A review of the spider genus Haplodrassus Chamberlin, 1922 in Crimea (Ukraine) and adjacent areas (Araneae, Gnaphosidae)

Abstract Eight species of Haplodrassus are recorded from Crimea: Haplodrassus bohemicus Miller & Buchar, 1977; Haplodrassus dalmatensis (L. Koch, 1866); Haplodrassus isaevi Ponomarev & Tsvetkov, 2006; Haplodrassus minor (O. P.-Cambridge, 1879); Haplodrassus kulczynskii Lohmander, 1942; Haplodrassus pseudosignifer Marusik, Hippa & Koponen, 1996; Haplodrassus signifer (C.L. Koch, 1839) and Haplodrassus umbratilis (L. Koch, 1866). The occurrence of Haplodrassus cognatus (Westring, 1861) in Crimea has not been confirmed. Haplodrassus bohemicus is a new species record for the Crimean fauna. Haplodrassus pseudosignifer is a new species record for Crimea and Ukraine as a whole, with Crimea as the westernmost point of its distribution range. Haplodrassus invalidus is recorded for the first time for the fauna of Azerbaijan, Caucasus and the former Soviet Union. Azerbaijan is the easternmost point of its known distribution range. All Crimean Haplodrassus species have only one peak of activity of adult specimens during the year. In Crimea we found syntopically two closely related species Haplodrassus dalmatensis and Haplodrassus isaevi in two localities (Sudak Distr., 10 km W Sudak, Mezhdurechie Vill., steppe; and Feodosiya Distr., Karadag Nature Reserve, steppes). These species differ in their phenology. The reproductive period of Haplodrassus dalmatensis isin May-July, and that of Haplodrassus isaevi occurs is in October-December. These phenological differences probably represent an additional mechanism of reproductive isolation between the two species. Diagnostic drawings are provided for all mentioned species as well as for Haplodrassus deserticola Schmidt & Krause, 1996 and Haplodrassus pugnans (Simon, 1880).

In the former Soviet Union 21 species have been recorded to date (Mikhailov 1997;Tuneva 2005;Ponomarev and Tsvetkov 2006;Marusik et al. 2007;Ponomarev 2008;Ponomarev et al. 2008), with six of them reported from Crimea (Kovblyuk 2006(Kovblyuk , 2011. While identifying gnaphosid material collected in Crimea during the last few years we recognized two additional species. During our research to identify them we studied Haplodrassus species known from adjacent territories and found one species new to the fauna of Caucasus. Some of the species occurring in Crimea are relatively poorly known, and/or closely resemble widespread species. Therefore, this paper aims to illustrate all of the species found in the Crimea and those from adjacent territories. In addition, we provide data for the distribution and seasonal activity dynamics of adult Haplodrassus specimens in Crimea, a key to all Crimean species.

Material and methods
Microphotographs were made using an SEM Jeol JSM-5200 in the Zoological Museum, University of Turku, Finland. Photographs were taken in dishes of different sizes with paraffin at the bottom. Specimens were photographed using an Olympus Camedia E-520 camera attached to an Olympus SZX16 stereomicroscope at the Zoological Museum, University of Turku. Digital images were montaged using "CombineZM" image stacking software.
Coloration was described from specimens preserved in an ethanol/water solution. Leg segments were measured after detaching them from the cephalothorax. All measurements are in mm: minimum-maximum; a figure in brackets represents the average. Illustrations were made using both reflecting-and transmitted-light microscopes. All scale bars equal 0.1 mm.
The morphological terminology follows Platnick and Shadab (1975) and Levy (2004). In the text we provide references only to the most useful publications, including books and revisions.
All specimens treated in this study are held in the following collections: CPpersonal collection of A.V. Ponomarev The following abbreviations are used in the text: AM, AL, PM, PL -anterior median, anterior lateral, posterior median and posterior lateral eyes; RTA -retrolateral tibial apophysis.

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Terminal apophysis toothed (Fig. 37)  Fovea rectangular (longer than wide) (Fig. 75)  Diagnosis. Male palp with large terminal apophysis, thick embolus, hooked median apophysis and RTA flattened, often shifted dorsally. Epigyne with thick sclerotized lateral pockets and with one anterior hood. Posterior median eyes close together, separated by their radius or less (Platnick and Shadab 1975;Levy 2004). Haplodrassus is most related to Orodrassus Chamberlin, 1922 with three species from the Nearctic, but differs by having a flattened RTA (bifid or laterally expanded in Orodrassus), lacking a median epigynal projection (present in Orodrassus) and by the presence of lateral epigynal sclerites (absent in Orodrassus) (Platnick and Shadab 1975).
In terms of habitus and coloration Haplodrassus resembles only two other genera that occur in Crimea and the eastern Mediterranean: Parasyrisca Schenkel, 1963 and, to a lesser extent, Drassodes Westring, 1851. However, Haplodrassus is easily distinguished by having a large terminal apophysis (absent in the other genera), a flat retrolateral tibial apophysis widened dorsally (conical or flat and tapering in Drassodes and Parasyrisca), a broad embolus (cylindrical or hidden in the other genera), and the presence of heavily sclerotized lateral epigynal pockets (absent in Drassodes and Parasyrisca).
Palp as in Figs 1-6. RTA with a "step" (St) on the promargin, terminal apophysis almost straight, without distinct ridge, embolus slightly twisted and lacks a tooth.
Records from Crimea. Bragina (1984); Kovblyuk (2004aKovblyuk ( ,b, 2006; Kovblyuk et al. (2008). Note. The earlier record of H. cognatus from Crimea was based on specimens of unknown sex and number from Karadag Nature Reserve (Bragina 1984). Haplodrassus cognatus is absent in our material from Crimea, although we have large collections, especially from the Karadag Reserve). It is reasonable to conclude that the earlier records of H. cognatus from Crimea represent a misidentified material. Diagnosis. Haplodrassus cognatus can be distinguished from all other Haplodrassus species by its straight terminal apophysis with a basal tooth in males, and by the shape of the fovea and wide anterior hood (Ah) in females.
Distribution. It has a trans-Palearctic boreo-nemoral range and occurs from France to Hokkaido, north to north Ural and Tomsk, and south to Greece (Marusik et al. 2000;Helsdingen 2010;Platnick 2012 (2012).
Records from Crimea. Apostolov and Onchurov (1998); Onchurov (1998);Mikhailov (2000); Kovblyuk (2004aKovblyuk ( ,b, 2006; Kovblyuk et al. (2008). Diagnosis. Haplodrassus dalmatensis can be easily distinguished from all other congeners by the shape of the terminal apophysis with two tooth-like apical processes and by the strong tooth on the embolus in males, and also by the shape of the epigynal fovea with a peculiar medial septum and converging lateral pockets in females.

