A revision of the spider genus Anemesia (Araneae, Cyrtaucheniidae)

The Central Asian spider genus Anemesia Pocock, 1895 is rediagnosed and revised. The genus was found to contain 14 species: ♂♀ A. andreevae sp. nov. (Uzbekistan, Tajikistan); ♂♀ A. birulai (Spassky, 1937) (Turkmenistan); ♂♀ A. castanea sp. nov.; ♂♀ A. incana Zonstein, 2001, ♂♀ A. infumata sp. nov.; ♂♀ A. infuscata sp. nov.; ♂♀ A. karatauvi (Andreeva, 1968) (all Tajikistan); ♂ A. koponeni Marusik, Zamani & Mirshamsi, 2014 (Iran); ♂♀ A. oxiana sp. nov.; ♂♀ A. pallida sp. nov.; ♂ A. parvula sp. nov. (all Tajikistan); ♂♀ A. pococki sp. nov. (Turkmenistan); ♂♀ A. sogdiana sp. nov. (Uzbekistan, Tajikistan) and ♂♀ A. tubifex (Pocock, 1889), the type species (Afghanistan, Turkmenistan). Nine species are newly described; others are redescribed from types and/or conspecific material. Males of A. tubifex and females of A. birulai, hitherto unknown, are described for the first time. Data on the variability, relationships, distribution, and ecology of all considered species are provided.


Introduction
The genus Anemesia was established by Pocock (1895) as a monotypic taxon comprising the single species, Nemesia tubifex which he had described a few years earlier (1889) from zoological material of the British Afghan delimitation commission. When establishing the new genus, Pocock used as a distinguishing character the fact that the burrow of the only collected female had an open entrance, whereas all then known Mediterranean representatives of Nemesia Audouin, 1826 were considered to build burrows with a trapdoor. The genus was originally placed in the family Ctenizidae Thorell, 1887. Simon (1903) assigned Anemesia to the ctenizid genera of the tribe Nemesiae Simon, 1889. For several decades, Anemesia was known only from the holotype of N. tubifex which had also been heavily damaged prior to the description, presumably when collected. Denis (1958) erroneously (see below) assigned a few female mygalomorphs collected in Afghanistan to this species. He noted that these spiders were considerably larger than the holotype. Most generic features, including the shape of the spinneret group, configuration of the female spermathecae and all male characters, were unknown until 2001. For almost a century the definitive characters of Anemesia were unclear. Raven (1985), who first elevated Simon's Nemeseae to family rank, placed the monotypic Anemesia in synonymy with Nemesia in the absence of reliable data that could indicate a group, rather than an individual character. Zonstein (1987) considered two of seven Central Asian species described in Brachythele Ausserer, 1871 (treated in a very broad sense until 1985) as belonging to Anemesia, however without providing a formal transfer and using the new combinations. These species, B. birulai and B. karatauvi, described by Spassky (1937) and  from Turkmenistan and Tajikistan, respectively, were first listed by Mikhailov (1996) as members of the nemesiid Anemesia. Later, the finding of adult male congeners enabled a redefinition of the generic diagnosis; Anemesia was assigned to the genera of the Cyrtaucheniidae Simon, 1889; an additional species, A. incana, was described concurrently . The fifth known member of the genus, A. koponeni Marusik, Zamani & Mirshamsi, 2014, was very recently described from Iran . Furthermore, several undescribed species of Anemesia were found in the spider material collected in the period 1986-1993 (with some further European Journal of Taxonomy 485: 1-100 (2018)

Structure
Structure, sectioning, and headers follow Schwendinger (2009), Schwendinger & Zonstein (2011) and Zonstein et al. (2018). To maintain the uniformity of species descriptions, males are described first, even in cases where the holotype is female.

Terminology
The terminology and descriptive format follows Raven (1981Raven ( , 1985, with a few exceptions. The term megaspines was first used by Raven (1981) to designate thickened and curved thornlike spines located on anterior leg tibiae in male mygalomorphs that form the mating spurs. The terms intercheliceral tumescence and preening combs correspond to a 'pallid and soft area at the lower basal interface of the chelicerae […] of some male mygalomorphs' (Raven 1985: 8) and to the apical combs of bristles located in some mygalomorphs on the metatarsi, respectively. The nomenclature of the spigot types is given according to Murphy & Roberts (2015).

Illustrations
Photographs were taken using a Canon 500D digital camera with a 100 mm Canon macro lens (majority of the totals) and a Zeiss Discovery V20 stereo microscope with a Canon PowerShot G9 digital camera attached to it, and prepared using the Helicon Focus ver. 6.3.2 Pro software (http://www.heliconsoft.com/). Field photographs showing the biotopes, live spiders, burrows and their aggregations were taken mainly with a Canon PowerShot G1 X Mark II camera. Scanning electron micrographs were made using a JEOL JSM-5200 scanning microscope at the Zoological Museum, University of Turku, Finland. Distribution maps were drawn using ®Adobe Illustrator CS5. Photographs and other figures were processed, grouped and arranged using ®Adobe Photoshop CS5. Female genitalia were dissected under a Leica M80 stereo microscope. Illustrations of the spermathecae were made following the maceration of the dissected female genital area in a 10% potassium hydroxide aqueous solution, cleaning in 85% lactic acid and exposure for a few minutes in a 70% alcohol solution of Chlorazol Black. Upon becoming transparent and gently stained, these structures were then placed in a dish with 85% lactic acid and fixed on a glass microsphere layer for image-making. Following examination, the preparations were removed, cleaned and then transferred to small microvials and placed together with the studied specimens.

Measurements
Measurements were taken through a Zeiss Discovery V20 stereo microscope to an accuracy to 0.01 mm. All measurements are given in millimetres. The total body length includes the chelicerae, but not the spinnerets. The sternum length was measured along a straight line between the posterior tip of the sternum and the hindmost part of the labium. The lengths of leg and palp segments were measured on the dorsal side, and the lengths of spinneret segments on the ventral side, from midpoint of anterior margin to midpoint of posterior margin. The diameter of the AME is usually given as the diameter of a sharply edged AME circle (the 'pupil'). When the AME cornea was well-separated and elevated, and its diameter could be measured, the corresponding data follow in brackets. Any eye interdistances incorporating this parameter are also given in brackets.

