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28 October 2020 Karakumosa gen. nov., a new Central Asian genus of fossorial wolf spiders (Araneae: Lycosidae: Lycosinae)
Dmitri V. Logunov, Alexander V. Ponomarev
Author Affiliations +
Abstract

A new genus, Karakumosa gen. nov., is established to accommodate nine central Asian species of burrowing wolf spiders, of which seven are diagnosed and described as new: K. badkhyzica sp. nov. (♂♀, southern Turkmenistan), K. gromovi sp. nov. (♂♀, southern Uzbekistan), K. repetek sp. nov. (♂♀, eastern Turkmenistan), K. shmatkoi sp. nov. (♂♀, northern Ciscaspian region and Azerbaijan), K. tashkumyr sp. nov. (♂, Kyrgyzstan), K. turanica sp. nov. (♂♀, Turkmenistan), K. zyuzini sp. nov. (♂♀, Uzbekistan). Two generic transfers are proposed: Karakumosa alticeps (Kroneberg, 1875), comb. nov. and K. medica (Pocock, 1889), comb. nov. (both ex HognaSimon, 1885). Lectotypes are designated for Tarentula alticepsKroneberg, 1875 (♂) and T. medicaPocock, 1889 (♂). The localities of all Karakumosa species are mapped, and an identification key is provided as well. A short overview of the fauna and diversity of the fossorial Lycosidae of Central Asia is given.

INTRODUCTION

Fossorial wolf spiders of the family Lycosidae Sundevall, 1833 of Central Asia have been studied inadequately and remain poorly understood taxonomically (for an overview see Logunov, 2010). Central Asia is here defined as the territory of traditional Middle Asia and Kazakhstan plus the northern Ciscaspian region (Dagestan, Kalmykia, etc.), eastern Azerbaijan, and the neighbouring territories of western China, including Tibet, northern Pakistan, Afghanistan and Iran. To date, 30 species of burrowing lycosids have been recorded/described from Central Asia (Table 1) (Mikhailov, 2013; Otto, 2020; Roewer, 1955a; Song et al., 1999; Zamani et al., 2020; present data). However, many of the earlier records were based on doubtful identifications which are difficult to interpret and/or impossible to verify because the corresponding material is not accessible or its whereabouts is unknown. For instance, the species Hogna alticeps (Kroneberg, 1875), as it is listed now in the World Spider Catalog (2020), has been recorded from Central Asia many times (Andreeva, 1975, 1976; Kroneberg, 1875; Mikhailov & Fet, 1994; Ovtsharenko & Fet, 1980; Schmidt, 1895; Simon, 1899; Vlassov & Sytshevskaja, 1937; Zyuzin et al., 1994), but the distribution and even the taxonomy of this species remain largely unresolved. As it is evident from our study, the species is clearly neither a member of Hogna Simon, 1885, nor of Lycosa Latreille, 1804, and only few of the earlier records of H. alticeps actually correspond to this species. The same holds true for Hogna medica (Pocock, 1889), which was described from north-western Afghanistan (Pocock, 1889: sub Tarentula medica) and then reported once from Uzbekistan (Schmidt, 1895: sub Lycosa medica). Since then nobody has recorded this species from Central Asia again. The present paper attempts to bring some clarity to the taxonomy and distribution of these species, and to rectify their generic placement. The paraphyletic genera Hogna and Lycosa still remain two of those lycosid genera that are “used as a ‘dumping ground’ for wolf spiders that could not be satisfactory placed in other genera” (Murphy et al., 2006: 585).

The aim of the present work is to propose a new genus of fossorial wolf spiders from Central Asia to accommodate nine distinct species. Two of these are here transferred from Hogna, and seven species are diagnosed and described as new. The localities of all species are mapped and an identification key to all of them is provided. Finally a short overview of the fauna of fossorial Lycosidae of Central Asia is given.

MATERIAL AND METHODS

A total of 114 specimens have been examined for the present paper. They were borrowed from or deposited in the following museums: BMNH = The Natural History Museum, London, UK (curator: J. Beccaloni); MHNG = Muséum d'histoire naturelle, Genève, Switzerland (curator: P.J. Schwendinger); MMUE = Manchester Museum, University of Manchester, Manchester, UK (curator: D.V. Logunov); ISEA = Institute of Systematics and Ecology of Animals, Novosibirsk, Russia (curator: G.N. Azarkina); PSU = Department of Invertebrate Zoology and Aquatic Ecology of the Perm State University, Perm, Russia (curator: S.L. Esyunin); ZISP = Zoological Institute of the Russian Academy of Sciences, St-Petersburg, Russia (curator: V.A. Krivokhatsky); ZMMU = Zoological Museum of the Moscow University, Moscow, Russia (curator: K.G. Mikhailov).

All SEM micrographs, except for Figs 9-11, were taken with the assistance of A.V. Nazarenko (Rostov-on-Don, Russia) by means of a Carl Zeiss EVO 40 XVP scanning electron microscope at the Center for Collective Usage ‘Joint Centre of scientific and technological equipment of the Southern Scientific Centre of the Russian Academy of Sciences (research, development, approbation)’, Rostov-on-Don, Russia. The SEM micrographs of K. alticeps (Figs 9-11) were taken by Yu.M. Marusik (Magadan, Russia) with a Jeol JSM-5200 stereo scanning electron microscope at the Zoological Museum, University of Turku, Finland; he also compiled some digital photos (Figs 17-21). The majority of digital photos (Figs 32-39, 44, 49-54, 59, 61, 70-72, 89-93, 103-106, 117, 122, 126, 127, 130, 136, 137, 140, 161-165, 171) were taken by V.Yu. Shmatko (Rostov-on-Don, Russia) using a Sony Alpha A6000 ILCE-6000 Camera attached either to a Mikmed-2 var. 2 microscope (for general appearances) or to a MBS-1 microscope with a VOLNA-9 Lens (for copulatory organs). Digital photos of the male bulbs of K. alticeps (Figs 12-16) were taken by A.A. Fomichev (Novosibirsk, Russia) with an Olympus DP74 camera attached to an Olympus SZX 16 stereomicroscope at the Altai State University, Barnaul, Russia. Some digital photographs (Figs 27, 77-80, 85, 159) were made by the first author at the World Museum of Liverpool (UK) using a Canon 6D Mark II Camera with a Canon MP-E 65mm lens with Helicon Remote ver. 3.9.7W to control the StackShot 3X Macro Rail and camera settings. Helicon Focus 6.8.0 was used as processing software. Distribution maps were produced by G.N. Azarkina (Novosibirsk, Russia) using the online mapping software SimpleMappr (Shorthouse, 2010).

The terminology of sclerites of the copulatory organs follows Zyuzin (1993), except for the term ‘median apophysis' which is used sensu Griswold (1993) and Logunov (2010). The term ‘palea' refers to a swollen, partially membranous part of the embolic division from which the synembolus and the embolus originate (Figs 9-13: Pl). Short spines occurring on the ventral surface of tarsi in some lycosine genera (e.g. Lycosa, Zyuzicosa Logunov, 2010; Figs 140-141) were referred to by Zyuzin (1990) as ‘spinules’, a term which is also adopted here. ‘Chelicera length’ refers to the frontal length of the basal cheliceral segment on intact specimens; chelicerae were not removed from the specimens. The clypeus is the area between the AMEs and the frontal margin of the carapace. The sequence of leg segment measurements is as follows: femur + patella + tibia + metatarsus + tarsus (total). The leg formula is given from longest to shortest leg. All measurements are in mm.

Abbreviations

AER

anterior row of eyes

AME

anterior median eye

ALE

anterior lateral eye

a.s.l.

above sea level

C

conductor

D

described

Distr.

district

E

embolus

IP

inner plate of the MA

MA

median apophysis

Mt

metatarsus

MT

median tooth of the MA

OP

outer plate of the MA

PE

proximal extension of the MA

Pl

palea

PLE

posterior lateral eye

PME

posterior median eye

Se

synembolus

SER

of second row of eyes (formed by PMEs)

St

subtegulum

T

tegulum

Tb

tibia

TL

type locality

Tr

tarsus

Vil.

village

TAXONOMIC PART

Family Lycosidae Sundevall, 1833
Subfamily Lycosinae Sundevall, 1833
Genus Karakumosa gen. nov.

  • Type species: Karakumosa repetek sp. nov. from the Repetek Reserve, Karakumy Desert, Turkmenistan, Central Asia (male holotype deposited in the ISEA).

  • Diagnosis: The genus Karakumosa gen. nov. belongs to the subfamily Lycosinae (sensu Zyuzin, 1993; see Dondale, 1986; Murphy et al., 2006; Piacentini & Ramírez, 2019) and is most similar to Zyuzicosa Logunov, 2010 (see Logunov, 2010, 2012). Both genera have a bipartite/biramous synembolus, the same SER/AER ratio (more than 1.3), tarsi of all legs with scopulae and spinules, and a prolatero-apical origin of the embolus. The new genus differs from all other Central Asian Lycosinae genera (see Logunov, 2010: table 2), including Zyuzicosa, by the following combination of characters: black ventral colour pattern of abdomen absent (Figs 25, 52, 91, etc.); median apophysis consisting of two flat plates (Figs 55, 118: OP, IP); synembolus with two acutely pointed lamellae (Figs 57, 98-99, 116, 118, 129, 149, 163, 175); epigynal atrium at least twice longer than wide; and septal pedicel absent (Figs 27, 44, 61, 72, 85, 96).

  • Etymology: The new generic name is composed of two parts: ‘Karakum’, referring to the regional name for the Karakumy Desert where the type species and some other species were discovered, combined with the ending of the generic name Lycosa (meaning ‘tear like a wolf'; see Cameron, 2005: 303), to which the majority of large burrowing Holarctic wolf spiders are currently assigned. The generic name is feminine in gender.

  • Description: Large to very large fossorial wolf spiders, with body lengths 20.23±3.23 (n=8) in males, and 22.9±4.05 (n=7) in females.

  • Carapace: In both sexes relatively low, with a clearly marked gradual descent from cephalic region towards abdomen (Figs 26, 35, 54, 78, 138, 158), densely clothed with white or yellowish white setae, and with prominent dense white or yellowish marginal pubescence; all three characters are typical of lycosid burrowers (see Zyuzin, 1990).

  • Chelicerae: Large, vertical, their proximal halves/two-thirds of frontal side densely clothed with white or yellowish setae (Figs 32, 53, 79, 136); cheliceral groove with three promarginal and three retromarginal teeth (Figs 144, 177).

  • Eyes: AER procurved and distinctly (1.3-1.4 times) shorter than SER (Figs 32, 53, 136); PME/AME ratio 1.7-2.5.

  • Clypeus: Narrow, its height equal or 1.3-1.7 times shorter than AME diameter.

  • Labium: Visibly wider than long (length/width ratio 0.6-0.8).

  • Sternum: Ovoid, densely covered with white setae in both sexes (e.g. Figs 23, 49, 52, 137).

  • Abdomen: Venter in both sexes without black pattern (e.g. Figs 25, 52, 91), thus distinct from the majority of Palaearctic burrowing lycosine genera described to date (see Simon, 1876; Logunov, 2010, 2012).

  • Legs: leg formula IV,I,II,III in both sexes, rarely IV,II,I,III in some males; all segments densely covered with white setae; in both sexes, metatarsi and tarsi I-II ventrally with well-developed scopulae and longitudinal rows of spinules (sometimes poorly visible; Figs 36, 63, 140), tarsi III-IV only with ventral longitudinal rows of spinules (in some specimens a scopula could be developed on lateral sides of segments only; Fig. 141). Female pedipalp: With a single tarsal claw (e.g. Figs 26, 35, 138).

  • Female copulatory organs: Epigyne with a well pronounced longitudinal atrium being at least two times longer than wide; edges at proximal end of atrium forming two round convergent lips, these usually markedly sclerotized and darker (brown or russet) than nearby cuticle (e.g. Figs 27, 61, 85); septal pedicel absent; median septum essentially reduced to posterior transverse, slightly convex plate of variable shape (e.g. Figs 27, 96); spermathecae tube-shaped, only slightly wider than or of same width as narrow and short insemination ducts (e.g. Figs 28, 43, 62, 121).

