New species and records of Quedius rove beetles (Coleoptera, Staphylinidae, Staphylininae) from Middle Asia

. This paper is devoted to the fauna of Quedius Stephens, 1829 of Middle Asia. It provides descriptions of two species new to science Quedius ( Distichalius ) gissaricus sp. nov. from Tajikistan and Q. ( Raphirus ) viator sp. nov. from Kyrgyzstan. Two more species are reported as potentially new to science, but not formally described: Q. ( Microsaurus ) sp. from the mutilatus group from Tarbagatay Mountains because it is known only from damaged specimens; Q. ( Distichalius ) sp. aff. gissaricus , known from a single male found sympatrically with Q. ( D. ) gissaricus sp. nov., remains undescribed because of the possibility of teratology. Earlier records of Q. umbrinus from Middle Asia are considered doubtful or assigned to Q. pseudoumbrinus . Taxonomic notes on the identity of that species as well as Q. cohaesus and Q. fulvicollis in Middle Asia are provided. Among new distribution records summarized for several species, Q. ( Velleius ) dilatatus , Q. ( Microsaurus ) tenellus and Q. ( Raphirus ) jenisseensis are recorded from the territory of Middle Asia for the first time .


Introduction
Three years ago, we published a revision of the poorly known fauna of Quedius Stephens, 1829 of Middle Asia (Salnitska & Solodovnikov 2018a).Back then, in the material from the Zoological Institute in St Petersburg (ZIN), which was one of the main data providers for our revision, we studied some specimens collected by the recently deceased colleague Vitaliy Alexandrovich Kastcheev (1953Kastcheev ( -2012)).V.A. Kastcheev was a well-known entomologist from Kazakhstan who collected extensive material of Staphylinidae Latreille, 1802 from Middle Asia, authored about 130 scientific publications and described 43 species new to science.The main part of his collection of Staphylinidae was acquired by ZIN and made available to us for study after the publication of our revision.Our study of that collection, supplemented by several specimens from the general collection of ZIN and the private collection of Alexander Ryvkin (Moscow) resulted in this paper.
The newly examined material revealed four new species, all represented by singletons requiring additional material to learn more about these species.Nevertheless, we decided to describe two of them, Q. gissaricus sp.nov.and Q. viator sp.nov., because both are distinctive new species.
Apart from the new species, this paper takes into account our recent molecular-based study of the Q. umbrinus complex (Salnitska & Solodovnikov 2021) and provides new distributional records for a number of species in Middle Asia as well as records of some widespread species from Middle Asia for the first time.A few of the new distributional records that provide notable range expansions or are of other significant novelty are included with our respective comments.Finally, the newly examined material provided additional records for species of the subgenus Quedius s. str.: for Q. altaicus Korge, 1962 andQ. fuliginosus (Gravenhorst, 1802) which were previously known from Middle Asia (Salnitska & Solodovnikov 2018a); and for Q. balticus Korge, 1960 recorded for the first time in this region.All these records are provided in detail in the revision of this subgenus (Hansen et al. 2022 in press).

Material and methods
This paper is based on material from the Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia (ZIN) and the private collection of Alexander Ryvkin, Moscow, Russia (cRyv).The holotypes of the two new species are deposited at ZIN. Specimens were examined with Lomo MSP-2 ver. 2 and MBS-2 dissecting scopes.Measurements were taken at 4 × magnification using an ocular micrometer and are given in mm.
Abbreviations are as follows: EL = length of elytra (from humerus to the most distal part of the posterior margin) EW = width of elytra (maximum, with elytra closed along suture) FB = forebody (from base of labrum to the most distal part of posterior margin of elytra) 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) TL = total length of the body (from base of labrum to apex of abdomen, with extended abdomen) Photographs of the habitus and genitalia were obtained with the following equipment: Canon EOS 5D Mark IV camera, Canon MP-E 65 mm lens, CANON Macro Twin Lite MT-26EX-RT flashand Cognisys StackShot 3X rail macro photo tripod.They were edited in Photoshop CS5.Illustrations of the aedeagi were made with Adobe Illustrator CS5.1 based on the photos taken from soft preparations of these structures in glycerine after dissecting, maceration in 10% KOH, and rinsing in distilled water.All dissected aedeagi are retained in glycerine in genitalia microvials pinned under their respective specimens.
All labels of the type specimens are cited verbatim in the 'Material examined' sections.
The distribution map in Fig. 2 was prepared using QGIS 2.12.0 free software by schematic free hand generalization of the species group distributions based on literature and by plotting geographic coordinates for the new species.