Geographical and ecological data
The altitude is shown in meters above sea level. Coordinates are given for material collected after 1994-1995 according to the GPS data; and before this date according to the corresponding former Soviet military maps of 1:100000 or 1:200000 scale. When used without brackets, the coordinates refer to the collecting place(s). The coordinates in brackets refer to the indicated nearest locality and follow it. When necessary, the earlier obtained data were checked, specified and improved, using Google Earth 4.2™. Names and limits of orographic and biogeographic units are given according to Kryzhanovsky (2002) and Nikolaev (2002). Some of the dug spider burrows were studied and measured, and fragments ZONSTEIN S., Revision of Anemesia (Araneae, Cyrtaucheniidae) of the prey consumed by spiders were obtained. The prey remains were identified by Y.S. Tarbinsky and S.V. Ovchinnikov (Institute of Biology, Kyrgyz Academy of Sciences).

Emended diagnosis
The genus differs from other known cyrtaucheniids in the following unique combination of characters: (a) a relatively broad carapace which is weakly if at all constricted posteriorly; (b) a fairly short and straight thoracic fovea; (c) a protruding triangular clypeus; (d) a long and narrow posterior pair of the sternal sigilla; (e) among types of the spigots only AC can be identified, while MAC are absent. Although these features can occur individually in other genera of the Cyrtaucheniidae, such a combination has no analogs within the family.
Prosoma. Carapace broadly oval, mostly hirsute in males, glabrous to slightly hirsute in females. Clypeus triangular. Thoracic fovea transverse, straight or weakly procurved. Eye tubercle, normally carrying eight eyes (posterior eyes sometimes reduced), low to very low in females, more developed in conspecific males, mound carrying AMEs always distinct. Chelicerae with rastellum composed of numerous spikes on dorsodistal cheliceral edge. Cheliceral furrow with large promarginal teeth and with smaller retromarginal teeth, minute to medium-sized in females, but reduced in size in conspecific males. Fang without serration. Male intercheliceral tumescence absent. Labium moderately long, trapezoidal to subquadrate, without cuspules. Sternum domed, narrowed anteriorly. Sternal sigilla mostly distant from sternal margin, elongate, sometimes poorly developed (in A. infumata sp. nov. and A. pallida sp. nov.). Posterior sternal sigilla generally long and narrowly oval. Maxillae broadly trapezoidal. Few to numerous maxillary cuspules confined to probasal edge of maxillae. Sexual dimorphism in size of cuspules absent. Serrula not evident.
structures of legs i-iV. Leg formula: 4123 or 1423. Femur I usually longer than femur IV. Legs equal in width or legs III and IV slightly stouter (in females of A. andreevae sp. nov.). Male tibia I with one retroventral megaspine. Metatarsal preening combs absent. Tarsi entire. Most species of Anemesia have spines on tarsi III-IV (females of A. pococki sp. nov. and A. tubifex possess spines only on tarsus III; in females of A. andreevae sp. nov. all tarsi are aspinose; in males of A. castanea sp. nov. and A. parvula sp. nov. in addition to spinose tarsi III-IV, spines are located also on tarsus II). Scopula thin, moderately long and dense; entire and distal on metatarsi I and II; entire or narrowly divided on tarsi I and II, and on female palpal tarsus; widely divided, mixed with setae or reduced to absent on tarsi III and IV in males; usually absent on tarsi III and IV in females. Trichobothria arranged in two converging rows on tibiae, one straight row on metatarsi and one relatively narrow zigzag row on tarsi. Trichobothrial bases with longitudinal ridges. Tarsal organ low, lingulate, with weak concentric ridges. Female palp with tarsal claw possessing few teeth located prolaterally. Paired claws on tarsi I and II biserially dentate with numerous teeth, on tarsi III and IV usually with fewer teeth, in females inner teeth row may be reduced completely. Unpaired tarsal claw small, curved.
male PalP. Tibia moderately long to very long, slightly to distinctly swollen or cylindrical, with no, few, or numerous spines. Cymbium asymmetrical with retrolateral lobe extended, with or without spines. Male palpal bulb pyriform or asymmetrically shaped. Embolus tapering and curved, in A. incana and related species with small subapical process.
sPinnerets. Two pairs of spinnerets. PMS medium-sized to small with functional spigots confined to apex only. Apical segment of PLS mostly triangular in females but usually longer (may be shortly digitiform) in conspecific males. Spigots: MAC not evident, only AC type present.

Species grouping
To assist with identifications, the species treated here are assigned to six species groups. These assignments are preliminary, because females in some species are unknown and they are not based on a phylogenetic grouping, though these groups may indeed reflect phylogenetic relationships.

Diagnostic characters
Males differ from other male congeners in possessing a distinctly swollen palpal tibia, almost or completely lacking spines , combined with a moderately long and broadly tapering embolus . Females have two or three branches in the spermathecae and relatively short stalks and weakly sclerotised receptacles (Figs 224-232).

Diagnosis
Anemesia birulai is closest in habitus to A. pococki sp. nov. and A. tubifex; however, it differs by darker colouration from the former, and by AMEs spaced closer from the latter. Males can be also distinguished from A. pococki sp. nov. by the spinose cymbium, and from A. tubifex by a shorter and more swollen palpal tibia and by a shorter embolus (Figs 173,(188)(189)(190)(193)(194)(195)(196) colour. Carapace and legs reddish brown, carapace with slightly darkened clypeus and eye tubercle brownish black; chelicerae dark reddish brown; sternum, labium and maxillae light reddish brown; cymbium brown; sternal sigillae brown; abdomen dorsally light brow with darker standard pattern ZONSTEIN S., Revision of Anemesia (Araneae, Cyrtaucheniidae) consisting of broad median spot and six pairs short chevron-like lateral spots inclined backward; dorsal abdominal surface and spinnerets yellowish brown.
PalP. Tibia, cymbium and palpal organ as shown in Figs 173,[188][189][190]. Palpal tibia swollen (Fig. 173). Embolus tapering slightly curved  colour. Differs in some details from that of male holotype: carapace and legs ferruginous-brown; anterior half of carapace darker, anterior edge and lateral margins of carapace as well as femora, patellae and tibiae of legs I and II are even darker, brown-coloured; eye tubercle with brownish-black spots around AMEs and lateral eyes; sternum, labium, maxillae, coxae ventrally, metatarsi and tarsi III and IV yellowish-brown; abdomen light greyish-yellow with mid-brown pattern consisting of moderately narrow median stripe and six paired chevrons; spinnerets even paler greyish-yellow.
sPermatHecae. Two or three branched, as shown in Fig. 224.