  • Male pedipalp: Femur length equal to that of patella+tibia (Figs 15-16); cymbium symmetrical, twice as long as wide, with almost round alveolus (Fig. 65), its length about equal to that of palpal tibia; distal part of cymbium 1.1-1.3 times shorter than alveolus length (Fig. 65); cymbium with a cluster of blunt, rigid and straight bristles on its tips (Figs 45, 69) as typical of lycosid burrowers (Zyuzin, 1990, 1993).

  • Male copulatory organs: Subtegulum round and comparatively small, situated in proximal-mesal position (Figs 9, 46, 56, 98: St); tegulum round and broad, in unexpanded palp clearly visible only on prolateral-proximal side of bulbus (Figs 55, 98, 118: T); median apophysis wide (usually wider than long) and broad (Figs 9, 55, 118: OP, IP), folded along its proximal edge and consisting of two (outer and inner) flat plates, the outer plate bearing a proximal extension and a median tooth (Figs 56, 98: MT, PE); median tooth rarely singular and finger-shaped (Figs 30, 42), usually consisting of a lateral claw and a median edge bearing micro-teeth (indicated by arrow in Figs 123, 128) and in some species with a low serrate flange at its foot (indicated by arrow in Figs 125, 134, 152); synembolus biramous, with two acutely pointed lamellae (Figs 57, 98, 118: Se); a triangular hyaline conductor present, well-developed and pointed ventrad (Figs 12, 56, 118: C); embolus thin, with a rather wide and prominent pars pendula (Figs 4, 116, 129), its origin in a prolatero-apical position, with only the embolic tip visible in between or beneath branches of the synembolus in unexpanded palp (Fig. 57).

  • Comments: Within the Lycosoidea the transverse median apophysis is considered a typical feature of the Lycosidae (Griswold, 1993) and a synapomorphy of the Lycosinae (Dondale, 1986), although its shape varies. In Karakumosa gen. nov. the median apophysis is large and composite, consisting of two distinct plates that appear fused along their proximal edges: the outer and the inner plates (e.g. Figs 9, 56, 118: OP, IP). Such a complex structure of the median apophysis is currently regarded as unique within the Lycosinae and within the entire family Lycosidae. Of the lycosid genera known to us, only two have a comparably complex structure of the median apophysis: Oculicosa Zyuzin, 1993 (Logunov & Gromov, 2011: figs 1-2) and Zyuzicosa (Logunov, 2010: figs 34-39). The outer plate of the median apophysis of Karakumosa gen. nov., with its proximal extension and median tooth of MA, looks like an apomorphic modification of the prominent transverse chitinous ridge of the median apophysis in Oculicosa. Obviously, both structures are homologous.

  • Another unique feature of Karakumosa gen. nov. is its biramous synembolus, consisting of two thin and very acutely pointed lamellae (e.g. Figs 4, 57, 67, 116). The only other lycosid genus with a comparable conformation of the synembolus is Zyuzicosa (Logunov, 2010: figs 64, 66), but in the latter genus it consists of one acutely pointed lamella and a wide, strongly sclerotized base. In the absence of a phylogenetic analysis of the Palaearctic genera of Lycosidae (but see Murphy et al., 2006; Piacentini & Ramírez, 2019), it is difficult to establish whether both unique features of Karakumosa gen. nov. are primitive or derived (they are likely to be derived). Yet, as a provisional hypothesis to be further tested, we consider both of them as the putative synapomorphies of Karakumosa gen. nov.

  • We speculate that if the genus Karakumosa gen. nov. was included in one of the two latest phylogenetic analyses of the Lycosidae based on DNA data (Murphy et al., 2006; Piacentini & Ramírez, 2019), it would likely be placed either within the clade E (sensu Murphy et al., 2006: figs 2-3), somewhere close to Lycosa tarantula (Linnaeus, 1758) and the clade E1, or within the Lycosinae (sensu Piacentini & Ramírez, 2019: fig. 4), in the branch containing the Palaearctic Lycosa species.

  • Composition: To date nine species are assigned to Karakumosa gen. nov.: K. alticeps (Kroneberg, 1875), comb. nov. (♂♀, Uzbekistan and southern Kazakhstan), K. badkhyzica sp. nov. (♂♀, Turkmenistan), K. gromovi sp. nov. (♂♀, southern Uzbekistan), K. medica (Pocock, 1889), comb. nov. (♂♀, north-western Afghanistan), K. repetek sp. nov. (♂♀, Turkmenistan), K. shmatkoi sp. nov. (♂♀, northern Ciscaspian region and Azerbaijan), K. tashkumyr sp. nov. (♂, Kyrgyzstan), K. turanica sp. nov. (♂♀, Turkmenistan), K. zyuzini sp. nov. (♂♀, Uzbekistan).

  • Distribution: Central Asia (Fig. 1): (semi)desert regions of the northern Ciscaspian Region, eastern Azerbaijan, Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan and north-western Afghanistan; the occurrence of this genus in northern and north-eastern Iran and Tajikistan is very likely.

  • The genus is essentially restricted to the so-called Turan zoogeographic province (sensu Kryzhanovsky, 2002) (Fig. 1). It is the third genus of fossorial wolf spiders within the currently known Lycosinae that is confined to Central Asia (Table 1). The other two are Oculicosa (see Zyuzin, 1993; Logunov & Gromov, 2011: fig. 6) and Zyuzicosa (see Logunov 2010, 2012: map), but their ranges are markedly smaller and of a different configuration. Karakumosa gen. nov. seems to be a typical Turan genus (sensu Pravdin & Mishchenko, 1980) and an endemic to Central Asia; its geographical range lies within the Turan lowlands of the desert zone. We are unaware of any other Central Asian spider genus that has a distributional pattern similar to that of Karakumosa gen. nov.

  • Fig. 1.

    Geographical range of the genus Karakumosa gen. nov.

    img-z4-8_275.jpg

    Key to the species of Karakumosa gen. nov.

    The female of K. tashkumyr sp. nov. remains unknown and thus is not included here.

    1A Males 2

    1B Females 10

    2A Proximal extension of median apophysis clearly hook-shaped (Figs 60, 118, 150) 3

    2B Proximal extension of median apophysis of a different shape (Figs 2, 46, 83, 98) 5

    3A Median tooth of median apophysis with a low serrate flange at its foot (indicated by arrow in Figs 125, 134, 152) 4

    3B Median tooth of median apophysis without such flange (Figs 66, 74-75) gromovi sp. nov.

    4A Inner plate of median apophysis as wide as outer plate, clearly visible in ventral view (indicated by arrow in Fig. 148); tips of both synembolic lamellae markedly bent downward (Fig. 149) tashkumyr sp. nov.

    4B Inner plate of median apophysis comparatively narrower and almost hidden beneath outer plate (indicated by arrow in Fig. 118); synembolic lamellae straight or with tips only slightly bent downward (Fig. 116) shmatkoi sp. nov.

    5A Proximal extension of median apophysis prominent (Figs 46, 83, 98) 6

    5B Proximal extension of median apophysis not prominent (Fig. 174) zyuzini sp. nov.

    6A Proximal extension of median apophysis spade/spatula-like, as wide as long (Figs 46, 98) 7

    6B Proximal extension of median apophysis wider than long (Figs 2, 83, 161) 8

    7A Median tooth of median apophysis finger-shaped (Figs 31, 42) badkhyzica sp. nov.

    7B Median tooth of median apophysis bifurcated at its tip (Fig. 100) repetek sp. nov.

    8A Median tooth of median apophysis flat, without a ventral bulge (Figs 5, 164) 9

    8B Median tooth of median apophysis with a ventral bulge (Fig. 87) medica

    9A Proximal extension of median apophysis twice as wide as long, with a markedly pointed prolaterad-directed shoulder (Fig. 161); median tooth quadrangular (Fig. 164) . turanica sp. nov.

    9B Proximal extension of median apophysis triangular, with an obtuse prolaterad-directed shoulder (Fig. 12); median tooth triangular, with a serrate prolateral edge (Figs 5-8) alticeps

    10A Edges of epigynal atrium subparallel or slightly bent outward (Figs 44, 61, 72, 85) 11

    10B Edges of epigynal atrium slanted to each other; atrium anteriorly clearly narrower than posteriorly (Figs 27, 96) 13

    11A Edges of epigynal atrium subparallel (Figs 61, 85), spermathecae as in Figs 62, 86 12

    11B Edges of epigynal atrium slightly bent outward (biconvex), atrium barrel-shaped (Fig. 44), spermathecae as in Fig. 43 badkhyzica sp. nov.

    12A Posterior transverse plate of epigyne in the shape of an inverted triangle (Figs 61, 72), spermathecae markedly widened anteriorly (Figs 62, 73) gromovi sp. nov.

    12B Posterior transverse plate of epigyne anchor-shaped (Fig. 85), spermathecae not widened anteriorly, worm-shaped (Fig. 86) medica

    13A Posterior transverse plate more or less anchor-shaped (Figs 27, 167-168), spermathecae curved anticlockwise or subparallel (Figs 28, 121, 166) 14

    13B Posterior transverse plate dumbbell-shaped (Fig. 96), spermathecae curved clockwise (Fig. 95) repetek sp. nov.

    14A Epigynal atrium markedly narrowed at its anterior end, pawn-shaped (Figs 27, 122) 15

    14B Epigynal atrium much less narrower at its anterior end, with almost subparallel lateral edges (Figs 167, 179) 16

    15A Posterior transverse plate of epigyne in the shape of a low inverted triangle, epigynal edges strongly sigmoid (Fig. 122); spermathecae markedly swollen in anterior portion (Fig. 121) shmatkoi sp. nov.

    15B Posterior transverse plate of epigyne slightly procurved, epigynal edges indistinctly sigmoid (Fig. 27); spermathecae not swollen anteriorly (Figs 28-29) alticeps

    16A Posterior transverse plate of epigyne with straight posterior margin (Fig. 179); spermathecae distinctly inclined towards each other (Fig. 182) zyuzini sp. nov.

    16B Posterior transverse plate of epigyne anchor-shaped, with convex posterior margin (Fig. 167); spermathecae more or less parallel to each other (Fig. 166) turanica sp. nov.

    Karakumosa alticeps (Kroneberg, 1875) comb. nov.
    Figs 2-29, 76
    Tarentula alticeps Kroneberg, 1875: 40, pl. 4, fig. 28 (description of male and female; syntypes in the ZMMU).
    Lycosa alticeps. – Schmidt, 1895: 449 (partim). – Zyuzin et al., 1994: 4, 9.
    Hogna alticeps. – Roewer, 1955b: 247.

  • Lectotype (designated here; Figs 17-21): ZMMU, Ta-1219; male; “Turkestan region” (no exact locality); no date; [A.P.] Fedchenko (Turkestan scientific expedition of the Imperial Society of Devotees of Natural Science).

  • Paralectotype: ZMMU, Ta-1218; 1 male; [KAZAKHSTAN], “Dyusebai” [well] (an unknown locality, apparently in Chardara Distr. of South Kazakhstan Area); no date; [A.P.] Fedchenko (Turkestan scientific expedition of the Imperial Society of Devotees of Natural Science). For information about other former syntypes of T. alticeps, see below under K. turanica sp. nov.

  • Other material: ISEA, 001.8403; 6 males, 1 female; KAZAKHSTAN, Zhambyl Area, Chu Distr., c. 9th km of road from Tole Bi (= Novotroitskoe) to Moyyunkum, Chu river valley (c. 43°45′N, 73°46′E), sands; 31.V.-2.VI.1990; leg. A.A. Fedorov & A.A. Zyuzin. – MHNG; 2 males; same data. – ISEA 001.8404; 1 male, 1 female; KAZAKHSTAN, South Kazakhstan Area, Chardara Distr., Kyzyl-Kum Desert, Karatau Mt. Range, Karamola Mt. (c. 42°20′N, 67°45′E); [late May] 1994; leg. A.A. Zyuzin. – ZMMU; 1 male, 1 female; same data. – MMUE; 1 male, 1 female; same data. – MMUE; 1 male; KAZAKHSTAN, Turkestan Area, Syr Darya River valley, near Zhankel' Vil. (42°32′46.1″N, 68°10′42.7″E), tugay (= gallery forest), 191 m a.s.l.; 28.V.2017; leg. Yu.V. Dyachkov. – MMUE; 1 male; KAZAKHSTAN, Kyzylorda Area, Syr-Darya River valley, near Tartogay Vil. (44°24′42.8″N, 66°16′40.1″E), tugay (= gallery forest), 142 m a.s.l.; 12.VI.2017; leg. Yu.V. Dyachkov. – MMUE; 1 male, 3 females; KAZAKHSTAN, Almaty Area, Ili Distr., near Kapchagay (on left side of road before reaching town (c. 43°50′17.3″N 76°58′14.9″E), sands; 25.-26.V.1990; leg. A.A. Zyuzin.

