra and appendages pale-brown to brown; scutellum punctate (Figs 19A,20A,21A).Aedeagus (in lateral view) (Figs 19B,21B): ventral contour of median lobe apically and basally from subapical tooth form one line, so that the tooth is protruding and median lobe apically from that does not look like an axe blade.Among all Middle Asian Raphirus, only Q. hauseri and Q. fulvicollis (see below) have the punctate scutellum.Quedius hauseri differs from Q. fulvicollis by the shape of the paramere (Figs 19C,21C,23C,respectively).
Distribution.Quedius hauseri is common and widely distributed in Middle Asia where it occurs from south-which is rather common and widespread (Fig. 22), we place Q. peneckei in synonymy with Q. hauseri.Coiffait (1969) described Q. ouzbekiscus from Uzbekistan based on the male holotype (Fig. 21) and 40 paratypes (4 males and 36 females).He considered it similar to the species from the boops-group and stated that Q. ouzbekiscus can be distinguished from other members of the group by the structure of aedeagus and proportions of the body.Also he noticed that Q. ouzbekiscus is especially similar to Q. fulvicollis.Our examination of the type material of Q. ouzbekiscus reveals that this species is conspecific with Q. hauseri and therefore we place the former in synonymy with the latter.eastern Kazakstan (southern border through Dzhungaskiy Alatau) to southern Tajikistan (Pamir Mountains, Schugnan) (Fig. 22).It was also recorded from Afganistan (Schülke and Smetana, 2015).
Bionomics.Based on the material examined here Q. hauseri usually inhabits various humid ground based plant debris or moss near water bodies.It occurs both in forested and open habitats.It also can be found under stones, bark and in dung, mostly at the medium to high elevations up to 3300 m.Material examined.One of the male paratypes of Q. ouzbekiscus (new synonym of Q. hauseri, see above), for details see material examined for Q. hauseri and Fig. 23 Comments on taxonomy, distribution and bionomics.One of the male paratypes of Q. ouzbekiscus (new synonym of Q. hauseri) was in fact a different species that we tentatively identify as Q. fulvicollis.It can be easily distinguished from Q. hauseri by the shape of the paramere (compare Fig. 23C and Figs 19C,21C,respectively).Quedius fulvicollis is considered a widely distributed Holarctic species, in Asia confined to Siberia and Russian Far East (Schülke and Smetana, 2015).The specimen from Chatkal Mountains in Uzbekistan examined here would be a distinct southernmost record for this species in the Palaearctic region and the first record for Middle Asia.In this respect it is noteworthy that it comes from ca. 1500 m of elevation.Also it is remarkable that this specimen from Middle Asia stands out from the variability range of Q. fulvicollis by the very narrow middle portion of the paramere and shorter and more irregular rows of peg setae.It well may be that our specimen represents a species new to science.Given the poorly studied variation of Q. fulvicollis, which itself maybe a complex of species and very limited material from Middle Asia, a decision on this matter is pending further study.

Quedius (Raphirus) scintillans Gravenhorst, 1806
Comments on taxonomy, distribution and bionomics.Quedius scintillans is widely distributed in Europe, Western and Middle Asia, and its diagnostic characters, distribution and biology were recently summarized in Assing and Schülke (2012).In Middle Asia, from the newly examined material here, the species is recorded in southern and eastern Turkmenistan and southwestern Tajikistan for the first time.
From all Middle Asian Raphirus species it can be easily distinguished by the presence of two additional punctures between anterior frontal punctures on the head.
Quedius scintillans prefers various wet ground-based debris mostly in lowland forests or open landscapes.In the mountains it can be found up to 1300 m elevation.