Variation
The length of the carapace varies from 4.65 to 5.30 in males and in females from 6.00 to 7.85. The coloration weakly varies across specimens from Badhyz Plateau; however, spiders collected in the surroundings of Ashgabat at a lower altitude (200-400 m vs 600-1100 m) look somewhat paler. The number of the maxillary cuspules ranges from 15 to 23 in males and from 13 to 21 in females. The variability in structure of the male palpal organ and the spermathecae is shown in Figs 188-190 and 224-227, respectively.

Habitat
The species occurs in foothill and low mountain steppes, often with shrubs or open park forest composed mainly of Pistacia vera L. The vicinity of the Kepele well, with hillsides inhabited by these spiders, is shown in Figs 307-308.

Distribution
South Turkmenistan. See Fig. 369. Vlasov & Sytshevskaya (1937) listed "Sterrhochrotus ferghanensis Croneb." among spider species collected from the burrows in surroundings of Ashgabat. The re-examination of two females deposited in ZMMU, herewith identified and labelled, revealed they are Anemesia. Since both these females are immature, they are provisionally assigned here to A. birulai, the only congener known in the neighbourhood; their studied characters do not contradict such assignment. Charitonov (1969) indicated this species as also distributed in Uzbekistan. A re-examination of the material used in his study (PSU), revealed that he dealt with representatives of an undescribed species (see description of A. sogdiana sp. nov.). The corresponding reference is thus excluded from the synonymy list of A. birulai. Marusik, Zamani & Mirshamsi, 2014 Figs 2, 49, 91, 118, 174, 191-192, 268, 369 Anemesia koponeni Marusik et al. 2014: 2, figs 1-8 (♂).

Diagnosis
By structure of the palp with a moderately swollen tibia and a widely tapering embolus, the relatively large male holotype of Anemesia koponeni resembles the large males of A. tubifex. However, it differs from them by the almost uniformly dark brown coloured body and legs (vs the lighter and more contrastingly coloured body and legs in A. tubifex; Fig. 2, cf. Fig. 4), as well as by a narrower eye group and by the closer spaced AMEs (Fig. 49, cf. Fig. 51).

Habitat
No data.

Distribution
The species is known from the type locality only (see Fig. 369).

Etymology
The specific name is a patronym in honor of Dr Reginald I. Pocock, the noted British arachnologist , who provided and published many studies on the Mygalomorphae; he described, in particular, both the genus Anemesia in 1895 and its type species a few years earlier (1889). colour. Carapace light brownish orange with anterior edge darker and thoracic part lighter, chelicerae light orange; palps and legs dorsally less intensive orange; sternum, labium, maxillae and legs ventrally pale brownish yellow; eye tubercle blackened; abdomen dorsally pale yellowish grey with brown pattern ZONSTEIN S., Revision of Anemesia (Araneae, Cyrtaucheniidae) consisting of moderately wide median lanceolate spot and six paired transverse and slightly inclined short stripes, ventral part of abdomen pale yellow, spinnerets pale brownish yellow.
legs. Tibia and metatarsus I as shown in Fig. 119. Scopula: distal on metatarsi I-II, entire on tarsi I and II, divided on tarsi III, vestigial on tarsi IV. Trichobothria: 2 rows of 8 each on tibiae, 12-14 on metatarsi, 12-13 on tarsi. PTC I-III with 9-10 teeth on both outer and inner margin, PTC IV with 8 and 5-7 teeth respectively.

Variation
The length of the carapace varies from 5.26 to 5.70 in males and from 5.15 to 6.37 in females. The colouration varies very narrowly throughout the specimens. In females the number of retromarginal cheliceral teeth ranges from 5 to 8 (as in Fig. 78). Variants in the shape of the spermathecae are shown in Figs 228-229.
PalP. Tibia, cymbium and palpal organ as shown in Figs 176,[195][196]. Tibia moderately long and weakly swollen (Fig. 176). Palpal organ with embolus tapering and noticeably curved . colour in alcoHol. Similar to that of male, but carapace and legs are paler, as well as dorsal abdominal pattern is paler and less contrasted.

Variation
The length of the carapace varies from 6.50 to 6.55 in males and from 7.52 to 8.63 in females; the number of the maxillary cuspules ranges between 11 and 17; both the colouration and the leg spination range very narrowly.

Habitat
The species was found to inhabit at least two types of habitats, in piedmont and montane areas of the Badhyz Plateau, respectively. The only visited piedmont habitat, located in the surroundings of ZONSTEIN S., Revision of Anemesia (Araneae, Cyrtaucheniidae) Kalaimor, is typical for the region: a true sand desert occupying the upper terrace on the left bank of Kushka River, 1-1.5 km from the floodplain. A sandy substrate features sparse herb vegetation dominated by the ephemerous desert sedge, Carex physodes M.Bueb. The visited montane habitats, confined to a relatively high south-western edging of the uplands, represent mountainous wormwood and cereal steppe on a rocky substrate, with sparse trees and shrubs (see Fig. 307).

Distribution
Northern Afghanistan and the far southern Turkmenistan (Fig. 369).

Notes
Although the holotype is in poor condition (a result of careless sampling rather than bad preservation), the identification of other specimens belonging to Anemesia tubifex is not problematic. Among all members of the considered species group, only Anemesia tubifex has such a broadly extended eye group and such broadly spaced AMEs. Within females of Anemesia, the somewhat comparable eye group can be observed only in the geographically distant A. andreevae sp. nov., A. castanea sp. nov. and A. karatauvi (see Figs 66,[70][71]. However, females of A. andreevae sp. nov. have legs III -IV barely stouter than legs I-II (vs evenly slender legs I-IV in A. tubifex), whereas females of A. castanea sp. nov. and A. karatauvi possess more numerous maxillary cuspules (40-70 vs 14 -18 cuspules in A. tubifex).
Examination of the spider material used by Denis (1958) for his work on the spider fauna of Afghanistan, revealed that he listed a new record of A. tubifex erroneously, since the latter was based on misidentified material (see page 48). The corresponding reference is therefore excluded from the synonymy list of this species.

