A revision of the spider genus Zaitunia (Araneae, Filistatidae)

the spider genus Zaitunia Lehtinen, 1967 (Araneae, Filistatidae) is revised. It was found to include 24 species distributed in the eastern Mediterranean, Middle east and Central asia: ♀ Z. afghana (Roewer, 1962) (Afghanistan), ♀ Z. alexandri Brignoli, 1982 (Iran), ♀ Z. akhanii Marusik & Zamani, 2015 (Iran), ♂♀ Z. annulipes (Kulczyński, 1908) (Cyprus), ♂♀ Z. beshkentica (andreeva & Tyshchenko, 1969) (Tajikistan, Uzbekistan), ♀ Z. brignoliana sp. nov. (Iran), ♂♀ Z. ferghanensis sp. nov. (Kyrgyzstan, Uzbekistan), ♀ Z. feti sp. nov. (Turkmenistan), ♀ Z. halepensis sp. nov. (Syria), ♀ Z. huberi sp. nov. (Afghanistan), ♀ Z. inderensis Ponomarev, 2005 (Kazakhstan), ♂♀ Z. kunti sp. nov. (Cyprus, Turkey), ♂♀ Z. logunovi sp. nov. (Kazakhstan, Kyrgyzstan), ♂♀ Z. maracandica (Charitonov, 1946) (Uzbekistan, Kazakhstan), ♂♀ Z. martynovae (Andreeva & Tyshchenko, 1969) (Tajikistan, Turkmenistan), ♀ Z. medica Brignoli, 1982 (Iran), ♂♀ Z. minoica sp. nov. (Greece), ♀ Z. minuta sp. nov. (Uzbekistan), ♀ Z. persica Brignoli, 1982 (Iran), ♂ Z. psammodroma sp. nov. (Turkmenistan), ♂♀ Z. schmitzi (Kulczyński, 1911), the type species (Egypt, Israel), ♂♀ Z. spinimana sp. nov. (Kazakhstan, Turkmenistan), ♂♀ Z. wunderlichi sp. nov. (Kyrgyzstan) and ♀ Z. zonsteini Fomichev & Marusik, 1969 (Kazakhstan). Twelve above-listed species are newly described, and males of Z. annulipes, Z. beshkentica, Z. maracandica and Z. martynovae are described for the first time. two new combinations are established: Z. annulipes (Kulczyński, 1908) comb. nov., ex Filistata, and Pholcoides monticola (Spassky, 1941) comb. nov., ex Zaitunia. new data on distribution of the considered taxa are provided.

Introduction the spider family Filistatidae is a group of cribellate spiders possessing a peculiar combination of both primitive and specialized characters. this is not a species-rich family: by 2015, Filistatidae included 18 genera and only 123 species (World Spider Catalog 2015). Nevertheless, the family has a worldwide distribution, occurring in tropical, subtropical, and warm-temperate areas (Gray 1995;). In the Palearctic region, it is mostly confined to the southern part: the Mediterranean, Middle East, Central Asia, and southern China and Japan (Helsdingen 2015;Platnick 2014;WSC 2015).
Until the revision of the cribellate spiders by Lehtinen (1967), all Palearctic filistatids were placed in the type genus Filistata Latreille, 1810 (for the complete list of synonymies, see Platnick 2014). Lehtinen (1967) described several new filistatid genera, including two from the West Palearctic: the widespread Pritha Lehtinen, 1967 and the monotypic Zaitunia Lehtinen, 1967. the genus Pritha was considered to include, among others, some species described by Simon (1868), Kulczyński in Chyzer & Kulczyński (1897), Strand (1914) and partially by Roewer (1962), whereas the monotypic Zaitunia included only Filistata schmitzi Kulczyński, 1911. Zaitunia was treated as a monotypic genus until Brignoli (1982) described three new species from Iran; however, the genus diagnosis was not modified. Zonstein (1990) transferred four species described by Spassky (1941), Charitonov (1946) and Andreeva & Tyshchenko (1969) from Filistata to Zaitunia. Ponomarev (2005), Fomichev & Marusik (2013) and Marusik & Zamani (2015) described three more species in Zaitunia, and Zonstein et al. (2013) transferred to this genus one species previously described by Roewer (1962) as a member of Filistata. thus, the genus currently is thought to have 12 species (World Spider Catalog 2015). However, only the type species is known from both sexes, whereas all other congeneric species are known exclusively from females or juveniles, and only from their original descriptions. additionally, the genus has never been revised, making its limits with Filistata unclear. Hence, clarifying this problem is one of the objectives of the present study. Zonstein (2009b) noted that a male of Z. schmitzi (Kulczyński, 1911) differed in many details from males of Central asian species, and suggested that the latter ones do not belong to Zaitunia but should be placed in a distinct genus. However, in the course of the present study, we recognised an intermediate state of several characters in males from Crete, Cyprus, and turkey. thus, all these groups are considered here as belonging to the same genus.
In this study, we examine all available material of this genus (340 specimens), revealing a total of 24 species of Zaitunia, 12 of which are described as new.