Discussion
This revision is the first focused summary on Quedius of Middle Asia.It clarifies the taxonomy of many poorly or very poorly known species such as Q. (s.str.) subunicolor, Q. (M.) capitalis, Q. (M.) fusicornis, Q. (M.) solskyi and Q. (R.) cohaesus, and it records from Middle Asia a few widely distributed species such as Q. (s.str.) fuliginosus, Q. (s.str.) sundukovi and Q. (R.) pseudonigriceps for the first time.It shows how confusing and incomplete the taxonomy was of the species that constitute the core of this fauna.In the course of this revision (including Salnitska and Solodovnikov 2018) the rate of new species dicovery was negligible compared to the rate of revealed misidentifications and synonymies.Many "endemic" species described from various regions of Middle Asia, mainly by H. Coiffait, turned out to be synonyms of the species described from this region at the border between XIX and XX centuries (Table 1).These species, with the newly examined material, expectedly turned out to be more widespread than they were previously thought.Several species, especially in the subgenus Microsaurus, remain very poorly known (e.g., Q. (M.) bucharensis, Q. (M.) fusicornis, Q. (M.) solskyi, Q. (M.) koltzei and Q. (M.) tajikiscus).Here they are represented by fragmentary, poorly georeferenced type material (often females only) and, at most, a few additional specimens.For Q. (s.str.) subunicolor and Q. (s.str.) sundukovi and Q. (M.) koltzei, new distributional records from Middle Asia change our idea of their distribution patterns and calls for their more thorough exploration.In general this revision made it obvious that, apart from a handful of species such as Q. (s.str.) fuliginosus, Q. (s.str.) vicinus, Q. (M.) ochripennis, Q. (R.) hauseri, Q. (R.) imitator, Q. (R.) limbatus, Q. (R.) novus, Q. (R.) pseudonigriceps and Q. (R.) scintillans, well represented in the examined material, the Middle Asian species are known from very scarce sampling.Because Middle Asia is mainly a warm and arid region that is not well suited to such a distinctly temperate and mesophilous genus, the fauna of Quedius is relatively poor.For example, the Quedius fauna of Denmark, a much smaller, geographically uniform and flat area, contains 41 species (http://danbiller.dk) as opposed to 28 species recorded from Middle Asia.It is not expected that the Middle Asian Quedius fauna will significantly grow with more explorations.But some increase of this number is likely, due to widespread species to be found there and new species for science to be discovered, especially from the mountain areas of Middle Asia.We hope that our work will encourage further field exploration of this diverse region by using collecting techniques targeting Staphylinidae, especially sifting.

Figure 6 .
Figure 6.Distribution of sister species Quedius altaicus (Altai Mts. in Kazakhstan and Russia) and Q. subunicolor (Northern Europe).Dots indicate material examined in this paper, colored areas summarise literature data.Black dots indicate type localities. Fig.2F Fig. 3A

Figure 7 .
Figure 7. Quedius capitalis, syntypes, males.A, B, habitus; C, D, aedeagus of the syntype in the photo A. E (of the syntype in the photo A), F (of the syntype in the photo B), labels.C, median lobe, lateral view; D, paramere, underside.Scale bars: A, B = 1 mm; C = 0.5 mm.

Figure 11 .
Figure 11.Quedius pseudonigriceps, distribution, median lobe of the aedeagus laterally, and variability of the paramere (as examples specimens from various localities numbered respectively on the map).Scale bars: 0.5 mm.
Aedeagus (Figs 12C, D; 13B, C): Median lobe parallel-sided with moderately acute apex, tooth situated close to its apex (Figs 12C, 13B).Paramere parallel-sided, slightly narrowing basad; its apex almost reaching apex of median lobe; sensory peg setae arranged in two irregular and wide longitudinal rows along each lateral margin of apical portion extending over pairs of lateral setae below apex (Figs 12D, 13C).

Figure 17 .
Figure 17.Quedius novus, distribution, median lobe of the aedeagus laterally, and variability of the paramere (as examples specimens from various localities numbered respectively on the map).Scale bars: 0.5 mm.

Figure 22 .
Figure 22.Quedius hauseri, distribution and variability of the paramere (as an example specimens from one locality, indicated by black dot).Scale bar: 1 mm.

Table 1 .
Alphabetical list of Quedius species recorded for Middle Asia, with new synonyms.Boldfaced species are those confirmed by material in our study; species in regular font not given in square brackets are those known from literature only, presumably absent in Middle Asia; species in regular font and given in square brackets are those previously recorded for the region in literature based on misidentifications and here excluded from the fauna.