Diagnostic characters
The distinguishing characters of this group are: 1) the specific form of the embolus which is relatively short and bent medially, possessing a very thin and narrow distal part; 2) the unique shape of 3-branched spermathecae, provided with strictly sclerotised receptacles.

Species included
Only one species, A. andreevae sp. nov.

Etymology
The specific name is a matronym in honour of the late Central Asian and Polish arachnologist Dr Ekaterina Andreeva (Dr Katarzyna Andrejeva-Prószyńska;1941-2008, who made a very significant contribution to the study of the spider fauna of Tajikistan; in the course of her numerous taxonomic European Journal of Taxonomy 485: 1-100 (2018) studies she described many regional spider taxa, including Brachythele karatauvi  assigned later to Anemesia.
Paratypes (17 ♀♀) UZBEKISTAN: 2 ♀♀, same collection data as for the holotype (SMNH). colour. Carapace light yellowish brown with anterior edge darker and thoracic part lighter; chelicerae, most part of palps and legs light yellowish brown; sternum, labium, maxillae and leg tarsi paler; eye tubercle blackened; abdomen dorsally pale yellowish brown with brown pattern consisting of moderately wide median lanceolate spot and few paired transverse and slightly inclined short stripes, ventral part of abdomen pale yellowish grey, spinnerets pale brownish yellow.
PalP. Tibia, cymbium and palpal organ as in Figs 177,[197][198]. Palpal tibia long and slender (Fig. 177). Embolus relatively short, curved, and bent, with very thin distal part  colour. Darker than that in male: carapace and legs dorsally ferruginous yellow; carapace with caput slightly darker and anterior edge including clypeus mid-brown; eye tubercle with blackish brown spots around AMEs and lateral eyes widely spaced; chelicerae reddish brown; sternum, legs ventrally, genital area and spinnerets light brownish yellow; labium, maxillae and palps with similar but slightly darker colouration; abdomen light ferruginous grey, dorsally with diffuse brownish pattern consisting of rather narrow median stripe and five pairs of lateral chevrons.

Variation
The length of the carapace varies from 4.30 to 5.53 in females and the number of maxillary cuspules ranges from 8 to 19. In females from Tajikistan (Varzob Canyon), the number of promarginal and retromarginal teeth may reach 8 and 6, respectively (see Fig. 80). The structure of the spermathecae appears to be almost the same in females collected from the different localities .

Habitat
The species occurs in the foothills covered with deciduous shrub with Prunus spp. and Pistacia vera L. (Fig. 310), and in the middle mountain belt. In the latter case the spiders were found inhabiting open forest biotopes dominated by Juniperus seravschanica Kom. and Acer spp. (Fig. 311).

Diagnostic characters
Males in the A. sogdiana group resemble males of the A. tubifex species group in possessing a visibly swollen palpal tibia. However, they differ from the members of the latter group in having a noticeably longer and narrowly tapering embolus. Females can be distinguished in having the specific shape of undivided spermathecae with relatively long stalks narrowed apically.

Species included
Two species: A. oxiana sp. nov. and A. sogdiana sp. nov.

Etymology
Oxiana (ancient Greek 'Ώξιανή') is the name of an historical area in Central Asia that existed in the antiquity and is applied to the territory located alongside the Oxus (i.e., the recent Amu-Darya River and its main component Panj River, forming the Afghanistan-Tajikistan border) which corresponds to the known range of this species. This specific epithet is a noun in apposition to the genus name, the gender is feminine.
European Journal of Taxonomy 485: 1-100 (2018) PalP. Tibia, cymbium and palpal organ as shown in . Palpal tibia moderately long, slightly swollen and spinose; cymbium with few apical spines (Fig. 178). Palpal organ with broad asymmetric bulb and with long, tapering and slightly curved embolus  colour. Differs from that of male in some details: carapace and legs light foxy brown; sternum, labium and maxillae slightly paler; caput darkened, but anterior part with pair of symmetric lighter, yellowish spots located laterally from eye tubercle; blackish brown spots surrounding AMEs and lateral eyes semifused; chelicerae dark carmine red; abdomen light greyish yellow, with dark brown pattern consisting of rather narrow transverse longitudinal spot crossed by wide transverse fascia and six pairs of lateral chevrons posteriorly; spinnerets light greyish yellow.

Variation
The length of the carapace varies from 7.10 to 8.13 in males and from 7.78 to 10.04 in females. Colouration, shape of the embolus and configuration of the spermathecae vary through specimens very narrowly (see [199][200][201][202][241][242][243][244]. In some females, the cheliceral furrow may be armed with 2-3 retromarginal teeth (see Fig. 81). The number of maxillary cuspules may range from 40 to 55 in males and from 60 to 85 in females. Tarsus III with 1-2 spines (most often occupying prolateral position); tarsus IV mostly aspinose (in some specimens with one prolateral spine).

Habitats
The most widespread and common congener in southern Tajikistan

Diagnosis
In habitus, Anemesia sogdiana sp. nov. is largely similar to A. pococki sp. nov. and especially to A. andreevae sp. nov., but differs from them by the long, thin and gradually curved embolus in males (197)(198) as well as by the entire and narrowed subapically spermathecae in females (233)(234).
sPinnerets. See Fig. 277 colour. Carapace and legs dorsally light yellowish foxy brown; cephalic part darkened, medium foxy brown; eye tubercle with blackish brown spots around AMEs and lateral eyes; chelicerae reddish brown; sternum, labium, maxillae and legs ventrally pale brownish yellow; abdomen light greyish brown with dark brown dorsal pattern consisting of wide median longitudinal spot and six pairs of lateral chevrons; spinnerets very pale greyish brown.

Variation
The length of the carapace varies from 4.87 to 5.21 in males and from 4.47 to 7.33 in females. Colouration varies through specimens very narrowly (see Fig. 30). Throughout the specimens, both PLE and PME may be reduced or even lost (Fig. 69). In some females, the cheliceral furrow may be armed with 2-3 retromarginal teeth (see Fig. 82). In conspecific males, the shape of the palpal organ is almost identical (Figs 203-205). Spermathecae show some variations and can feature with straight or bent stalks (Figs 235-240).