  • Etymology: According to Parker (1980) the specific epithet could be translated as ‘high headed’, originating from the Latin ‘altus' meaning high, and ‘caput’ meaning the head.

  • Diagnosis: The male of K. alticeps is most similar to that of K. turanica sp. nov., but can be easily distinguished by the narrower, obtuse prolaterad-directed shoulder of the proximal extension of the median apophysis of the palpal organ (wider and markedly pointed in K. turanica sp. nov.; Figs 2, 12 cf. Fig. 161) and by the comparatively smaller, triangular median tooth with a serrate median edge (wide and quadrangular in K. turanica sp. nov.; Figs 5-8 cf. Fig. 164). The female of K. alticeps is most similar to that of K. shmatkoi sp. nov., from which it can be distinguished by almost straight lateral edges of the epigynal atrium and by the narrow, slightly procurved posterior transverse plate (sigmoid edges and a low inverted triangle-shaped posterior transverse plate in K. shmatkoi sp. nov.; Fig. 27 cf. Figs 122, 139), as well as by the narrow, not swollen spermathecae (markedly swollen in K. shmatkoi sp. nov.; Fig. 28 cf. Fig. 121).

  • Description: Male (from Chu river valley; ISEA, 001.8403). Measurements: Carapace 10.00 long, 7.50 wide. Eye sizes and interdistances: AME 0.43, ALE 0.35, PME 1.10, PLE 1.05, AME-AME 0.25, AME-ALE 0.15, PME-PME 0.90 PME-PLE 1.90. Width of anterior eye row 2.10, of second row 2.80, of third row 3.40. Clypeus height 0.25; chelicera length 4.75. Abdomen 9.80 long, 6.50 wide. Length of leg segments: I 10.00 + 4.20 + 8.30 + 8.80 + 4.20 (35.50); II 8.50 + 4.20 + 7.60 + 8.70 + 4.50 (33.50); III 8.70 + 3.40 + 6.50 + 9.00 + 4.40 (32.00); IV 10.70 + 3.80 + 8.20 + 11.30 + 4.90 (38.90). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 22-23): Carapace brownish, with a wide median longitudinal yellowish white band of setae along the entire length of the carapace and two wide lateral brownish longitudinal bands of setae; carapace sides with wide marginal bands of white setae. Sternum light brown, densely covered with yellowish white setae. Maxillae brown. Labium brown, with a yellow tip. Chelicerae brown, proximal half of frontal side densely covered with yellowish white setae. Abdomen: dorsum densely covered with white setae, with a long, wide, yellow brownish cardiac mark outlined by a brown line; sides and venter densely covered with yellowish white setae. Book-lung covers yellow, covered with yellowish white setae. Spinnerets light brown. All legs yellow brownish, densely covered with white setae; Mt I and Tr I ventrally brown, but dorsally densely covered with white setae. Palp yellow, with brownish cymbium, densely covered with white setae.

  • Palp structure (Figs 2-20): Acutely pointed synembolic lamellae subparallel to each other; median tooth medium-sized, notched at its tip and with a serrate median edge; proximal extension wide and obtuse at its prolaterad-directed shoulder; inner plate large and ovoid, clearly visible in ventral view; conductor triangular, acutely pointed and bent at its tip.

  • Female (from Chu river valley; ISEA, 001.8403). Measurements: Carapace 10.70 long, 8.00 wide. Eye sizes and interdistances: AME 0.50, ALE 0.45, PME 1.20, PLE 1.15, AME-AME 0.35, AME-ALE 0.15, PME-PME 1.15, PME-PLE 2.55. Width of anterior eye row 2.35, of second row 3.15, of third row 3.80. Clypeus height 0.20; chelicera length 5.75. Abdomen 12.00 long, 9.80 wide. Length of leg segments: I 9.30 + 4.10 + 7.20 + 6.50 + 3.80 (30.90); II 8.70 + 4.10 + 6.40 + 6.10 + 3.80 (29.10); III 7.70 + 3.50 + 5.60 + 6.70 + 4.00 (27.50); IV 10.00 + 4.00 + 7.20 + 9.20 + 4.70 (35.10). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 24-26): Carapace brownish, densely covered with yellowish white setae and with two longitudinal bands of brownish setae; carapace sides with wide marginal bands of white setae. Sternum brownish yellow, densely covered with white setae. Maxillae and labium brown. Chelicerae dark brown, proximal half of frontal side densely covered with yellowish white setae. Abdomen: dorsum densely covered with yellowish white setae and with a large, wide, brownish cardiac mark; sides and venter, including book-lung covers, densely covered with white setae. Spinnerets light brown. All legs and palps yellow, densely covered with white setae; tarsi of all legs and palps darker (brownish). Palps with a single claw at their tips.

  • Epigyne and vulva (Figs 27-29): Epigynal atrium twice as long as wide, markedly narrower at its anterior end and with only slightly sigmoid lateral edges; posterior transverse plate narrow and slightly procurved; spermathecae tube-shaped, not incrassate, directed antero-mediad, inclined towards each other.

  • Comments: The original type series of Tarentula alticeps, deposited in the ZMMU, consists of two species. Two syntype males (Ta-1218 and Ta-1219) were collected from today's southern Kazakhstan and are conspecific. In order to stabilize the species-group name Tarentula alticeps, one of them (Ta-1219; Figs 17-21) is here designated as the lectotype. Both males are conspecific with those collected from southern Kazakhstan together with females in the last two decades (see above under ‘Other material'). Thus, there is no doubt about matching the sexes in this species.

  • The female paralectotype (Figs 165, 168) and juveniles of Tarentula alticeps were collected from present-day Uzbekistan (Ulus and Samarkand). This adult female differs from those of K. alticeps in having a much wider epigynal atrium (Fig. 168 cf. Fig. 27) and subparallel spermathecae (convergent in K. alticeps; Fig. 166 cf. Fig. 28). This female conforms to the description of K. turanica sp. nov. from Turkmenistan (see below; Figs 166-167). Hence, we assign the female paralectotype of Tarentula alticeps and its Uzbekistani records (Fig. 155) to K. turanica sp. nov.

  • Moreover, the immature syntypes of T. alticeps examined (Ta-1216) contain specimens of two different genera. One immature female has a brown, contrastingly coloured venter, which is typical of Allohogna Roewer, 1955a, Lycosa (s.str.), Zyuzicosa and some other burrowing wolf spiders, but that was never observed in Karakumosa gen. nov. It is additional evidence that the original type series of Tarentula alticeps is indeed not conspecific.

  • Distribution: Southern Kazakhstan (Kroneberg, 1875; Zyuzin et al., 1994; present data) (Fig. 76); all the known records of K. alticeps lie in the same South-Turkestan Phytogeographic Province (see Pravdin, 1978 for further details). At one of the localities examined (Karamola Mt.), K. alticeps was found together with another large burrowing wolf spider, Zyuzicosa turlanica Logunov, 2010; see Logunov (2010, 2012) for further details about the latter species and its records.

  • Many of the earlier records of K. alticeps are in need of confirmation by re-examining the corresponding material, which was not available for the present study. For instance, the records from Uch-Adzhi (c. 38°05′N, 62°48′E) and Turkmenbashi (c. 40°02′N, 52°59′E) in Turkmenistan by Schmidt (1895: sub. Lycosa alticeps) most likely belong to K. turanica sp. nov., while his record from the Fergana region of Uzbekistan (no exact locality) could belong to K. tashkumyr sp. nov. The records of K. alticeps from Tajikistan (Kondara, Kvak, Varzob Canyon, Hissar Mt. Range, Ramit, Gandzhina) by Andreeva (1975, 1976: sub Lycosa alticeps) are likely based on misidentifications. To date we have been able to re-examine only two samples from Tajikistan that are deposited in the ZMMU (Ta-2916: 1 male from Kondara; Ta-2915: 3 females, 4 juveniles from Yavan-su Vil.) and earlier identified by Jan Buchar as Lycosa alticeps. Both samples turned out to be misidentified and in fact belong to Zyuzicosa laetabunda (Spassky, 1941).

  • Figs 2-11.

    Karakumosa alticeps (Kroneberg, 1875), left male palp; specimens from Chu river valley (2-4, 7-11) and Karamola Mt. (5-6). (2, 9) Bulbus, ventral view. (3, 11) Ditto, retrolateral view. (4) Embolic division, ventral view. (5-8) Median tooth of MA, proximal view. (10) Bulbus, apical view. Scale bars 0.25 mm (2-3), 0.1 mm (4-11). Abbreviations as explained in Material and methods.

    img-z7-1_275.jpg

    Figs 12-21.

    Karakumosa alticeps (Kroneberg, 1875); male specimens from Tartogay Vil. (12-15) and Zhankel' Vil. (16), and the male lectotype (17-21). (12, 17) Left bulbus, ventral view. (13, 20) Ditto, retrolateral view. (14, 19) Ditto, posterior view. (15, 16) Left male palp, retrolateral view. (18) Left bulbus, apical view. (21) Original data labels of the lectotype. Scale bars 0.5 mm (15, 16), 0.2 mm (12-14, 17-20). Abbreviations as explained in Material and methods.

    img-z8-1_275.jpg

    Figs 22-29.

    Karakumosa alticeps (Kroneberg, 1875); male (22-23) and female (24-28) from Chu river valley, and female (29) from Karamola Mt. (22, 24) Body, dorsal view. (23, 25) Ditto, ventral view. (26) Ditto, lateral view. (27) Epigyne, ventral view. (28-29) Vulva, dorsal view. Scale bars 1 cm (22-26), 0.25 mm (27-29).

    img-z9-1_275.jpg

    Karakumosa badkhyzica sp. nov.
    Figs 30-48, 76
    Lycosa alticeps (Kroneberg, 1875). – Ovtsharenko & Fet, 1980: 443. – Mikhailov & Fet, 1994: 508 (misidentification of specimens from Badkhyz).

  • Holotype: ISEA; male; TURKMENISTAN, Mary Velayat, c. 73 km NW of Serhetabat (= Kushka), Badkhyz (= Badhyz) Reserve (c. 35°52′N, 61°40′E), cordon Kyzyl-Dzhar, clay slope; 10.-14.IV.1993; leg. A.A. Zyuzin.

  • Paratypes: ISEA; 3 females; collected together with the holotype. – ZMMU; 1 male, 1 female; same data. – MMUE; 1 male, 1 female; same data.

  • Etymology: The species epithet is latinized adjective derived from the name of the type locality, the Badkhyz (= Badhyz) Reserve in Turkmenistan.

  • Diagnosis: The male of K. badkhyzica sp. nov. is most similar to that of K. repetek sp. nov. (Figs 94, 98, 102), but differs in having a smaller and narrower proximal extension of the MA (wide, spade-shaped in K. repetek sp. nov.; Figs 40, 46 cf. Fig. 98) and an undivided, finger-shaped median tooth (bifurcated in K. repetek sp. nov.; Figs 30-31, 42 cf. Figs 97, 100). The female of this species is similar to that of K. medica (Figs 85-86), but can be readily distinguished by a shorter epigynal atrium (Fig. 44) and by the shape of the spermathecae: anteriorly markedly swollen in K. badkhyzica sp. nov. (Fig. 43), worm-shaped, anteriorly not swollen in K. medica (Fig. 86).