Remarks
In our revision (Salnitska & Solodovnikov 2018a), we considered Q. cohaesus and Q. pseudonigriceps Reitter, 1909 as separate species distinguished by a subtle, hard to express, difference in the aedeagus and by the fact that in Middle Asia Q. cohaesus has the apical seam of the palisade fringe on tergite VII and normally developed elytra, while Q. pseudonigriceps was brachypterous and without the palisade fringe.Our concept implied that Q. pseudonigriceps is a species widely distributed in southern Europe and the south-western part of Asia, brachypterous in Middle Asia.For Q. cohaesus it implied that it was a fully winged species distributed only in the Middle East and Middle Asia: Iran, Turkmenistan, Tajikistan and Afghanistan.Assing (2019) synonymized Q. pseudonigriceps with Q. cohaesus because he (1) considered the concept of a macropterous species (Q.cohaesus) replacing the macropterous form of another, wing dimorphic and very similar species (Q.pseudonigriceps) implausible; and (2) he did not find in his large collection the discrete differences between the two species mentioned in Salnitska & Solodovnikov (2018a).Thus, Assing (2019) assumed that Q. cohaesus is distributed both west and east of the Caspian Sea where it is wing-dimorphic in Middle Asia and wing-monomorphic everywhere else.Here, we follow Assing (2019) since we agree that it may be impossible to distinguish the two 'species' using the structure of the aedeagus, and that Q. cohaesus (= Q. pseudonigriceps) could be wing-dimorphic.However, the case needs further investigation with more data and a molecular approach.It is worth mentioning that in the new material from southern Kazakhstan and northern Kyrgyzstan reported here, there are only wingless specimens that we would have formerly identified as Q. pseudonigriceps and that Assing (2019) would identify as the wingless form of Q. cohaesus (= Q. pseudonigriceps).

Remarks
In Middle Asia, Q. fulvicollis was hitherto known only from a male specimen from the Chatkal Mountain Range in Uzbekistan (Salnitska & Solodovnikov 2018a).The identification of this specimen was tentative due to the form of its paramere (with very narrow middle portion and shorter and more irregular rows of peg setae), which was somewhat different from other Q. fulvicollis.Also, the locality of that specimen was rather remote from the main distribution area of the species.Based on the material examined here, we provide new records of Q. fulvicollis from Kazakhstan and Kyrgyzstan.All examined specimens can be identified as Q. fulvicollis based on the structure of the aedeagus.Apparently, Q. fulvicollis is a boreo-montane species confined to the northern part of the Holarctic and to the mountains in the south.Nevertheless, considering the slightly different shape of the paramere in the mentioned specimen from Uzbekistan and the isolated records of Q. fulvicollis from Middle Asia with respect to the main broad latitudinal range of this species, this case needs a more thorough molecular study.

Remarks
Quedius novus is widely distributed in Middle Asia, where it appears most commonly in southern Kazakhstan, eastern Uzbekistan, western Kyrgyzstan and northeastern Tajikistan (Salnitska & Solodovnikov 2018a).Here, we provide the first record of Q. novus from Turkmenistan.

Remarks
According to the molecular data, partially supported by the shape of the paramere (Salnitska & Solodovnikov 2021), the former single species Q. umbrinus Erichson, 1839 (for example, sensu Salnitska & Solodovnikov 2018a) is divided into three species, Q. volkeri Salnitska & Solodovnikov, 2021, Q. umbrinus and Q. pseudoumbrinus.The first species is a local endemic of the Western Caucasus, while Q. umbrinus and Q. pseudoumbrinus are widespread West Palaearctic species with largely overlapping distributions.Unfortunately, neither the material of the former 'Q.umbrinus' from Middle Asia reported in Salnitska & Solodovnikov (2018a), nor the material from this region used in Salnitska & Solodovnikov (2021) are of DNA-grade.Among the material from Middle Asia studied in Salnitska & Solodovnikov (2021), only the male specimens from Chilik River of Kungey Alatau in Kazakhstan can be identified as Q. pseudoumbrinus based on the more parallel-sided, not apically rhomboid paramere.The shape of the paramere in males from the Dzhungar Mountains in Kazakhstan and from N of Issyk Kul Lake in Kyrgyzstan in Salnitska & Solodovnikov (2021) does not allow their confident identification as either Q. umbrinus or Q. pseudoumbrinus.Based on the shape of the paramere, the material reported as 'Q.umbrinus' in Salnitska & Solodovnikov (2018a) is likely to be Q.pseudoumbrinus (Salnitska & Solodovnikov 2021), though this needs to be tested with molecular data.It is noteworthy that all material of 'Q.umbrinus' from Middle Asia from both of our abovementioned studies with precise bionomic data was collected at higher elevations in the mountains.Thus, the Middle Asian populations of the Q. umbrinus complex may be somewhat isolated from the main ranges of Q. umbrinus and Q. pseudoumbrinus.To properly shed light on the identity of 'Q.umbrinus' in Middle Asia in view of the data in Salnitska & Solodovnikov (2021), DNA-grade material is needed.