Habitat
The known records mostly correspond to the midland mountain belt although the species can also occur in both the piedmont semi-desert (Zafarabad) and the highlands (

Distribution
Central and southern Uzbekistan, and western Tajikistan (Fig. 369). Charitonov (1969) listed five males of this species collected by K. Arnoldi and D. Fedotov in 1942 in the surroundings of Ishkent as Brachythele birulai Spassky. A detailed examination of the spider collection at the Perm University (1987,1988) revealed the presence of only two appropriate male specimens sampled and labelled according to the mentioned record data.

Diagnostic characters
Males of the A. castanea species group resemble those of the A. pallida species group in possessing lateral spines on the cymbium. However, they differ from the members of the latter group in possessing a considerably longer palpal tibia and embolus (206)(207)(208)(209)(210)(211)(212)(213)(214). Females are distinguished by the specific shape of the normally entire spermathecae provided with long cylindrical stalks (see .

Species included
A. castanea sp. nov. and A. karatauvi .

Etymology
The specific name is a derivative (using the feminine gender) of the Latin adjective 'castaneus' meaning 'chestnut-brown'; the name is derived from the colouration of the specimens. colour. Carapace almost uniformly light chestnut brown; chelicerae, leg I entirely and most part of palps and legs II-IV light-medium to dark brown; sternum, labium, maxillae and leg tarsi paler, brownish orange; eye tubercle black; abdomen dorsally yellowish grey with dark brown pattern consisting of dentate median stripe anteriorly and few paired transverse and slightly inclined chevrons posteriorly; ventral part of abdomen and spinnerets pale yellowish grey.

Variation
The length of the carapace varies from 5.80 to 6.25 in males and from 7.73 to 8.39 in females. The colouration varies through specimens very narrowly. The number of maxillary cuspules ranges from 32 to 40 in males and from 65 to 70 in females. The variation in the shape of the palpal organ and the length of the embolus seems to be almost imponderable (see .

Habitat
The species occurs in the piedmont sandy desert (Tigrovaya Balka Nature Reserve) and in the woodless steppe foothills near Ganjina. Curiously, despite intensive collecting, A. castanea sp. nov. has never been found in the less harsh montane zone of the Aruktau Mts, occupied by A. incana and some other congeners.

Distribution
The species is known from far southwestern Tajikistan only (Fig. 371).

Notes
Andreeva (1968) included both conspecific females in the type series of Brachythele karatauvi. To avoid a situation of the same specimen being listed as a paratype in two different species, these females are not included in the types of Anemesia castanea sp. nov., despite definitely belonging to the latter congener.

Variation
The length of carapace varies from 5.78 to 5.90 in males and from 5.82 to 9.23 in females. The colouration varies through specimens very narrowly. The number of maxillary cuspules ranges from 28 to 32 in males and from 40 to 55 in females. No significant variation in structure of the palpal organ is evident. The spermathecae are generally uniform (Figs 246,(248)(249); however, occasionally the stalks are divided subapically (see Fig. 247).

Habitats
The species occurs in woodless steppe foothills at 700-1000 m, as well as in the scarcely forested midland area above, with shrubs and low trees represented by Acer, Prunus and Pistacia (see Figs 313,317).

Distribution
Contrary to the distribution provided with the original description, the range of this species is confined to the Vahsh Karatau Mts in southwestern Tajikistan only, as shown in Fig. 371 (see below).

European Journal of Taxonomy 485: 1-100 (2018)
Notes A captured live male is shown in Fig. 31 (a positive slide made in 1989). The cheliceral rastellum, trichobothrial bases and tarsal organ of leg I, and spigots are shown in Figs 145-146, 169-170, and 298-301, respectively. Andreeva ( , 1976 listed as paratypes of B. karatauvi 17 ♀♀ collected together with the holotype and 8 ♀♀ collected in 1966-1967 in other regions of southern Tajikistan. As noted above, the type series comprises only 7 ♀♀ from Karatau Mts. All other females originally included in the type series are reidentified and considered here as belonging to other species of Anemesia.

Diagnostic characters
The group comprises small, pale-coloured species. Males in this group resemble those of the castanea species group in possessing a laterally spinose cymbium, but differ from them in the considerably shorter palpal tibia and embolus (211)(212)(213)(214)(206)(207)(208)(209)(210). Females of Anemesia pallida sp. nov. can be distinguished from other congeners in the reduced dorsal abdominal pattern (Fig. 23) and in the specific shape of the deeply divided spermathecae (Fig. 250).

Species included
Anemesia pallida sp. nov. and A. parvula sp. nov.

Etymology
The specific name is derived from the Latin adjective 'pallidus' meaning 'pale', and reflects the general light colouration of the specimens; the gender is feminine. Paratypes (11 ♂♂, 2 ♀♀) TAJIKISTAN: 4 ♂♂, 1 ♀, same collection data as for preceding but 13-19 Apr. 1986 (SMNH); 1 ♂, same collection data as for preceding but 9 Apr. 1988 (SMNH); 3 ♂♂, same collection data as colour. Carapace pale yellowish orange with brownish clypeus and blackish-brown spots surrounding AMEs and lateral eyes; chelicerae, palps and legs dorsally yellowish orange with femora and patellae slightly darker; sternum, labium, maxillae, palps and legs ventrally pale brownish yellow; abdomen dorsally greyish-yellow with simple brown pattern consisting of relatively narrow median lanceolate spot only, ventral part of abdomen pale yellowish grey, spinnerets pale brownish yellow.

Variation
The length of the carapace varies from 4.10 to 4.55 in paratype males, the dorsal abdominal pattern may be developed somewhat better and represent the more contrasted variant than that shown in Fig. 10. The number of tarsal spines varies as follows: leg II: 0-3; leg III: 6-12; leg IV: 9-20; the number of the maxillary cuspules ranges as 21-27 in males and 28 -32 in females.

Etymology
The specific name is derived from the Latin adjective 'parvulus' meaning 'small', and reflects the minute size of the specimens; the gender is feminine.

Variation
The carapace length is 4.44 in the only paratype male; the colouration, including the dorsal abdominal pattern, looks completely the same as in the holotype and distinctive from that in males of A. pallida sp. nov.

Habitat
See the corresponding data for A. infuscata sp. nov.

Distribution
Known only from the type locality in the central part of Tajikistan (Fig. 372).