  • Description: Male (paratype, MMUE). Measurements: Carapace 7.65 long, 4.50 wide. Eye sizes and interdistances: AME 0.40, ALE 0.34, PME 0.68, PLE 0.75, AME-AME 0.20, AME-ALE 0.10, PME-PME 0.80 PME-PLE 0.85. Width of anterior eye row 1.63, of second row 2.30, of third row 2.20. Clypeus height 0.10; chelicera length 2.75. Abdomen 5.75 long, 3.25 wide. Length of leg segments: I 8.20 + 3.30 + 7.10 + 7.20 + 4.40 (30.20); II 7.80 + 3.40 + 6.20 + 7.50 + 3.70 (28.60); III 7.40 + 3.00 + 5.30 + 8.00 + 3.90 (27.60); IV 8.50 + 3.20 + 7.00 + 9.80 + 4.50 (33.00). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 37-38): Carapace brownish, with a wide median longitudinal yellowish white band of setae along entire length of carapace and with two wide lateral brown longitudinal bands of setae; carapace sides with wide marginal bands of white setae. Sternum light brown, densely covered with white setae. Maxillae and labium brownish, with yellow tips. Chelicerae brown, proximal half of frontal side densely covered with white setae. Abdomen: dorsum densely covered with white setae, with a long, wide, yellow-brown cardiac mark; sides and venter densely covered with white setae. Book-lung covers yellow, covered with white setae. Spinnerets brown. All legs yellow, densely covered with white setae; Mt I and Tr I brown, also densely covered with white setae. Palp yellow, with the brownish cymbium, densely covered with white setae.

  • Palp structure (Figs 30-31, 39-42, 45-48): Acutely pointed synembolic lamellae diverging from each other; median tooth finger-shaped, in some specimens with a basal tooth; proximal extension narrow, spatula-like; inner plate large, with an extended retrolateral shoulder; conductor triangular, pointed at its tip.

  • Female (paratype, MMUE). Measurements: Carapace 8.70 long, 6.50 wide. Eye sizes and interdistances: AME 0.45, ALE 0.38, PME 0.88, PLE 0.95, AME-AME 0.25, AME-ALE 0.15, PME-PME 0.93, PME-PLE 1.20. Width of anterior eye row 1.93, of second row 2.68, of third row 3.30. Clypeus height 0.30; chelicera length 4.35. Abdomen 8.76 long, 6.40 wide. Length of leg segments: I 6.60 + 3.20 + 4.90 + 4.50 + 2.80 (22.00); II 6.50 + 3.20 + 4.30 + 4.30 + 2.70 (21.00); III 5.30 + 2.50 + 3.60 + 5.10 + 2.80 (19.30); IV 7.50 + 3.00 + 5.30 + 6.70 + 3.50 (26.00). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 32-35): Carapace yellow-brown, densely covered with white setae and with two paramedian longitudinal bands of brown setae. Sternum brownish yellow, densely covered with white setae. Maxillae and labium brown, with yellow tips. Chelicerae dark brown, proximal half of frontal side densely covered with yellowish white setae. Abdomen: dorsum densely covered with white setae, with a large, wide, brownish cardiac mark and with a brownish pattern of transverse lines; sides and venter, including book-lung covers, densely covered with white setae. Spinnerets brownish. All legs and palps yellow, densely covered with white setae; tarsi of all legs darker (brownish). Palps with a claw at their tips.

  • Epigyne and vulva (Figs 43-44): Epigynal atrium twice as long as wide, with lateral edges slightly biconvex; posterior transverse plate short, anchor-shaped; spermathecae tube-shaped, directed antero-mediad, inclined towards each other, distinctly swollen in anterior portion.

  • Distribution: Only known from the type locality, the Badkhyz Reserve in Turkmenistan (Fig. 76).

  • Habitat: This species occurs in sparse pistache woodland, with an undergrowth dominated by bulbous bluegrass (Poa bulbosa) and sedge (Carex pachystylis) (Ovtsharenko & Fet, 1980: sub Lycosa alticeps).

  • Figs 30-38.

    Karakumosa badkhyzica sp. nov.; male paratype (30-31, 37-38) and female paratype (32-35), MMUE. (30-31) Median tooth of MA, posterior view. (32) Female prosoma, frontal view. (33, 37) Body, ventral view. (34, 38) Ditto, dorsal view. (35) Female prosoma, lateral view. (36) Female tarsus I, ventral view. Scale bars 0.5 cm (33-35, 37-38), 0.3 mm (36), 0.25 cm (32), 0.1 mm (30, 31).

    img-z11-4_275.jpg

    Figs 39-48.

    Karakumosa badkhyzica sp. nov.; male paratype, MMUE (39-42), male holotype (45-48) and female paratype (43-44). (39-40, 46) Bulbus, ventral view. (41) Ditto, retrolateral view. (42, 48) Median tooth of MA, posterior view. (43) Vulva, dorsal view. (44) Epigyne, ventral view. (45) Cymbium tip, dorsal view. (47) Bulbus, apical view. Scale bars 0.5 mm (39, 44), 0.25 mm (40-41, 43), 0.2 mm (45-47), 0.1 mm (42), 0.05 mm (48). Abbreviations as explained in Material and methods.

    img-z12-1_275.jpg

    Karakumosa gromovi sp. nov.
    Figs 49-76

  • Holotype: ISEA; male; UZBEKISTAN, Surxondaryo Region, Baisun [= Boysun] Distr., c. 44 km SW of Denau [= Denov], SE foothills of Dzhetymkalyas Mt. Range (38°01′10.6″N, 67°28′06.3″E), c. 640 m. a.s.l.; 13.-14.V.1994; leg. A.V. Gromov.

  • Paratypes: ISEA; 1 female; collected together with the holotype. – ZMMU; 1 male, 1 female; same data. – MMUE; 1 male, 1 female; same data. – ISEA; 1 female; from the type locality; 14.V.1994; leg. A.V. Gromov & A.A. Zyuzin.

  • Etymology: This species is dedicated to our friend and colleague, Mr Alexander V. Gromov (Bingen-am-Rhein, Germany), who collected the type specimens.

  • Diagnosis: In the hook-shaped proximal extension of the MA, the male of K. gromovi sp. nov. is most similar to males of K. shmatkoi sp. nov. (Figs 115-120) and K. tashkumyr sp. nov. (Figs 145-154), but can be readily distinguished by the absence of a low serrate flange at the foot of the median tooth (present in K. tashkumyr sp. nov.; Figs 74-75 cf. Figs 107-112 and Figs 152-154). In the straight, subparallel lateral edges of the epigynal atrium, the female of K. gromovi sp. nov. is most similar to that of K. medica, from which it can be distinguished by the triangular posterior transverse plate of the median septum and by the wider and shorter spermathecae (anchor-shaped posterior transverse plate and worm-shaped spermathecae in K. medica; Figs 61-62, 72-73 cf. Figs 85-86).

  • Description: Male (paratype in MMUE). Measurements: Carapace 9.50 long, 7.40 wide. Eye sizes and interdistances: AME 0.50, ALE 0.45, PME 1.20, PLE 1.00, AME-AME 0.20, AME-ALE 0.15, PME-PME 1.25, PME-PLE 1.50. Width of anterior eye row 2.00, of second row 2.80, of third row 3.50. Clypeus height 0.35; chelicera length 3.90. Abdomen 8.75 long, 6.25 wide. Length of leg segments: I 9.80 + 3.90 + 8.00 + 8.40 + 4.40 (34.50); II 9.40 + 3.90 + 7.50 + 8.80 + 4.00 (33.60); III 8.20 + 3.10 + 6.50 + 9.00 + 3.50 (30.30); IV 9.80 + 3.50 + 8.50 + 11.50 + 4.70 (38.00). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 49-50): Carapace brown, densely covered with white setae, with two wide longitudinal bands of yellow brownish setae. Sternum brownish yellow, densely covered with white setae. Maxillae and labium orange-coloured, with yellow tips. Chelicerae brown, proximal half of frontal side and lateral sides densely covered with white setae. Abdomen: dorsum densely covered with white setae, with a wide, yellow brownish cardiac mark and with a pattern consisting of yellow brownish transverse stripes and patches; sides and venter densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets yellowish grey. All legs brownish yellow, densely covered with white setae; Mt I-II and Tr I-II darker (brownish), also densely covered with white setae. Palp yellow, with brownish cymbium, densely covered with white setae.

  • Palp structure (Figs 55-60, 64-69, 74-75): Acutely pointed synembolic lamellae subparallel to each other; median tooth markedly bifurcated; proximal extension wide and markedly hook-shaped; distance between proximal extension and median tooth short, equal to width of proximal extension; inner plate transverse, with an extended, obtuse retrolateral shoulder; conductor triangular, obtuse at its tip.

  • Female (paratype in MMUE). Measurements: Carapace 12.00 long, 7.80 wide. Eye sizes and interdistances: AME 0.55, ALE 0.45, PME 1.30, PLE 1.20, AME-AME 0.25, AME-ALE 0.18, PME-PME 1.50, PME-PLE 1.70. Width of anterior eye row 2.00, of second row 3.15, of third row 4.00. Clypeus height 0.40; chelicera length 6.00. Abdomen 12.60 long, 9.40 wide. Length of leg segments: I 8.90 + 3.90 + 6.80 + 6.00 + 3.10 (28.70); II 8.90 + 3.80 + 6.10 + 6.00 + 3.00 (27.80); III 6.70 + 2.90 + 5.20 + 6.30 + 2.90 (24.00); IV 8.50 + 3.50 + 6.90 + 8.00 + 3.80 (30.70). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 51-54, 70-71): Carapace brown, densely covered with white setae and with a well-marked pattern of two wide yellowish brownish longitudinal paramedian bands. Sternum yellow, densely covered with white setae. Maxillae and labium brown, with yellow tips. Chelicerae dark brown, proximal half of frontal side and lateral sides densely covered with yellowish white setae. Abdomen: dorsum densely covered with white setae, with a brownish cardiac mark and a pattern consisting of brownish transverse stripes, patches and speckles; sides and venter yellow, densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets greyish yellow. All legs yellow brownish, densely covered with white setae. Palp yellow brownish, densely covered with white setae, with a tarsal claw.

  • Epigyne and vulva (Figs 61-62, 72-73): Epigynal atrium twice as long as wide, with straight subparallel lateral edges; posterior transverse plate markedly triangular, its posterior margin angular; spermathecae pear-shaped, their tips directed medio-anteriad, inclined towards each other.

  • Distribution: Only known from the type locality in southern Uzbekistan (Fig. 76) where this species occurs together with another large burrowing wolf spider: Zyuzicosa baisunica Logunov, 2010; see Logunov (2010) for further information on the latter species.

  • Figs 49-54.

    Karakumosa gromovi sp. nov.; male paratype (49-50) and female paratype (51-54). (49, 52) Body, ventral view. (50-51) Ditto, dorsal view. (53) Female prosoma, frontal view. (54) Female body, lateral view. Scale bars 1 cm (49-52, 54), 0.5 cm (53).

    img-z14-6_275.jpg

    Figs 55-58.

    Karakumosa gromovi sp. nov., bulbus of male paratype. (55) Ventral-retrolateral view. (56) Ventral view. (57) Apical view. (58) Retrolateral view. Scale bars 0.2 mm. Abbreviations as explained in Material and methods.

    img-z15-5_275.jpg

    Figs 59-69.

    Karakumosa gromovi sp. nov.; male paratypes (59-60, 64-69) and paratype female (61-63). (59-60) Bulbus, ventral view. (61) Epigyne, ventral view. (62) Vulva, dorsal view. (63) Female tarsus I, ventral view. (64, 66) Median tooth of MA, posterior view. (65) Cymbium, ventral view. (67) Embolic division, ventral view. (68) Bulbus, retrolateral view. (69) Cymbium tip, dorsal view. Scale bars 0.5 mm (59, 61), 0.3 mm (63, 65), 0.25 mm (62), 0.1 mm (60, 66-69), 0.05 mm (64). Abbreviations as explained in Material and methods.

    img-z16-1_275.jpg

    Figs 70-75.

    Karakumosa gromovi sp. nov.; male paratypes (74-75) and female paratype (70-73). (70) Body, dorsal view. (71) Ditto, ventral view. (72) Epigyne, ventral view. (73) Vulva, dorsal view. (74-75) Median tooth of MA, posterior view. Scale bars 1 cm (70-71), 0.5 mm (72), 0.25 mm (73), 0.1 mm (74-75).

    img-z17-1_275.jpg

    Karakumosa medica (Pocock, 1889) comb. nov. Figs 76-88
    Tarentula medica Pocock, 1889: 110, pl. 13, figs 1a-g (description of male and female; 1 adult male, 1 adult female and 1 immature syntypes in the BMNH).
    Hogna media. – Roewer, 1955b: 249 (misspelling).

  • Lectotype (designated here): BMNH, BM 1887.51; male; AFGHANISTAN, Meshed/Tirp-hul [‘Kalla-idast-dargarri'; c. 34°42′N, 61°03′E]; no date [apparently June 1884]; leg. [J.E.T.] Aitchison.