Diagnosis. Males of H. invalidus can be distinguished from all other
Haplodrassus species by the peculiar thin embolus with an inner spur-like process, and also the peculiar shape of the RTA, which is not indented and has a claw-like tip (Figs 32-34).
Comments. Haplodrassus invalidus is a new species record for the fauna of Azerbaijan, Caucasus and the former Soviet Union. Azerbaijan is the easternmost point of the known distribution range. Diagnosis. Haplodrassus isaevi is most similar to H. dalmatensis but differs by the shape of the terminal apophysis, which has only one tooth-like process (two tooth-like apical processes in H. dalmatensis), in lacking a tooth on the embolus (embolic tooth present in H. dalmatensis), and also by the proportions of the epigyne. Differences also occur in the spination of certain leg segments: male metatarsus I with two ventral spines in H. isaevi, but without spines in H. dalmatensis; female metatarsus IV with 4-5 retrolateral spines in H. isaevi, but with 3 spines in H. dalmatensis.
Male palp as in Figs 25-27. Terminal apophysis sharply turned, without ridge, but with tooth (Tt) in subterminal part, embolus without tooth.
Comments. In Crimea we found both closely related species, H. dalmatensis and H. isaevi, to be syntopical in two localities (Sudak Distr., 10 km W Sudak, Mezhdurechie Vill. and Feodosiya Distr., Karadag Nature Reserve). However, these species have quite different phenologies and adults of the two different spe- cies do not co-occur. The reproductive period of H. dalmatensis is in May-July with the peak in June, and in H. isaevi adults can be found in October-December, with their peak of activity in December. Distribution. West Palaearctic -Far East disjunct nemoral-subtropical range: Central and Southern Europe to Urals, Caucasus, Turkey, Eastern China, Far East Russia and Korea (Mikhailov 1997;Tuneva and Esyunin 2003;Helsdingen 2010;Platnick 2012).
Diagnosis. Haplodrassus pseudosignifer is very similar to H. signifer. The two species have no distinct differences in coloration, size or leg spination, but H. pseudosignifer can be differentiated from H. signifer by having an almost straight and shorter terminal apophysis and thinner embolus, and by the shape of the lateral pockets and fovea of the epigyne.
Coloration grey. Male palp as in Figs 68-70. Terminal apophysis short (length/width ratio ~ 2) and straight, ridge poorly developed; embolus almost straight and with a tooth.
Distribution. Steppe zone of Eurasia: from Crimea and Nikolaev Area of Ukraine east to Altai (Marusik et al. 1996;present data).
Comments. Haplodrassus pseudosignifer is a new species record for Crimea and Ukraine. Crimea is the westernmost point of the known distribution range.
Phenology. ♂♀ -V-VII, ♂♂ -IV, X. In Crimea the peak of activity in adults occurs in May. Diagnosis. The species can be easily recognized by the shape of the terminal apophysis, which has a peculiar long ridge, and also by the shape of the epigyne. Haplodrassus signifer is very similar to H. pseudosignifer (see the diagnosis for H. pseudosignifer).
Comments. Haplodrassus signifer is the largest and most abundant Haplodrassus species in Crimea.

Species diversity
The number of Haplodrassus species found in Crimea is rather high. Eight species, as in Crimea are known in the larger area of Bulgaria and Hungary (Helsdingen 2010). Several large countries such as Austria, Germany, Poland and Romania have 9 species of Haplodrassus. 10 species are known to occur in the Czech Republic, Switzerland and France. The highest species diversity in Europe, and probably in the Holarctic, is in Italy, with 13 species known from the mainland (Helsdingen 2010). Although Crimea was intensively investigated during only a short period we do not expect additional Haplodrassus species on the peninsula. The neighboring mainland Ukraine and Caucasus have no species that are absent in Crimea. There are several species that occurs in Romania or Bulgaria that are absent in Crimea: H. cognatus, H. moderatus (Kulczyński, 1897), and H. silvestris (Blackwall, 1833). In terms of species diversity per unit area of a country or region, Crimea rates as the most diverse place in Europe and in the Mediterranean (8 species in ~ 26 000 km 2 ). Only Israel has a similar number of species per unit area (8 species in ~27 000 km 2 ).

Phenology
Many specimens were collected using pitfall traps, which were regularly checked once in two week during one or two years. Thus, it was possible to analyze the seasonal dynamics of adult activity. All Crimean Haplodrassus species have one peak of activity of adults during the year. The maximum number of individuals and peak of activity for the adults of H. kulczynskii occurred in April; for H. bohemicus, H. minor, H. pseudosignifer and H. signifer