Diagnostic characters
This group comprises intensely dark-coloured medium-sized and large species. Members of this species group can be easily distinguished from other congeners due to the unique membranous process near the embolus apex in males (see Figs 148,150), and to a specific shape of the entire spermathecae with unusually wide stalks in females (Figs 251-265). Unlike most other members of Anemesia, all three species of the considered group have relatively large PMS (288)(289)(290)274,(277)(278)(279)(280)(281)(282)(283)(284).
legs. Scopula distal on metatarsi I and II, entire on palpal tarsus, very narrowly divided on tarsus I, narrowly divided on tarsus II, elsewhere absent. Trichobothria: 8-10 in each row on tibiae, 16-20 on metatarsi, 15-18 on tarsi. Palpal claw with 4 promarginal teeth. PTC I-II with 6-7 teeth on each margin. PTC III with 6 teeth on outer margin, 3 teeth on inner margin; PTC IV with 3 teeth and one tooth, respectively.

Variation
The length of the carapace varies from 6.30 to 7.92 in males and from 5.96 to 8.43 in females. The colour of the carapace and legs varies from very dark reddish brown to deep blackish-brown. Number of maxillary cuspules varies from 2 to 11.

Habitat
Foothills and low mountain ridges at an altitude of 600-1700 m, covered with ephemerous grasslands and subshrubs (mostly species of Artemisia), often with tall shrubs or low open forest (Pistacia vera L., different species of Prunus, Crataegus and Acer). The typical biotopes preferred by these spiders are shown in Figs 318-321.

Notes
Some illustrations of Anemesia incana used in this study are partially based on the additional (non-type) material. Live spiders are presented in Figs 32-33, 35, 43-44. The dentition of the cheliceral furrow in female and male paratypes is shown in Figs 84 and 85, respectively. The separated female palps and legs I-IV are shown in Figs 138-142. The SEM micrographs include those of the palpal organ (147, 149), the tip of the embolus (Figs 148, 150), the tarsal claws (Figs 156-157), the trichobothrial base and the tarsal organ of leg I (Figs 171 and 172, respectively), and spigots of PLE (Fig. 302) and PME (Figs 303 -305).

Etymology
The specific epithet is derived from the Latin adjective 'infumatusi', meaning 'smoked', referring to the mostly darkened (blackened) colouration of these spiders. The gender is feminine. colour. Carapace intensely dark mahogany, with darker brown clypeus, margins, thoracic grooves, and numerous small spots forming reticulate pattern on mahogany background; eye tubercle brownish black;

Holotype
European Journal of Taxonomy 485: 1-100 (2018) chelicerae, palps, leg I entirely, leg II from femur to tibia, femora III and IV dark chestnut brown; other segments of legs III and IV, and metatarsus and tarsus II light to medium brownish orange; maxillae orange, sternum and labium medium brownish orange; abdomen greyish, dorsally with dark coffee brown pattern consisting of median longitudinal stripe, very broad in anterior half, but narrow and fused with several paired chevrons in posterior half; spinnerets pale brownish yellow.

Variation
The length of the carapace varies from 6.65 to 6.78 in males and in females from 5.58 to 8.09. There are small variations in the eye size and arrangement as shown in Figs 74-75. The shape of the palpal organ in both collected male congeners looks practically the same. The configuration of the spermathecae does not show any noticeable difference (Figs 255-257).

Habitat
As in A. incana (see above); however, A. infumata sp. nov. has been hitherto encountered in low and middle montane areas between 800 and 1700 m, with slopes covered by rarely standing trees (Acer, Crataegus), shrubs and steppe vegetation (Figs 322-323).

Distribution
Known currently from a few localities in South Tajikistan only, see Fig. 273.

Holotype
colour. Carapace, palps and legs predominantly intensive dark reddish brown; anterior edge and lateral margins of carapace, chelicerae and femora even darker; whole eye tubercle brownish black; sternum, labium, maxillae and leg coxae light coffee brownish coloured, metatarsi and tarsi III-IV brownish orange; abdomen yellowish brown with dark chocolate brown pattern consisting of very broad median longitudinal stripe crossed medially and posteriorly with six wide procurved fasciae, and numerous small and partially fused spots located between them; ventral abdominal surface with darker horse-shoe spot covering genital area; book-lungs paler, light yellowish brown; spinnerets even paler, light yellow.
PalP. Tibia, cymbium and palpal organ as in Figs 187,[220][221]223. Tibia cylindrical with numerous thick ventral bristles (Fig. 187). Embolus tapering, slightly curved, embolic tip with small membranous keel 223 TBL 22.30,CL 8.55,CW 7.32,LL 1.12,LW 1.82,SL 4.70,SW 4.11. colour. Similar to that of male, but carapace and leg more intensive reddish coloured; dark brown dorsal abdominal pattern is even better developed; ventral brownish yellow surface of abdomen with numerous dense and tiny brown spots located irregularly between last pair of book lings to spinnerets.

Variation
The length of the carapace varies from 6.48 to 6.97 in males and from 6.89 to 9.47 in females. The colour varies narrowly from somewhat lighter and more reddish to an even darker brown than that of the holotype. The thoracic fovea are sometimes slightly recurved. Variations in shape, arrangement and structure are shown for the eyes (76)(77), the cheliceral furrow (Figs 87-89), the sternum (Figs 115-116), the male tibia and the metatarsus I (Figs 131-132), the male palpal tibia and the cymbium (Figs 186-187), the palpal organ (Figs 220-223), and the spermathecae (Figs 258-265). The number of maxillary cuspules ranges from 10 to 14.

Notes
Some illustrations of Anemesia infuscata sp. nov. are based on the additional paratype specimens; these include the images showing variations (see above) and photos of live spiders presented in Figs 38-41 and 45-47.

Data on natural history
Burrows All known species of Anemesia, living mostly in large colonies, build open burrows lacking the wafer-lid or trap-door (see . The length of the burrows generally varies from 25 to 40 cm  and own data), although in some populations of A. tubifex inhabiting the sandy desert biotopes, it can reach 60 cm in length. In adult congeners the diameter of the burrow entrance, always with a raised rim, ranges from 0.5 to 1.5 cm. In A. incana this rim is more developed and resembles a tower-like or mushroom-like surface soil turret 2-3 cm in height and 1.5-4 cm in diameter (Figs 340-344). The entrance leads to a narrow burrow neck which then becomes wider; this dilated burrow section, lined by silk, ends with a wide living chamber . A variant of the general form is observed in A. oxiana sp. nov., in which the burrow is provided with a lateral chamber, either ending with an additional entrance located close to the main one (Fig. 332), or blind and closely reaching the soil surface.