  • Paralectotypes: BMNH, BM 1887.51; 1 female, 1 immature; collected together with the lectotype.

  • Etymology: Although the author (Pocock, 1889) did not explain the origin of the name, the species epithet presumably is the Latin adjective ‘medicus, -a, -um' (= medical), likely referring to the fear of this spider expressed by local people; see under ‘Comments’ below.

  • Diagnosis: This species is similar to K. badkhyzica sp. nov. (Figs 39-48), but differs in having a wider proximal extension of the MA (Fig. 83), a longer epigynal atrium (Fig. 85) and a different conformation of the spermathecae: worm-shaped in K. medica (Fig. 86) and markedly swollen anteriorly in K. badkhyzica sp. nov. (Fig. 43). See also comments above under ‘Diagnosis’ of K. gromovi sp. nov.

  • Description: Male (lectotype). Measurements: Carapace 13.00 long, 10.00 wide. Eye sizes and interdistances: AME 0.55, ALE 0.50, PME 1.30, PLE 1.25, AME-AME 0.30, AME-ALE 0.20, PME-PME 1.00, PME-PLE 2.45. Width of anterior eye row 2.50, of second row 3.40, of third row 4.20. Clypeus height 0.35; chelicera length 5.00. Abdomen 9.00 long, 6.40 wide. Length of leg segments: I 13.00 + 5.30 + 11.00 + 12.50 + 6.00 (47.80); II 12.60 + 5.30 + 10.50 + 12.50 + 5.50 (46.40); III 11.40 + 4.60 + 8.60 + 12.70 + 5.50 (42.80); IV 13.80 + 4.50 + 11.20 + 16.20 + 6.50 (52.20). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 81-82): Specimen damaged and shabby. Carapace dark russet, almost without white setae. Sternum orange-coloured, densely covered with white setae. Maxillae orange-coloured. Labium orange-coloured, with a yellow tip. Chelicerae dark brown, their frontal sides covered with white setae. Abdomen: dorsum densely covered with white setae, with a yellow cardiac mark outlined by a brown line; sides and venter densely covered with yellowish white setae. Book-lung covers orange-coloured, densely covered with yellowish white setae. Spinnerets brown. All legs orange-coloured brownish, densely covered with white setae. Palps orange-coloured, densely covered with white setae.

  • Palp structure (Figs 83-84, 87-88; mirrored image of right palp): Acutely pointed synembolic lamellae slightly convergent towards each other; medina tooth markedly bifurcated; proximal extension wide and relatively short; inner plate transverse-ovoid; conductor triangular, pointed at its tip.

  • Female (paralectotype). Measurements: Carapace 14.70 long, 11.00 wide. Eye sizes and interdistances: AME 0.70, ALE 0.55, PME 1.70, PLE 1.20, AME-AME 0.35, AME-ALE 0.25, PME-PME 1.10, PME-PLE 3.10. Width of anterior eye row 3.00, of second row 4.25, of third row 5.10. Clypeus height 0.65; chelicera length 6.50. Abdomen 13.00 long, 9.50 wide. Length of leg segments: I 12.30 + 5.00 + 9.50 + 9.00 + 4.70 (40.50); II 11.50 + 5.30 + 8.80 + 9.20 (no segment) (?); III 10.30 + 5.20 + 7.40 + 9.50 + 5.20 (37.70); IV 13.30 + 5.50 + 9.80 + 13.00 + 6.10 (47.70). Leg formula: IV, I, II, III. Colouration in alcohol (Figs 77-80): Specimen damaged and shabby. Carapace dark russet, almost without white setae. Sternum orange-coloured, densely covered with white setae. Maxillae brownish orange-coloured. Labium brown-orange-coloured, with a yellow tip. Chelicerae dark brown, proximal part of frontal side and lateral sides covered with yellowish white setae. Abdomen damaged: dorsum densely covered with yellowish white setae, with no visible cardiac mark and colour pattern; sides and venter yellow, densely covered with yellowish white setae. Book-lung covers yellow, densely covered with yellowish white setae. Spinnerets brown. All legs orange-coloured, densely covered with yellowish white setae. Palps orange-coloured, densely covered with white setae.

  • Epigyne and vulva (Figs 85-86): Epigynal atrium three times as long as wide, with subparallel lateral edges; posterior transverse plate short, its posterior margin slightly bent (making the plate anchor-shaped); spermathecae tube-shaped, bent mediad and then directed anteriad.

  • Comments: In the original paper (Pocock, 1889: 111), the particulars of collecting the Tarentula medica types are described as follows: “…taken between Tirphul and Meshed in Persia”. Later in the text, based on the collector's observations (J.E.T.A.), Pocock (1889) added that “At Kalla-idast-dargarri, June 8th, amongst the brick debris of the old buildings, and running in and out of the clay fissures in the soil, this spider occurred in large numbers, much to the horror of the natives - the Afghans and Persians holding it in great fear, as, they say, should it drop any of its excreta on the skin, the result is acute erysipelas, usually ending in death”. Based on the map published by the Afghan Delimitation Commission (Aitchison, 1889), ‘Kalla-idast-dargarri' seems to be a site lying some 14 km WNW of the present-day Kohsān, on the left bank of the Hari River in the Herat Province of Afghanistan (c. 34°42′N, 61°03′E). This site is here accepted as the type locality of Tarentula medica. We failed to find the geographic name ‘Kalla-idast-dargarri’ on contemporary maps.

  • Distribution: Only known from the type locality in north-western Afghanistan (Fig. 76). The record of this species by Schmidt (1895: 450; sub. Lycosa medica) from Uzbekistan, Qarshi (= Karschi; c. 38°51′N, 65°47′E) is doubtful and needs verification, because it more likely can be attributed to either K. alticeps or K. gromovi sp. nov. Unfortunately the whereabouts of Schmidt's specimens is unknown and they may be lost.

  • Fig. 76.

    Localities of five species in the genus Karakumosa gen. nov.

    img-z18-1_275.jpg

    Figs 77-82.

    Karakumosa medica (Pocock, 1889); female paralectotype (77-80) and male lectotype (81-82). (77, 81) Body, dorsal view. (78) Ditto, lateral view. (79) Prosoma, frontal view. (80, 82) Body, ventral view. Scale bars 1 cm (77-78, 80-82), 0.5 cm (79).

    img-z19-1_275.jpg

    Karakumosa repetek sp. nov.
    Figs 76, 89-102

  • Holotype: ISEA, 001.4309; male; TURKMENISTAN, Repetek (c. 38°35′N, 63°11′E), on light near a house; 7.VIII.1972; leg. G.T. Kuznetsov.

  • Paratypes: ISEA, 001.4310; 2 males; from the type locality; 31.VIII.1972; leg. G.T. Kuznetsov. – ISEA, 001.4213; 1 female; from the type locality; 21.VII.1985; leg. V.Ya. Fet. Etymology: The species epithet is a noun in apposition referring to the name of the type locality, the Repetek Reserve in Turkmenistan.

  • Diagnosis: The male of K. repetek sp. nov. is most similar to that of K. badkhyzica sp. nov. (Figs 40-42, 46-48), but differs in having a longer and wider, spade-like proximal extension of the MA (Fig. 98) and a bifurcated median tooth (spatula-like proximal extension and finger-like median tooth in K. badkhyzica sp. nov.; Fig. 42 cf. Fig. 100). In the slightly sigmoid lateral edges of the epigynal atrium, the female of K. repetek sp. nov. is most similar to that of K. zyuzini sp. nov., from which it differs in the shape of the posterior transverse plate (dumbbell-shaped vs. with straight posterior margin; Fig. 96 cf. Fig. 179) and in the curved vs. straight spermathecae (Fig. 95 cf. Fig. 182).

  • Description: Male (holotype). Measurements: Carapace 10.80 long, 8.00 wide. Eye sizes and interdistances: AME 0.50, ALE 0.38, PME 1.10, PLE 0.90, AME-AME 0.13, AME-ALE 0.10, PME-PME 0.85, PME-PLE 1.05. Width of anterior eye row 1.90, of second row 2.70, of third row 3.35. Clypeus height 0.33; chelicera length 4.00. Abdomen 9.20 long, 5.30 wide. Length of leg segments: I 9.80 + 4.20 + 9.10 (distal segments missing); II 9.70 + 3.80 + 9.00 + 11.70 + 5.00 (39.20); III 9.70 (distal segments missing); IV 11.40 + 4.20 (distal segments missing).

  • Colouration in alcohol (Figs 89-90): Specimen damaged and shabby. Carapace yellow brownish, densely covered with white setae. Sternum yellow, densely covered with white setae. Maxillae and labium yellow, with white tips. Chelicerae yellowish brownish, their frontal and lateral sides densely covered with white setae. Abdomen: dorsum densely covered with white setae, with a poorly marked cardiac mark; sides and venter densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets yellow brownish.

  • All legs yellow brownish, densely covered with white setae. Palp yellow, densely covered with white setae.Palp structure (Figs 93-94, 97-102; mirrored image of right palp): Acutely pointed synembolic lamellae convergent to each other or bent proximad; median tooth bifurcated at its tip; proximal extension wide and long, spade-like; inner plate transverse and relatively narrow, with an extended, obtuse retrolateral-proximal shoulder; conductor triangular, pointed at its tip.

  • Female (paratype). Measurements: Carapace 9.20 long, 7.20 wide. Eye sizes and interdistances: AME 0.53, ALE 0.35, PME 1.03, PLE 1.05, AME-AME 0.28, AME-ALE 0.10, PME-PME 1.15, PME-PLE 1.15. Width of anterior eye row 2.10, of second row 2.80, of third row 3.35. Clypeus height 0.40; chelicera length 5.25. Abdomen 11.00 long, 7.30 wide. Length of leg segments: I-II absent; III 7.20 + 3.50 + 4.80 + 6.50 + 4.00 (26.00); IV absent.

  • Colouration in alcohol (Figs 91-92): Damaged specimen with only one legs remaining. Carapace brownish yellow, densely covered with white setae; its dorsal side with a brownish colour pattern consisting of two wide longitudinal paramedian bands. Sternum yellow, densely covered with white setae. Maxillae and labium yellow-brown, with yellow tips. Chelicerae brown, their frontal and lateral sides densely covered with white setae. Abdomen: dorsum densely covered with white setae, with a yellowish brown cardiac mark and a brownish colour pattern consisting of transverse stripes; sides and venter densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets yellow brownish. All legs brownish yellow, densely covered with white setae. Palps lost.

  • Epigyne and vulva (Figs 95-96): Epigynal atrium twice as long as wide, narrower at its proximal end and with slightly sigmoid lateral edges; posterior transverse plate dumbbell-shaped; spermathecae curved like a “C”, bent mediad and tips directed anteriad.

  • Distribution: Only known from the type locality, the Repetek Reserve in Turkmenistan (Fig. 76).

  • Figs 83-88.

    Karakumosa medica (Pocock, 1889); male lectotype (83-84, 87-88) and female paralectotype (85-86). (83) Bulbus, ventral view. (84) Ditto, retrolateral view. (85) Epigyne, ventral view. (86) Vulva, dorsal view. (87) Median tooth of MA, posterior view. (88) Synembolus, apical view. Scale bars 0.5 mm (83-86), 0.1 mm (87-88).

    img-z20-1_275.jpg

    Figs 89-92.

    Karakumosa repetek sp. nov.; male paratype (89-90) and female paratype (91-92). (89, 92). Body, dorsal view. (90, 91) Ditto, ventral view. Scale bars 0.5 cm.

    img-z21-1_275.jpg

    Karakumosa shmatkoi sp. nov.
    Figs 1035-141, 155
    Lycosa alticeps (Kroneberg, 1875). – Schmidt, 1895: 450 (partim, misidentification). – Dunin, 1984: 55. – Minoransky & Ponomarev, 1984: 85-86 (all misidentifications).
    Lycosa” sp. 1. – Ponomarev & Abdurakhmanov, 2014: 92-93.
    Lycosa” sp. 2. – Ponomarev & Abdurakhmanov, 2014: 93.

  • Holotype: ZMMU; male (Figs 105-106, 128, 132-133); KAZAKHSTAN, Atyrau Province, c. 20 km NE of Ganyushkino Vil., Zhuzguntyube stow (46°43′1.4″N, 49°25′32.1″E); 6.VII.1977; leg. F.A. Saraev.