Remarks
Based on the newly examined material, Q. dilatatus is recorded for the first time from Middle Asia (Kazakhstan).It is a widely distributed Palaearctic species that had already been recorded from several provinces of Russia near the border with Kazakhstan (Salnitska & Solodovnikov 2019).This is the first record of Q. dilatatus from the Altai Mountains.

Diagnosis
Among other members of Distichalius Casey, 1915, the new species can be distinguished by the following combination of characters: body relatively small, head with interocular punctures (red arrows in Fig. 1A), elytra with rough and very sparse punctation arranged in irregular rows; median lobe of symmetrical aedeagus simple, without apical lobes, with distinct apex; paramere with several sensory peg setae apically arranged in short rows along parameral margins with apical setae.

Etymology
The species name 'gissaricus' (latinized adjective) is derived from the mountain range where this specimen was collected.
Colouration.Body black (Fig. 1A) with reddish appendages; head and pronotum completely black; elytra and abdomen black with paler margins.
Head.About as wide as long (HL/HW: 0.97), surface with microsculpture of transverse waves on disk and between eyes.Eyes large, moderately convex, temples 1.5 times as long as longitudinal diameter of eye.Head chaetotaxy as in Fig. 1A.Anterior frontal puncture touching anteriomedian margin of eye, with barely visible additional interocular punctures between anterior frontal punctures (red arrows in Fig. 1A); posterior frontal puncture situated closer to posterior margin of eye than to nuchal ridge, pair of smaller basal punctures situated close to nuchal ridge; each temple with two temporal punctures, posterior one situated halfway between posterior margin of eye and nuchal ridge; temples additionally with some fine setiferous punctures bearing short yellowish setae.Antennae moderately long, first segment shorter than second and third segments together; third distinctly longer than second, fourth and fifth slightly longer than wide, sixth to tenth wider than long, gradually increasing in width towards apex of antenna, last fusiform segment distinctly longer than penultimate segment.
Pronotum.Barely wider than long (PL/PW: 0.96), widest at about middle; with nearly parallel sides at middle, posterior lateral portions barely explanate.Disk of pronotum with two dorsal rows of punctures, each with three punctures, third (basalmost) puncture situated behind the level of large lateral puncture; sublateral area with three punctures (one additional between dorsal and sublateral row), posterior two of which situated on one line slightly between level of basalmost puncture of dorsal row; basal margin with row of three variably sized setae on each side; microsculpture of transverse waves, sparser than on head.
Abdomen.Punctated more densely than elytra, punctation becoming sparser toward apex; tergite II with very fine and distinct punctures; posterior margin of tergite VII with whitish palisade fringe.

Male
First four protarsomeres markedly dilated, each densely covered with modified pale setae ventrally; second segment as wide as, or slightly narrower than apex of tibia.Sternite VIII with two long setae on each side; apical margin with weak triangular medio-apical emargination (Fig. 1E).

Female
Unknown.

Remarks
In addition to the holotype of Q. gissaricus sp.nov., there was another specimen of Distichalius collected at the same locality, but some days earlier (for details see Q. (D.) sp.aff.gissaricus below).That specimen differs from the holotype of Q. gissaricus sp.nov.by the distinctly paler pronotum, elytra and appendages, lacking interocular punctures between frontal punctures, less regular punctation of elytra, tergite II with fine and shallow punctation, and median lobe of the aedeagus with smaller subapical tooth located considerably further away from the apex.Given the co-occurrence of the two specimens and their subtle difference from each other, they may be considered either as variants of one morphologically variable species, or one of them could be an aberrant specimen, or they may represent closely related sympatric species.Since a firm conclusion is impossible until more material is examined, we tentatively identify the second specimen as a separate species and list it below.