Phenology
Data on the phenology of the congeners are sparse and based completely on fragmentary field observations. The collecting data indicate that the wandering adult males of Anemesia occur in March-April in lowland localities at 200-1300 m and from mid-April to at least the end of May in midland localities at 1300-2300 m. Females with egg sacs containing 30-40 eggs were found only in October (A. infuscata sp. nov.). They have never been observed during the spring months, despite intensive collecting.

Feeding
Several burrows built by adult Anemesia females (11 burrows of A. karatauvi and 45 burrows of A. incana) were examined to reveal the character of the prey remains accumulated at the burrow bottom. Almost all these burrows, with minor exceptions, contained fragments (the head capsules, parts of the legs, the separate thoracal and abdominal sclerites) of the carpophagous foraging ants belonging to two species of Messor Forel, 1890: M. inermis Kuznetsov-Ugamsky, 1929 andM. structor (Latreille, 1798).

Notes
The re-examination of these specimens, identified by Denis (1958) as Anemesia tubifex, shows that they actually represent an undescribed alpine species of the nemesiid Raveniola Zonstein, 1987, closely related to R. concolor from the highlands of northern India. The characters missed by Denis (a short and wide labium, a broadly rounded sternum, a fairly rounded and submarginal posterior pair of the sternal sigilla, the characteristic divided spermathecae, as well as many less significant features) contradict his identification and confirm the given re-assignment. The species is planned to be described in a revision of Central Asian Raveniola spp. (Zonstein, in prep.).

Introductory notes
In this review Anemesia has been compared with the putatively related genera of the Cyrtaucheniidae, the related family Euctenizidae, and the similarly looking members of two subfamilies of the Nemesiidae: Nemesiinae s. str. (without Raveniola, tentatively included in this subfamily for want of a better assignment), and Bemmerinae. Several characters of other nemesiids have also been included. The Bemmerinae are considered here without Pionothele Purcell, 1902 which was tentatively included in this subfamily by Raven (1985). Prior to 1985, all these taxa were considered as members of the same family, Ctenizidae s. lat. Raven (1985) elevated the earlier tribes of Simon, Cyrtaucheniae and Nemesiae, to family rank, Bemmereae to the subfamily of the Nemesiidae, and established the Euctenizinae as a new cyrtaucheniid subfamily. Bond et al. (2012b) raised the latter taxon to family rank. Currently, the monophyletic status of both Cyrtaucheniidae and Nemesiidae is debated (Goloboff 1993;Bond & Opel 2002;Bond et al. 2012b;Garrison et al. 2016;Wheeler et al. 2017). These families are expected to be radically revised in the near future. Therefore, all the obtained data and results listed below should be considered as approximate and preliminary.
clyPeus. The anteriorly protruded clypeus is shared by almost all Anemesia spp.; this character seems less developed in males than in the conspecific females (see Figs 48-77). It does not represent, however, the unique feature of this genus since a somewhat similar though less pronounced clypeus is observed in some Eucteniza Ausserer, 1875 (Euctenizidae) (Bond & fig. 5a), most genera of Cyrtaucheniidae (save Anemesia and some Ancylotrypa Simon, 1889) and the majority of nemesiids lack a pronounced clypeus (see Raven 1985). Curiously, Raveniola ZONSTEIN S., Revision of Anemesia (Araneae, Cyrtaucheniidae) cyrtaucheniids Acontius, Ancylotrypa, Cyrtauchenius Thorell, 1869 (Figs 397, 400, 402, respectively) and Rhytidicolus (Goloboff 1993: fig. 7), as well as for the euctenizid Aptostichus (Bond 2012: figs 42, 63). It appears equally dissimilar to the domed TO of Nemesia (Fig. 407) and to the craterous TO of the nemesiid Entypesa Simon, 1902 (Fig. 403). Unfortunately, the scarcity of the corresponding data (structure of the tarsal organ and the trichobothrial bases remains unknown for most cyrtaucheniids and euctenizids) makes these characters limited in value when being used in relation to Anemesia spp.
sPermatHecae. Raven (1985) listed the multilobate spermathecae (which have been found in several species of Anemesia) among the diagnostic characters of the Cyrtaucheniidae. This feature may have a limited taxonomic significance because the multi-branched spermathecae have been recently described also for some nemesiids (see Goloboff 1995: figs 66E, 68, 71, 74A;Ferretti & Pompozzi 2016: fig. 10). Within the latter family, however, both the Bemmerinae and Nemesiinae possess entire or sometimes divided, but never multilobate, spermathecae (Raven 1985(Raven : fig. 54, 1987  . For some genera of Cyrtaucheniidae, however, the structure of the vulva still remains either unknown or known and shown only for single species (Raven 1985: figs 201, 208;Rudloff 1996: fig. 5). Regarding Anemesia, the pronounced differences in shape of the spermathecae may be considered a criterion for clustering congeners into the well-defined species groups, as it has been shown for the nemesiid Raveniola (see Zonstein et al. 2018).