  • Paratypes: ZMMU; 1 female; same locality as for the holotype; 1.-10.VII.1977; leg. F.A. Saraev. – ISEA; 2 males; KAZAKHSTAN, Atyrau Province, Embinsk Distr., near Tengiz (46°23′49.9″N, 53°25′22.6″E); 8.VI.1986; leg. F.A. Saraev. – ZMMU; 1 male; KAZAKHSTAN, Atyrau Province, Embinsk Distr., near Koschagyl (46°48′17.3″N, 53°43′50″E); 19.-24.VI.1987; leg. F.A. Saraev. – ZMMU; 6 males; KAZAKHSTAN, Mangistau Province, Mangyshlak, c. 10 km S of Fort Shevchenko, shore of Tyub-Karagan Peninsula (44°26′19.9″N, 50°15′19.9″E), hilly sands, pitfall traps; 17.VI.2013; leg. G.M. Abdurakhmanov. – ZMMU; 3 males; KAZAKHSTAN, Mangistau Province, Mangyshlak, Island Kulaly (44°53′31.4″N, 50°03′13.2″E), fine hilly sands, hand collecting; 19.VI.2013; leg. G.M. Abdurakhmanov. – MHNG; 2 males; same data. – PSU; 1 male; KAZAKHSTAN, Aktobe Province, c. 25 km ENE of Embi Vil., Mugodzhary Mts (c. 48°57′N, 58°39′E); 28.VI.2013; leg. A.O. Shkurikhin. – MMUE; 1 male; RUSSIA, Kalmykia, Yashkul Distr., c. 55 km NE of Yashkul Vil., near Utta Vil. (46°22′33.9″N, 46°01′18.5″E), hilly sands with sparse vegetation; 12.VI.1975; leg. A.V. Ponomarev. – ZMMU; 2 males; RUSSIA, Kalmykia, Chernozemel'sk Distr., c. 65 km SE of Komsomol'sky Vil., near Artezian Vil. (44°57′29.5″N, 46°37′53.5″E); 1.-4.VII.1976; leg. E.A. Khachikov.

  • Other material: ZMMU; 1 female (Figs 136-141); AZERBAIJAN, Absheron Peninsula (no exact locality); no date and collector given.

  • Etymology: The species is dedicated to our friend and colleague, Mr Vladimir Yu. Shmatko (Rostov-on-Don, Russia), who has been collaborating with the second author (AVP) in spider studies for many years and who took the majority of digital photos of Karakumosa species given in the present paper.

  • Diagnosis: The male of K. shmatkoi sp. nov. is most similar to those of K. gromovi sp. nov. (Figs 59, 60, 64-69) and K. tashkumyr sp. nov. (Figs 142-154); it can be distinguished from the former by the presence of a low serrate flange at the foot of the median tooth (absent in K. gromovi sp. nov.; Figs 125, 134 cf. Figs 74-75), and from the latter by a narrower inner plate of the median apophysis, which is almost hidden under the outer plate (Figs 118-119 cf. Figs 148, 150) and by the shape of the synembolus (Fig. 116 cf. Fig. 149). In the sigmoid lateral edges of the epigynal atrium, the female is similar to those of K. repetek sp. nov. (Figs 95-96) and K. zyuzini sp. nov. (Figs 179, 182), from which it differs in the shape of the posterior transverse plate (developed as a low, inverted triangle; Fig. 122 cf. Fig. 96 and Fig. 179) and by the spermathecae being visibly swollen (Fig. 121 cf. Fig. 95 and Fig. 182). See also comments above under ‘Diagnosis’ of K. alticeps.

  • Description: Male (holotype). Measurements: Carapace 11.00 long, 9.30 wide. Eye sizes and interdistances: AME 0.45, ALE 0.40, PME 0.90, PLE 0.90, AME-AME 0.38, AME-ALE 0.15, PME-PME 1.20, PME-PLE 1.15. Width of anterior eye row 2.25, of second row 2.95, of third row 3.65. Clypeus height 0.35; chelicera length 4.00. Abdomen 10.50 long, 7.20 wide. Length of leg segments: I 10.00 + 5.00 + 9.80 + 10.00 + 4.80 (39.60); II 11.00 + 4.50 + 9.50 + 10.30 + 4.90 (40.20); III 10.00 + 3.80 + 7.60 + 9.80 + 4.20 (35.40); IV 12.00 + 4.10 + 9.70 + 13.50 + 5.40 (44.70). Leg formula: IV, II, I, III.

  • Colouration in alcohol (Figs 103-106, 126-127): Carapace dark brown, densely covered with yellowish white setae, without two wide brownish longitudinal paramedian bands. Sternum yellow-orange-coloured, densely covered with white setae. Maxillae and labium yellowish brown, with yellow tips. Chelicerae dark brown, their frontal and lateral sides densely covered with yellowish white setae. Abdomen: dorsum densely covered with white setae, with an indistinct cardiac mark outlined by brown lines; sides and venter yellow, densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets white. All legs yellow brownish, densely covered with white setae; metatarsi and tarsi of all legs ventrally darker (brown). Palp yellow, densely covered with white setae.

  • Palp structure (Figs 107-112, 115-120, 123-125, 128-135): Acutely pointed synembolic lamellae visibly convergent; median tooth developed as a large median claw with additional small teeth on its prolateral side and a prominent serrate ventral flange; proximal extension wide and hook-shaped; distance between proximal extension and median tooth wide, equal to two proximal extension widths; inner plate large, transverse-ovoid, its retrolateral shoulder extended and bent ventrad; conductor triangular, obtuse at its tip (pointed in other specimens).

  • Female (paratype). Measurements: Carapace 11.50 long, 8.60 wide. Eye sizes and interdistances: AME 0.45, ALE 0.35, PME 1.15, PLE 1.00, AME-AME 0.30, AME-ALE 0.18, PME-PME 1.20, PME-PLE 1.45. Width of anterior eye row 2.00, of second row 3.15, of third row 3.90. Clypeus height 0.50; chelicera length 4.85. Abdomen 14.00 long, 11.00 wide. Length of leg segments: I 9.80 + 4.10 + 7.00 + 6.20 + 3.50 (30.60); II 8.70 + 3.50 + 6.70 + 6.30 + 3.60 (28.80); III 8.00 + 3.60 + 5.80 + 6.80 + 3.60 (27.80); IV 10.00 + 4.50 + 7.00 + 9.50 + 3.90 (34.90). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 113, 114): Damaged and shabby specimen with abdomen detached from carapace. Carapace brown, densely covered with yellowish white setae and with two longitudinal lateral bands of brownish setae; carapace sides with wide marginal bands of white setae. Sternum brownish, densely covered with white setae. Maxillae and labium brown, with yellowish tips. Chelicerae dark brown, proximal half of frontal side densely covered with yellowish white setae. Abdomen: dorsum densely covered with yellowish white setae and with a large, wide, brownish cardiac mark; sides and venter, including book-lung covers, densely covered with white setae. Spinnerets brown. All legs and palps brownish yellow, densely covered with white setae. Palps with a claw at their tips.

  • Epigyne and vulva (Figs 121-122, 139): Epigynal atrium twice as long as wide, with markedly sigmoid lateral edges (almost S-shaped); posterior transverse plate developed as a low, inverted triangle, its central part slightly elevated and pointed backwards; spermathecae straight and visibly swollen in anterior portion, directed antero-mediad, inclined towards each other.

  • Comments: The males of K. shmatkoi sp. nov. examined display a noticeable variation in the shape of the proximal extension of the pedipalp which has either an obtuse (Fig. 132) or pointed (Figs 118-119) median shoulder. In the latter case, the proximal extension varies in its width, being visibly narrower (Fig. 118) or wider (Fig. 119). It is possible that more than one closely related species are here placed under the name of K. shmatkoi sp. nov. Unfortunately, the available material does not allow us to scrutinize the problem further; currently only one variety is known from both sexes. The problem needs special attention in the future when more material of both sexes from a larger number of localities is available.

  • Distribution: Known from two localities in Kalmykia, Russia (Minoransky & Ponomarev, 1984: sub Lycosa alticeps; present data), and from several localities in western Kazakhstan (Ponomarev & Abdurakhmanov 2014: sub “Lycosa” sp. 1 & “Lycosa” sp. 2; present data) and Azerbaijan (Absheron Peninsula, Baku and Chilov Island) (Schmidt, 1895; Dunin, 1984: sub L. alticeps; present data) (Fig. 155).

  • Figs 93-102.

    Karakumosa repetek sp. nov.; male holotype (97-98), male paratype (93-94, 99-102) and female paratype (95-96). (93-94, 98). Bulbus, ventral view. (95) Vulva, dorsal view. (96) Epigyne, ventral view. (97) Median tooth of MA, ventral view. (99) Embolic division, ventral view. (100) Median tooth of MA, posterior view. (101) Embolic division, apical view. (102) Bulbus, retrolateral view. Scale bars 0.5 mm (93, 96), 0.25 mm (94, 102), 0.2 mm (98), 0.1 mm (95, 97, 99-101). Abbreviations as explained in Material and methods.

    img-z22-1_275.jpg

    Figs 103-114.

    Karakumosa shmatkoi sp. nov.; male holotype (105-106), male paratype (103-104, 107-112) and female paratype (113-114); specimens from Kulaly Island (103-104, 109), Zhuzguntyube stow (105-106, 113-114), Mugodzhary Mts (107), Artezian Vil. (108, 110), Tengiz (111), Tyub-Karagan Peninsula (112). (103, 105, 113) Body, dorsal view. (104, 106, 114) Ditto, ventral view. (107-112) Median tooth of MA, posterior view. Scale bars 1 cm (103-106, 113-114), 0.1 mm (107-112).

    img-z23-1_275.jpg

    Figs 115-125.

    Karakumosa shmatkoi sp. nov.; male paratypes (115-120, 123-125) and female paratype (121-122); specimens from Tyub-Karagan Peninsula (115-116, 118, 120, 123, 125), Island Kulaly (117), Zhuzguntyube stow (121-122) and Tengiz (119, 124). (115, 118-119) Bulbus, ventral view. (116) Embolic division. (117) Left male palp, ventral view. (120) Bulbus, retrolateral view. (121) Vulva, dorsal view. (122) Epigyne, ventral view. (123-125) Median tooth of MA, posterior view. Scale bars 0.25 mm (115, 117, 120, 122), 0.2 mm (118-119), 0.1 mm (116, 121, 123-125). Abbreviations as explained in Material and methods.

    img-z24-1_275.jpg

    Figs 126-135.

    Karakumosa shmatkoi sp. nov.; male holotype (128, 132-133) and male paratype from Kalmykia, Utta Vil. (126-127, 129-131, 134-135). (126) Body, dorsal view. (127) Ditto, ventral view. (128, 134) Median tooth of MA, posterior view. (129) Embolic division. (130) Left male palp, ventral view. (131-132) Bulbus, ventral view. (133) Synembolus, apical view. (135) Bulbus, retrolateral view. Scale bars 1 cm (126-127), 0.5 mm (130-131, 135), 0.2 mm (132), 0.1 mm (128, 134).

    img-z25-1_275.jpg

    Figs 136-141.

    Karakumosa shmatkoi sp. nov., female from Azerbaijan, Absheron peninsula. (136) Prosoma, frontal view. (137) Ditto, ventral view. (138) Ditto, lateral view. (139) Epigyne, ventral view. (140) Tarsus I, ventral view. (141) Tarsus IV, ventral view. Scale bars 2 mm (137-138), 1 mm (136), 0.5 mm (140-141), 0.2 mm (139).

    img-z26-1_275.jpg

    Figs 142-154.

    Karakumosa tashkumyr sp. nov., male paratypes, in ISEA. (142) Body, ventral view. (143) Ditto, dorsal view. (144) Left chelicera, ventral view. (145, 147, 152-154) Median tooth of MA, posterior view. (146) Left male palp, ventral view. (148, 150) Bulbus, ventral view. (151) Ditto, retrolateral view. (149) Embolic division. Scale bars 1 cm (142-143), 0.5 cm (144, 146, 150-151), 0.2 mm (148), 0.1 mm (145, 147, 149, 152-154).

    img-z28-1_275.jpg

    Karakumosa tashkumyr sp. nov.
    Figs 142-155

  • Holotype: ISEA, 001.8406; male; KYRGYZSTAN, Dzhalalabad Area, N of Taşkomur (=Tash-Kumyr) (c. 41°23′N, 72°14′E), clay hills; 14.-15.VI.1992; leg. A.A. Fedorov & A.A. Zyuzin.