Ecology
Quedius gissaricus sp.nov. is hitherto known only from the Gissar Mountain Range in Tajikistan where it was collected at an elevation of 2500 m (Fig. 2).

Comparison
Without a complete revision of species currently in Distichalius it is impossible to place Q. gissaricus sp.nov. in a species group.It differs from other smaller Western Palaearctic Distichalius (e.g., Q. punctatellus (Heer, 1839) and allied ones with which it shares the sparse punctuation of the elytra arranged in rows) by the presence of interocular setae on the head and by a smaller number of sensory peg setae on the paramere, which do not extend behind the apical setae towards the parameral base.As can be judged from Smetana (2017), it is distinct from any of the Chinese species at least as follows: it differs from the Q. ladas, Q. bipictus and Q. daedalus groups by a sparse irregular punctation of the elytra which form rows; from the Q. annectens group by the aedeagus without apical lobes; from the Q. laetipictus group at least by a uniform dark body colouration; from the Q. regularis group by a pointed, not widely arcuate apex of the aedeagal median lobe.Our new species differs from all Far Eastern and Japanese species by a smaller body size and the shape of the aedeagus.

Diagnosis
Among other species of Raphirus Stephens, 1829, Q. viator sp.nov. is most closely related to Quedius coloratus Fauvel, 1875 and all allied species, from which it can easily be distinguished by the structure of the aedeagus, namely by the wider apical portion of the median lobe with a sharper apex, rows of peg setae converging with each other in the middle portion (not parallel-sided), and the apex of the paramere almost reaching the apex of the median lobe (Fig. 3B-D) (for more details see section 'Comparison').

Etymology
The name is the Latin noun 'viator' (= 'traveller') that refers to the fact that the newly described species, presumably closely related to a group of Mediterranean species, appears far away, in Middle Asia (Fig. 2).Colouration.Body mainly dark brown (Fig. 3A) with paler appendages; head piceous-black, abdomen dark brown, pronotum and elytra brownish with paler lateral margins; antennae and legs uniformly light brown.

Holotype
Head.Slightly transverse (HL/HW: 0.90), surface with microsculpture of transverse waves on disk and isodiametric between eyes.Eyes moderately large and convex, 1.87 times as long as temples.Anterior frontal puncture situated at inner margin of eye; fine posterior frontal puncture situated closer to posterior margin of eye than to nuchal ridge, one basal puncture situated close to nuchal ridge; each temple with one temporal puncture; temples additionally with some shallow setiferous punctures bearing short yellowish setae.Antennae moderately long, first segment slightly shorter than second and third segments together; third distinctly longer than second, fourth to tenth longer than wide, gradually increasing in width towards apex of antenna, last segment distinctly longer than preceding segments.
Pronotum.Transverse, distinctly wider than long (PL/PW: 0.88), widely rounded basally, widest at about middle, posterior lateral portions vaguely explanate.Disk of pronotum with two dorsal rows of punctures, each with three punctures, third (basalmost) puncture situated far behind the level of large lateral puncture; sublateral area with two punctures at right side and without punctures at left side; basal margin with row of 2-3 setae on each side; microsculpture of transverse waves as on head.

Male
First four protarsomeres markedly dilated, each densely covered with modified brown setae; second protarsomere as wide as, or slightly narrower than apex of tibia.Sternite VIII with a long seta on each side; apical margin with distinct triangular medio-apical emargination (Fig. 3E).Sternite IX moderately elongate (Fig. 3G), asymmetrical, basal portion narrow and glabrous, apical portion wider with obtuse almost square apex and finely setose.Tergite X triangular (Fig. 3F), narrowing apicad, with four strong setae at apical margin and with very few smaller setae at apical portion.3B-D): median lobe (in parameral or anteparameral view) symmetrical, with distinct constriction medially, gradually narrowing anteriad, with obtusely pointed apex; (in lateral view) with obtusely pointed and gradually narrowed apex, minute tooth located far basad from apex (Fig. 3D).Paramere (in parameral view) wider than median lobe, symmetrical, gradually narrowing apicad to moderately sharp apex (Fig. 3B); (underside) apical portion with 13-17 peg setae (Fig. 3C) arranged in two longitudinal irregular rows converging with each other closer to parameral midline; (in lateral view) almost reaching apex of median lobe (Fig. 3D).Internal sac without large sclerotized structures.

Female
Unknown.