Relationships
As noted earlier , females of Anemesia spp. possess several characters listed by Raven (1985), as indicative for the family Cyrtaucheniidae: (1) the carapace glabrous or only slightly hirsute; (2) an incomplete biserial dentition of PTC III and IV; (3) the characteristic comb of spines on the prodistal edge of femur IV; (4) the multilobate spermathecae (at least in some species); (5) the leg tarsi with few to numerous spines, and tarsi I-II with fewer (if any) spines than tarsi III-IV. Some exceptions include a hirsute carapace in some species of Cyrtauchenius, the aspinose leg tarsi I-IV, and the biserially and multidentate PTC III-IV in some Aporoptichinae (Op. cit.). Nevertheless, the placing of Anemesia in the Cyrtaucheniidae is problematic.
Superficially, Anemesia resembles the aporoptichine genera. It differs, however, significantly in the structure of the chelicerae, labium, sternum and sigilla which makes a close relationship between these groups improbable. Regarding other cyrtaucheniid genera, the situation looks less obvious. Amblyocarenum, Cyrtauchenius and Ancylotrypa share with Anemesia the anteriorly narrowed sternum, with the posterior pair of sigilla located relatively far apart the sternal edge, and lack of the metatarsal preening combs (entirely or in some species). Even so, in comparison with Anemesia, these genera possess a more elevated cephalic portion of the posteriorly narrowed carapace, a predominantly procurved thoracic fovea, the distinctly thickened female legs III-IV, and a considerably better developed cheliceral rastellum (Figs 374, 376;Raven 1985: figs 191-205;Dippenaar-Schoeman 2002: -47, 90-116, 143-146). Hence, the relationships between Anemesia and the three above-listed genera do not seem to be close.
Among the remaining cyrtaucheniid genera, the monotypic Asian Angka (Cyrtaucheniidae incertae sedis) shares with Anemesia some characters such as a retrolateral localisation of megaspines on the male tibia I, the almost uniformly slender legs I-IV lacking the metatarsal preening combs, the maxillae with a deep and rounded heel, a straight thoracic fovea, and a moderately arched cephalic portion of the carapace both in males and females (Raven & Schwendinger 1995: figs 6B, 6E, 6F, 6J, respectively). Concurrently, Angka differs from Anemesia in having a rounded (as in most nemesiids) sternum with small submarginal sigilla, the aspinose leg tarsi with equally and evenly bipectinate PTC (op. cit., fig. 3D), and tiny PMS. It can also be distinguished from the latter by the absence of an eye tubercle or any developed clypeus, and the apical spine cluster on the female femur IV. No characters demonstrating a common specialisation shared by these genera, except possibly the same type of megaspine localisation, are evident. In other words, despite several characters shared by Angka and Anemesia, most of them are here considered symplesiomorphic and thus not indicating the close relationship between these genera.
Owing to the presence of the second teeth row, Anemesia shows some resemblance to Homostola; additional similarity is based on the fact that in the latter genus legs I-II either equal in width to legs European Journal of Taxonomy 485: 1-100 (2018) III-IV, or only slightly thicker than the posterior legs, and the congeners appear habitually to resemble Anemesia   fig. 1B;   fig. 2M). Unfortunately, in this case no male characters could be compared, because all five species of Homostola still remain known exclusively from females (information concerning males of H. zebrina is based on misidentifications; see Raven 1985). The differences between Homostola and Anemesia relate mainly to the shape of the thoracic fovea and the apical segment of PLS (mostly procurved and domed in the former genus vs more or less straight and triangular to shortly digitiform in the latter one). Apart from Anemesia, members of Homostola possess the scarcely toothed PTC I-IV (not only PTC III-IV, as in the former genus), and the metatarsal preening combs on legs III-IV. Within the Cyrtaucheniidae, Homostola appears currently to be the least distant genus from Anemesia.
The remaining variants can be presented here as follows. The Euctenizidae are strictly distinguishable from Anemesia in a great array of parameters, including the family level features listed by Raven (1985), Bond & Opel (2002) and Bond et al. (2012b). Unlike Anemesia, the genera of Nemesiinae generally have a normally hirsute carapace, a rounded sternum with small submarginal sigilla, unarmed leg tarsi with equally multi-dentate PTC, a shortened male palpal tibia, and predominantly ventral or prolateral megaspines (only males representing the monotypic Mexentypesa Raven, 1987 possess a sessile retroventral megaspine; see Raven 1987: fig. 2). The members of Bemmerinae, although they also have the sigilla located distantly from the sternal edge, differ from Anemesia in having male PTC with a single S-shaped teeth row, and megaspines located ventrally and/or prolaterally. Thus, until both families, Cyrtaucheniidae and Nemesiidae, are revised and the character polarity is specified for these groups, Anemesia could be tentatively considered as a representative of the Cyrtaucheniidae incertae sedis, with the reservation that the genus seems to be more closely related to Homostola rather than to other cyrtaucheniid genera.

Biogeography
As a member of the Cyrtaucheniidae, the Central Asian Anemesia is isolated geographically from the prime range of this family which covers, according to the World Spider Catalog (2018), Central and South America, the entire African continent and the Western Mediterranean. Hence, the genus appears to be separated from this main family range by a vast area, including the fairly well-studied Eastern Mediterranean, where cyrtaucheniids have not been detected. In relation to Anemesia, the geographically closest family members are known from Algeria (Cyrtauchenius), mainland Italy, Sardinia and Sicily (Amblyocarenum), and Ethiopia (Ancylotrypa). Such isolation, however, does not represent anything extraordinary. In comparison with Anemesia, the South-East Asian cyrtaucheniid Angka appears to be isolated from other genera of this family to a much greater extent.
Biogeographically, the distribution of Anemesia is limited to the area of Turkmen-Khorassan and the Turkestan multi-zonal provinces belonging to the arid Sahara-Gobian Region of the Palearctic realm (see Kryzhanovski 1965Kryzhanovski , 2002. The known range of the genus is thus confined to the north-eastern part of the Turkmen-Horassan biogeographic province (Kopetdag and Piropamiz Mts, Badhyz and Karabyl Plateaus) and to the adjoining south-western part of the Turkestan biogeographic province (the western Hissar-Alaj Mts and Afghan-Tajik Depression). Within this range, two centres of species diversity can be currently observed; one in each of the noted provinces, respectively.
The first of these is restricted to the south-western Badhyz, where three of the four known members of tubifex species group are sympatric. The second centre of species diversity, even more defined, is located in the territory of the Afghan-Tajik Depression. Regarding the Tajik part of this region, many of the investigated localities are known to host three and some of them (e.g., Ganjina) four sympatric congeners. Three of six species groups and eight of fourteen species of Anemesia are represented only here (see . Outside the mentioned centres and at the range periphery, the genus is ZONSTEIN S., Revision of Anemesia (Araneae, Cyrtaucheniidae) represented by a few sub-allopatric species (in the localities situated closer to the diversity centres, these species, A. andreevae sp. nov., A. birulai and A. sogdiana sp. nov., are sympatric with some other congeners).
Unfortunately, the scarcity of data regarding the spider fauna of Iran and Afghanistan (Logunov & Zamanpoore 2005;Mirshamsi 2005;Logunov 2010) leaves some questions open. It is thus impossible at present to determine whether these two diversity centres are indeed isolated from one another, or whether we are able to see here only the two visible endings of a single invisible arc stretching across the northern face of the Hindu Kush, from Horassan in the west to the Pamirs in the east.