  • Paratypes: ISEA; 7 males; collected together with the holotype. – ZMMU; 2 males; same data. – MMUE; 2 males; same data. – MHNG; 1 male; same data.

  • Etymology: The species epithet is a noun in apposition referring to the name of the type locality, Tash-Kumyr in Kyrgyzstan.

  • Diagnosis: In the presence of a low serrate flange at the foot of the median tooth of the MA, this species is most similar to K. shmatkoi sp. nov. (Figs 107-112, 115-120, 123-125, 128-135). The males of K. tashkumyr sp. nov. can be readily distinguished by the relative size of the inner plate of the MA, which in the ventral view is comparatively narrower and almost hidden beneath the outer plate in K. shmatkoi sp. nov. and wider and clearly visible in K. tashkumyr sp. nov. (Fig. 148 cf. Fig. 118 and Fig. 119), and by the shape of the synembolus: both tips of the lamellae markedly bent downwards in K. tashkumyr sp. nov. vs. upper tip straight in K. shmatkoi sp. nov. (Fig. 149 cf. Fig. 116 and Fig. 129). The female of K. tashkumyr sp. nov. is unknown.

  • Description: Male (paratype, ISEA). Measurements: Carapace 12.00 long, 8.60 wide. Eye sizes and interdistances: AME 0.55, ALE 0.40, PME 1.10, PLE 1.20, AME-AME 0.30, AME-ALE 0.20, PME-PME 1.00, PME-PLE 1.50. Width of anterior eye row 2.55, of second row 3.35, of third row 4.00. Clypeus height 0.43; chelicera length 5.50. Abdomen 10.50 long, 6.80 wide. Length of leg segments: I 11.50 + 5.10 + 11.00 + 12.40 + 5.20 (45.20); II 11.50 + 4.80 + 10.20 + 12.40 + 5.40 (44.30); III 11.00 + 4.40 + 8.80 + 12.50 + 5.50 (42.20); IV 13.00 + 4.70 + 10.10 + 15.40 + 6.00 (49.20). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 142-143): Carapace brown, densely covered with yellowish white setae and with two wide brownish longitudinal bands. Sternum yellow-orange-coloured, densely covered with white setae. Maxillae half brownish and half yellow-orange-coloured. Labium brownish yellow, with a yellow tip. Chelicerae dark brown, their frontal and lateral sides densely covered with white setae. Abdomen: dorsum densely covered with white setae, with an indistinct brownish cardiac mark; sides and venter yellow, densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets yellow brownish. All legs yellow brownish, densely covered with white setae; metatarsi and tarsi of all legs ventrally darker (brownish). Palps yellow-orange-coloured, densely covered with white setae.

  • Palp structure (Figs 145-154): Acutely pointed synembolic lamellae bent downwards; median tooth developed as a large median claw with additional small teeth on its prolateral side and a prominent ventral flange looking like an additional tooth; proximal extension wide and hook-shaped; distance between proximal extension and median tooth wide, equal to two proximal extension widths; inner plate large, transverse-ovoid, its retrolateral shoulder bent ventrad; conductor triangular, obtuse at its tip.

  • Female. Unknown.

  • Distribution: Only known from the type locality, north of Taşkomur in Dzhalalabad Area of Kyrgyzstan (Fig. 155).

  • Karakumosa turanica sp. nov.
    Figs 155-168
    Tarentula alticeps Kroneberg, 1875: 40, pl. 4, fig. 28 (partim; misidentification of paralectotypes from Uzbekistan).
    Lycosa alticeps (Kroneberg, 1875). – Simon, 1899: 479. – Vlassov & Sytshevskaja, 1937: 250. – Mikhialov & Fet, 1994: 508 (misidentification of specimens from Ashgabat).

  • Holotype: ZISP; 1 male; TURKMENISTAN, [Balkan Velayat], between Serdar (=Kyzyl-Arvat) and Iskander (c. 39°03'N, 56°07'E); 6.VI.1936; leg. S. Nenyukov.

  • Paratypes: ZISP; 2 females; “ex vil. Melnikova” near Ashgabad (c. 37°59′N, 58°20′E); summer and autumn, 1933; collector unknown.

  • Other material: ZMMU, Ta-1215; 1 female (paralectotype of Tarentula alticeps; Figs 165, 168); [UZBEKISTAN], “Ulus” [c. 39°34′N, 66°22′E]; no date; [A.P.] Fedchenko (Turkestan scientific expedition of the Imperial Society of Devotees of Natural Science). – ZMMU, Ta-1216; 1 subadult male, 3 immature females (paralectotypes of Tarentula alticeps); [UZBEKISTAN], Samarkand [c. 39°37′N, 66°57′E]; no date; leg. [A.P.] Fedchenko (Turkestan scientific expedition of the Imperial Society of Devotees of Natural Science).

  • Etymology: The specific epithet is latinized adjective derived from the name ‘Turan’, a historic region of Central Asia.

  • Diagnosis: The male of K. turanica sp. nov. is most similar to that of K. alticeps, but can be easily distinguished by a markedly pointed and wider median shoulder of the proximal extension (narrower and obtuse in K. alticeps; Fig. 161 cf. Fig. 2 and Fig. 12) and by the larger, quadrangular median tooth (smaller, triangular median tooth with a serrate prolateral edge in K. alticeps; Fig. 164 cf. Figs 5-8). The female of K. turanica sp. nov. is most similar to that of K. zyuzini sp. nov., from which it can be distinguished by an anchor-shaped posterior transverse plate (parallel-sided in K. zyuzini sp. nov.; Figs 167-168 cf. Fig. 179) and by spermathecae directed anteriad (inclined antero-mediad in K. zyuzini sp. nov.; Figs 165-166 cf. Fig. 182).

  • Description: Male (holotype). Measurements: Carapace 10.70 long, 8.50 wide. Eye sizes and interdistances: AME 0.45, ALE 0.45, PME 1.20, PLE 1.15, AME-AME 0.35, AME-ALE 0.15, PME-PME 1.00, PME-PLE 2.15. Width of anterior eye row 2.35, of second row 3.15, of third row 3.80. Clypeus height 0.45; chelicera length 4.70. Abdomen 8.80 long, 5.80 wide. Length of leg segments: I 11.10 + 4.00 + 9.40 + 11.50 + 4.60 (40.60); II 10.80 + 4.40 + 8.50 + 10.50 + 4.60 (38.80); III 11.10 + 3.80 + 7.50 + 11.10 + 4.30 (37.80); IV 12.10 + 3.80 + 9.10 + 12.60 + 5.50 (43.10). Leg formula: IV, I, II, III. Colouration in alcohol (Figs 156-158): Specimen in poor condition, damaged, shabby, with some legs and left palp missing. Carapace dark brown, densely covered with yellowish white setae; colour pattern of longitudinal brown stripes indiscernible due to poor condition. Sternum brown, densely covered with yellowish white setae. Maxillae and labium yellowish brown. Chelicerae dark brown, their frontal and lateral sides densely covered with yellowish white setae. Abdomen: dorsum densely covered with grey yellowish setae, with a poorly marked brownish cardiac mark; the frontal part of abdomen brownish; sides and venter, including book-lung covers, brownish, densely covered with yellowish white setae. Spinnerets yellow light brown. All legs light brown, densely covered with yellowish white setae. Palps yellow-orange-coloured, densely covered with yellowish white setae.

  • Palp structure (Figs 161-164; mirrored image of right palp): Acutely pointed synembolic lamellae markedly convergent towards each other; median tooth large and wide; proximal extension wide and pointed at its median angle, distance between proximal extension and median tooth narrow, equal to or less than a proximal extension width; inner plate rather narrow, almost hidden beneath outer plate; conductor triangular, pointed at its tip.

  • Female (paratype). Measurements: Carapace 12.60 long, 9.60 wide. Eye sizes and interdistances: AME 0.55, ALE 0.60, PME 1.25, PLE 0.65, AME-AME 0.30, AME-ALE 0.15, PME-PME 1.40, PME-PLE 2.60. Width of anterior eye row 2.90, of second row 3.60, of third row 4.35. Clypeus height 0.45; chelicera length 6.60. Abdomen 10.00 long, 8.60 wide. Length of leg segments: I 10.00 + 4.60 + 8.00 + 7.50 + 4.00 (34.10); II 9.50 + 4.40 + 6.90 + 7.60 + 4.00 (32.40); III 8.50 + 3.90 + 6.30 + 7.70 + 3.80 (30.20); IV 10.80 + 4.50 + 8.30 + 11.00 + 4.70 (39.30). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 159-160): Carapace brown, densely covered with yellowish white setae, without two longitudinal paramedian bands of brownish setae; carapace margins with wide bands of yellowish white setae. Sternum brown, densely covered with yellowish white setae. Maxillae and labium brown, with yellowish tips. Chelicerae dark brown, proximal part of frontal side densely covered with yellowish white setae. Abdomen: dorsum brownish, densely covered with yellowish white setae, with a poorly marked brownish cardiac mark; sides and venter, including book-lung covers, densely covered with yellowish white setae. Spinnerets brown. All legs and palps brownish yellow, densely covered with white setae; tips of all leg tarsi darker (brown). Palps with a claw at their tips.

  • Epigyne and vulva (Figs 165-168): Epigynal atrium twice as long as wide, with slightly sigmoid lateral edges; posterior transverse plate narrow, anchor-shaped, its posterior margin widely procurved; spermathecae tube-shaped and straight, directed anteriad.

  • Comments: Male and female specimens of K. turanica sp. nov. were not collected together and hence are matched provisionally, on the ground that (1) both sexes were collected from lowland sites of Turkmenistan lying relatively close to each other (Fig. 155), and (2) two other Karakumosa species known from the plains of Turkmenistan (K. badkhyzica sp. nov. and K. repetek sp. nov.; Fig. 76) are described from both sexes collected together; these are distinct from K. turanica sp. nov.

  • The female paralectotype of K. alticeps (Figs 165, 168) turned out to be conspecific with female paratypes of K. turanica sp. nov. (Figs 166-167), and therefore its record from Samarkand is included in the account of K. turanica sp. nov. See also ‘Comments’ above under K. alticeps.

  • Distribution: Known from a few localities in Turkmenistan and Uzbekistan (Fig. 155), but it is believed that the species is more widespread across the plains of Central Asia, within the so-called Turan zoogeographic province (sensu Kryzhanovsky, 2002). Besides, some of the old records of Lycosa alticeps from Turkmenistan by Schmidt (1895) could also belong to this species: viz., the records from Uch-Adzhi (c. 38°05′N, 62°48′E) and Turkmenbashi (c. 40°02′N, 52°59′E).

  • Fig. 155.

    Localities of four species in the genus Karakumosa gen. nov.

    img-z30-1_275.jpg

    Figs 156-160.

    Karakumosa turanica sp. nov., male holotype (156-158) and female paratype (159-160). (156, 160) Body, dorsal view. (157, 159) Ditto, ventral view. (158) Ditto, lateral view. Scale bars 1 cm.

    img-z31-1_275.jpg

    Karakumosa zyuzini sp. nov.
    Figs 155, 169-186

  • Holotype: ISEA, 001.8405; male; UZBEKISTAN, Navoiy Region, Kanimekh Distr., near Chengeldy (c. 40°56′44.1″N, 64°18′06.0″E), flat clay clough between sandy plots; 22.-23.V.1994; leg. A.A. Zyuzin.

  • Paratypes: ISEA, 001.4188; 11 males, 3 females; collected together with the holotype. – ZMMU; 2 males, 2 females; same data. – MMUE; 3 males, 3 females; same data. – MHNG; 2 males, 2 females; same data.

  • Etymology: The species is dedicated to our friend and colleague, Dr Alexei A. Zyuzin (Almaty, Kazakhstan), an expert on the Lycosidae, who collected the type specimens of this species.

  • Diagnosis: The male of K. zyuzini sp. nov. can be readily distinguished from males of all other known Karakumosa species by the shape of the large, poorly marked proximal extension of the MA (Figs 174, 176). The female of K. zyuzini sp. nov. is most similar to that of K. repetek sp. nov. from which it differs in the shape of the posterior transverse epigynal plate (with straight posterior margin vs. dumbbell-shaped, Fig. 179 cf. Fig. 96) and the straight vs. C-curved spermathecae (Fig. 182 cf. Fig. 95). See also comments under ‘Diagnoses’ of K. shmatkoi sp. nov. and K. turanica sp. nov.