Distribution and bionomics
Quedius viator sp.nov. is known only from the holotype, which was collected in the Kyrgyz-Alatoo Mountains at an elevation of 3000 m (Fig. 2), most likely under stones (I. A. Belousov, pers. com.).Assuming the phylogenetic affiliation of Q. viator sp.nov.with Q. coloratus and allied species is correct, the geographic disjunction between our new species, and Q. coloratus and allied confined to the Mediterranean is noteworthy.

Comparison
As a large species of Raphirus with moderately large eyes and characteristic structure of the aedeagus, Q. viator sp.nov.appears similar to Q. coloratus and allied species (sensu Assing 2017) and to some extent to Q. boluensis Korge, 1971.The obtusely pointed and slightly curved apex of the median lobe and lanceolate paramere with two irregular longitudinal groups of peg setae situated closer to the parameral midline enforce such a resemblance.We assume that Q. viator sp.nov. is phylogenetically closer to Q. coloratus and allied species rather than to Q. boluensis.An affiliation of Q. viator sp.nov. to Q. coloratus and allied species is additionally supported by the colouration of the body and ecological preference.According to Assing (2017), Q. coloratus and allied species are subterranean, rare in collections and mainly collected by carabidologists from under stones.The single known specimen of Q. viator sp.nov.was also collected in this.Within the complex of species affiliated with Q. coloratus in Assing (2017), Q. viator sp.nov. is most similar to Q. coloratus based on the head chaetotaxy without additional punctures near the posterior frontal puncture and the structure of the aedeagus (for distinguishing characters see section 'Diagnosis').
According to the identification key for species of Raphirus from China (Smetana 2017), Q. viator sp.nov.falls within the Q. kalganensis group and in fact its habitus is very similar to that of Q. wanyan Smetana, 1996 from that group.This group consists of two species currently known only from female specimens.One of them is described from Hebei Province, another from Qinghai Province of China.To prove the monophyly of the Q. kalganensis group and its affinity to Q. viator sp.nov.despite remote distributions, better knowledge of the Asian Quedius fauna, or at least males of Q. kalganensis group are needed.

Remarks
Prior to this study, the northernmost record for the peculiar mutilatus group was presented by three specimens from the Dzhungarian Alatau (Salnitska & Solodovnikov 2018b).However, at that time we considered that record as questionable, possibly based on mislabelling, because of its very distant location from the core distribution area for this group.The specimens examined here come from a locality even further northwards.Thus, it can be argued that the formerly questioned record from Dzhungarian Alatau was in fact not erroneous.These findings show that the distribution of the mutilatus group extends significantly northwards.Unfortunately, as in the case of the material from Dzhungarian Alatau (Salnitska & Solodovnikov 2018b), both male specimens from the Tarbagatay Mountains reported here are damaged by dermestids and their aedeagi are lost.Based on the external morphology, both specimens clearly belong to the Q. mutilatus group, but a more precise identification at the moment is not possible.

Discussion
This is the first supplement to our revision of the fauna of Quedius of Middle Asia (Salnitska & Solodovnikov 2018a).It shows the relative completeness of that revision as far as the fauna of the lower elevations areas is concerned.Despite the fact that quite extensive material was examined here, only three species, Q. dilatatus, Q. tenellus and Q. jenisseensis, are recorded for Middle Asia for the first time.Two new species described here, Q. gissaricus sp.nov.and Q. viator sp.nov., were in fact already mentioned but not described in the revision (Salnitska & Solodovnikov 2018a).Our study of the additional collections confirms that material of Quedius from the higher elevations of the mountains of Middle Asia is biogeographically very interesting, but unfortunately very fragmentary and represented by singletons only.Apart from the new species, the most interesting finds are a considerable extension northwards for the distribution of the mutilatus group and additional data corroborating Q. fulvicollis as a boreo-montane species (or potential species complex) that occurs in Middle Asia.Our understanding of Q. fulvicollis would greatly benefit from molecular data, which are almost absent but highly desirable for Quedius in Middle Asia.A similar situation is that of the Q. umbrinus complex where molecular data provided by Salnitska & Solodovnikov (2021) clarified the previously morphologically blurred species limits, but where, again, the highly interesting Middle Asian material is non-DNA-grade.
As a result of the present study, there are 33 species of Quedius currently known in Middle Asia.This is likely to be close to the real fauna at least for the better-known lowlands.Targeted expeditions aimed at collecting Staphylinidae and fresh material for DNA sequencing would undoubtedly yield more new species, especially from the highlands.As shown here by Q. gissaricus sp.nov.and Q. viator sp.nov., such species may have unexpected biogeographic connections to be explored.