  • Description: Male (paratype, MMUE). Measurements: Carapace 10.50 long, 8.00 wide. Eye sizes and interdistances: AME 0.51, ALE 0.35, PME 1.00, PLE 0.90, AME-AME 0.25, AME-ALE 0.30, PME-PME 1.25, PME-PLE 1.40. Width of anterior eye row 2.00, of second row 3.00, of third row 3.90. Clypeus height 0.37; chelicera length 4.30. Abdomen 10.50 long, 6.30 wide. Length of leg segments: I 11.00 + 4.00 + 9.90 + 9.80 + 5.00 (39.70); II 10.50 + 3.80 + 9.10 + 10.00 + 4.80 (38.20); III 9.30 + 3.30 + 7.10 + 10.50 + 4.10 (34.30); IV 11.50 + 3.80 + 9.60 + 13.80 + 5.00 (43.70). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 169-170): Carapace brown, densely covered with white setae and with two poorly-marked wide brownish longitudinal paramedian bands. Sternum yellow, densely covered with white setae. Maxillae brownish yellow. Labium brownish yellow, with yellow tip. Chelicerae brown, their frontal and lateral sides densely covered with white setae. Abdomen: dorsum densely covered with white setae, with a brownish colour pattern consisting of longitudinal lines outlining cardiac mark and transverse lines in rear half of dorsum; sides and venter yellow, densely covered with white setae. Book-lung covers yellow, densely covered with white setae. Spinnerets yellow brownish. All legs yellow brownish, densely covered with white setae; metatarsi and tarsi of all legs ventrally darker (brownish). Palp yellow brownish, densely covered with white setae. Palp structure (Figs 173-178, 180-181, 183-186): Acutely pointed synembolic lamellae both curved downwards and subparallel to each other; median tooth bifurcated, with a serrated prolateral edge; proximal extension large and ovoid, not distinct from margin of outer plate; inner plate large, transverse-ovoid; conductor triangular, pointed at its tip.

  • Female (paratype, MMUE). Measurements: Carapace 12.00 long, 9.70 wide. Eye sizes and interdistances: AME 0.50, ALE 0.40, PME 1.00, PLE 1.00, AME-AME 0.40, AME-ALE 0.25, PME-PME 1.40, PME-PLE 1.60. Width of anterior eye row 2.40, of second row 3.00, of third row 4.25. Clypeus height 0.65; chelicera length 6.20. Abdomen 16.20 long, 12.00 wide. Length of leg segments: I 10.00 + 4.20 + 8.10 + 6.50 + 3.70 (32.50); II 9.40 + 4.10 + 7.40 + 7.10 + 3.60 (31.60); III 8.50 + 3.90 + 6.10 + 7.50 + 3.90 (29.90); IV 11.00 + 4.00 + 8.60 + 10.50 + 4.60 (38.70). Leg formula: IV, I, II, III.

  • Colouration in alcohol (Figs 171-172): Carapace dark brown, densely covered with yellowish white setae and with a poorly marked pattern of two wide brownish longitudinal paramedian bands. Sternum yellow-orange-coloured, densely covered with white setae. Maxillae and labium brown, with yellow tips. Chelicerae dark brown, their frontal and lateral sides densely covered with yellowish white setae. Abdomen: dorsum densely covered with yellowish white setae, with a brownish cardiac mark; sides and venter yellow, densely covered with white setae. Book-lung covers yellow, covered with white setae. Spinnerets yellow brownish. All legs yellow brownish, densely covered with white setae; metatarsi and tarsi of all legs darker (brownish). Palp yellow brownish, densely covered with white setae, carrying a tarsal claw.

  • Epigyne and vulva (Figs 179, 182): Epigynal atrium twice as long as wide, with almost subparallel, slightly sigmoid lateral edges; posterior transverse plate narrow and straight; spermathecae tube-shaped, slightly swollen in anterior part and straight, directed antero-mediad, inclined towards each other.

  • Distribution: Only known from the type locality, Chengeldy in Navoiy Region of Uzbekistan (Fig. 155).

  • Figs 161-168.

    Karakumosa turanica sp. nov., male holotype (161-164), female paratype (166-167) and paralectotype of Tarentula alticeps newly identified as K. turanica (165, 168). (161) Bulbus, ventral view. (162) Ditto, retrolateral view. (163) Synembolus, apical view. (164) Median tooth of MA, posterior view. (165-166) Vulva, dorsal view. (167-168) Epigyne, ventral view. Scale bars 0.25 mm (161-162, 166), 0.2 mm (165, 167-168), 0.1 mm (163-164).

    img-z32-1_275.jpg

    Figs 169-172.

    Karakumosa zyuzini sp. nov., male paratype (169-170) and female paratype (171-172), in MMUE. (169, 171) Body, ventral view. (170, 172) Ditto, dorsal view. Scal bars 1 cm.

    img-z33-1_275.jpg

    Figs 173-186.

    Karakumosa zyuzini sp. nov., male paratype (173-176, 178, 180-181, 183-186) and female paratype (177, 179, 182), in MMUE. (173-174, 176) Bulbus, ventral view. (175) Embolic division. (177) Left female chelicera, ventral view. (178) Bulbus, retrolateral view. (179) Epigyne, ventral view. (180-181, 183-186) Median tooth of MA, posterior view. (182) Vulva, dorsal view. Scale bars 0.5 mm (173, 176-179), 0.25 mm (182), 0.2 (174), 0.1 mm (175, 180-181, 183-186).

    img-z34-1_275.jpg

    DISCUSSION

    Karakumosa gen. nov. is part of a rather diverse and poorly studied fauna of the burrying wolf spiders of Central Asia. To date 30 species in nine genera of fossorial Lycosidae have been recorded/described from Central Asia (Table 1), but their regional diversity is likely to be much higher. The only comparable regional Palaeartic fauna of fossorial lycosids is that of Europe, including the Mediterranean, from where 32 species in five genera are currently known (Nentwig et al., 2020). The striking difference between the Central Asian and European faunas is the diversity of Lycosa Sundevall, 1833 accounting for 25 species in Europe and only three in Central Asia. At least 12 west-Mediterranean species of the tarantula-group of Lycosa have been recently revised (Planas et al., 2013) and thus the observed regional differences in Lycosa diversity are not accidental. Each region has a unique, monotypic genus of fossorial lycosids: Donacosa Alderweireldt & Jocqué, 1991 known only from south-western Spain (see Alderweireldt & Jocqué, 1991) and Oculicosa confined to the Ustyurt Plateau and the Kyzylkum Desert (see Logunov & Gromov, 2011).

    Currently only three species of fossorial lycosids occur between Europe and Central Asia (Marusik et al., 2000; World Spider Catalog, 2020; Zyuzin & Logunov, 2000): viz., the trans-Palaearctic Alopecosa fabrilis (Clerck, 1757), the eastern European - Central Asian Allohogna singoriensis (Laxmann, 1770) and the east-Mediterranean - Central Asian Lycosa praegrandis C.L. Koch, 1836. However, the taxonomy and distributional ranges of all of them are still in need of revision (e.g., see comments at the end of Table 1). Two common species are restricted to the eastern geographic limits of Europe only: Geolycosa dunini Zyuzin & Logunov, 2000 known from Transcaucasia (see Otto, 2020) and Karakumosa shmatkoi sp. nov. known from the eastern Caucasus and cis-Caspian regions (present data).

    Among the Central Asian fossorial lycosids (Table 1) 13 species (42%) are only known from the respective type localities and/or the original descriptions. This can partly be attributed to the inadequate knowledge of Central Asian Lycosidae, but it could also reflect the presence of truly restricted ranges in many fossorial lycosids (e.g., McCrone, 1963; Fairweather, 1993; Marshall, 1995; etc.) as a result of their lower vagility and habitat specificity. These species are often confined to certain (semi)desert habitat types. In the absence of further survey data to clarify biogeographical and biological information about the majority of fossorial lycosids from Central Asia, except for A. singoriensis (see Wagner, 1868; Marikovski, 1956), the underlying causes of currently recognized localised distribution patterns in these spiders will remain unresolved.

    Some earlier records of fossorial lycosids from that region are doubtful. For instance, Hogna bergsoei was described (but not illustrated) by Thorell (1875a: sub Tarentula bergsoei) from a single male from Derbent (Dagestan). Later it was reported from Baku (Azerbaijan) (L. Koch, 1878: sub Lycosa bergsoei). Based only on similarity in the body colouration of the five studied females from Turkmenistan (Kuldzha, Ashgabat and Kyzylkum desert) and the type described by Thorell, Schmidt (1895: 447) unreasonably identified and described them as H. bergsoei. Almost a hundred years later Dunin (1984) confirmed the occurrence of H. bergsoei in the Absheron Peninsula (Azerbaijan), but provided no illustration of the specimens he had studied. Neither Dunin (1984) nor previous authors explained how they were able to identify this species without a re-examination of its type. Despite the fact that these records have been accepted by some regional spider catalogues (Abdurakhmanov et al., 2012; Mikhailov, 2013) and by the World Spider Catalog (2020) giving the distribution of this species as ‘Caucasus (Russia, Azerbaijan), Central Asia’, it is obvious that the taxonomic validity and all post-description records of H. bergsoei are in need of revision.

    The case of Vesubia vivax (Thorell, 1875a) is similar to that of H. bergsoei. The species was described by Thorell (1875a, b: sub. Tarentula vivax) from a single female taken from an unknown locality in ‘Südrussland’ (German: southern Russia). Later Schmidt (1895: 457, sub Lycosa vivax) recorded this species from Nukus (Uzbekistan) on the basis of a male and female which he decided to be “in complete agreement with Thorell's description” apart from some varying measurements. It is unclear why the World Spider Catalog (2020) lists the distribution of V. vivax as ‘Ukraine, Russia (Europe), Turkmenistan (?)’, since neither Ukraine nor Turkmenistan were mentioned in any papers dealing with this species. As in the previous example, the taxonomic validity and the post-description records of V. vivax are in need of revision. Based on unpublished information from the colleague who re-examined the holotype of V. vivax (Yu. M. Marusik, personal communication, 27 March 2020), this species is actually a member of the genus Alopecosa.

    Table 1.

    The fossorial Lycosidae of Central Asia and their distribution.

    img-z36-2_275.gif

    img-z37-1_275.gif

    ACKNOWLEDGEMENTS

    We are most grateful to the following colleagues who provided help during the preparation of the present work: Mr Alexander V. Gromov (Bingen-am-Rhein, Germany) and Dr Alexei A. Zyuzin (Almaty, Kazakhstan) for tracing some localities of the studied species, Dr Anton A. Nadolny (Sevastopol, Russia) for critical commenting on an earlier draft of the manuscript, Dr Yuri M. Marusik (Magadan, Russia) for taking some SEM micrographs and digital photos (Figs 9-11, 17-21) and for critical comments on the early manuscript version, Dr Galina N. Azarkina (Novosibirsk, Russia) for producing the maps (Figs 76, 149) and commenting on an early manuscript version, Mr Tony Hunter (Liverpool, UK) for allowing one of us (DVL) to use the digital facilities at the World Museums of Liverpool (UK), Mr Phillip Rispin (Manchester, UK) for help with digital photography of three Karakumosa species (Figs 22-26, 113-114, 150-154), and Mr Alexander A. Fomichev (Novosibirsk, Russia) for sharing additional records of K. alticeps and allowing us to use some of his digital photos (Figs 12-16). We also wish to sincerely thank Mr Vladimir Yu. Shmatko and Mr Alexander V. Nazarenko (both Rostov-on-Don, Russia) for their kind assistance and help with producing most of the digital photos and SEM micrographs. Finally, Drs Volker W. Framenau (Murdoch, Australia) and Peter J. Schwendinger (Geneva, Switzerland) are acknowledged for their critical comments that help to improve the final manuscript version.

    The research by the second author (AVP) was conducted within the framework of the state task to the Southern Scientific Centre of the Russian Academy of Sciences, under project no. AAAA-A19-119011190176-7.

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    Accepted: 6 September 2020; Published: 28 October 2020
    KEYWORDS
    descriptions
    identification key
    taxonomy
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