Photographs
Photographs were taken using a Zeiss Discovery V20 stereo microscope with a Canon PowerShot G9 camera and an olympus SZX16 stereo microscope with an olympus e-520 camera, and prepared using the CombineZP software. Scanning electron micrographs were made using the SeM JeoL JSM-5200 scanning microscope at the Zoological Museum, university of turku, Finland. illustrations of the endogyne, abdomen and spinnerets were made after maceration in a 20% potassium hydroxide aqueous solution and exposure for a few minutes in an alcohol/water solution of Chlorazol Black. it is worth noting that the receptacles in all Zaitunia species are very small and weakly sclerotized and therefore can easily be overlooked. Differential staining with Chlorazol Black is essential both for observing and photographing receptacles. in most cases, to make photographs of the endogyne and sometimes the palps more clear and to show more details, we placed a dark background in the case of transmission microscopy, or pieces of black plastic (or black paper), or minutia pins near specimens to be photographed.

ZONSTEIN S. & MARUSIK Y.M., Revision of Zaitunia (Araneae, Filistatidae)
Photographs of the somatic characters and some of the palps were taken in dishes with paraffin on the bottom to hold the specimens in the right position. endogynes were photographed on slides either under an Olympus SZX16 or an Olympus BH-2. Small pieces of cotton were used to fix the specimens in the correct position. Background maps were taken from the website http://www.maps-for-free.com.

Measurements
Measurements were made to an accuracy of 0.01 mm. Lengths of leg and palp segments were measured on the dorsal side, from the midpoint of the anterior margin to the midpoint of the posterior margin. all measurements are given in millimetres.

Spination
Since leg spination in most species appears to be almost invariable and thus has no significant taxonomic value, a generalised survey is provided in the description of the genus and particular variants are given only when they noticeably differ from the prevalent case. all femora with one basodorsal spine and 1-2 smaller spines in pro-and retrodistal position. Patellae unarmed. Tibiae with 2-6 ventral spines (can be absent on tibiae I-II in females). Metatarsi usually long and slender, with 6-9 ventral spines, but in males of Z. psammodroma sp. nov. metatarsi shortened, dilated and covered with numerous ventrodistal spines. Tibiae and metatarsi with few (1-3) inclined trichobothria, which are 1.2-2 times shorter than diameter of the segment. trichobotrial bases low and inclined, smooth outside and densely rifled inside ( Fig. 2H-I). Tarsi ventrally with few small spines, and with short and dense bristles. Male tarsi curved and pseudosegmented. tarsal organ with wide inclined opening ( Fig. 2F-G, J). Calamistrum short, present in females only, formed by one composite row of a few thick, curved and flattened setae on a raised keel (crest); median part of calamistrum without setae ( Fig. 2A-E). Paired tarsal claw narrow and slightly curved, with row of 8-15 long, dense teeth (Fig. 1H). Unpaired claw curved, with 5-8 dense teeth (Fig. 1I).
abdomen. abdomen elongate oval, overhangs posterior part of carapace. Spinnerets shifted anteriorly. Spinneret group set relatively far from posterior edge of abdomen. Cribellum small, bipartite trapezoidal, present in both sexes (Figs 3A,D,4D) but reduced in size and lacking functional spigots in males. ALS and PLS subequal in size (Figs 3A,C,4C); PMS much smaller, with two probably paracribellar spigots ( Fig. 4F, I). Tracheal spiracle wide and located in posterior ⅓ of abdomen between epigastral furrow and spinnerets ( Fig. 3A-C). Tracheae thin and short (about as long as spiracle width). Two pairs of tracheal stems: median branches longer and inclined sideward, lateral branches much shorter (Figs 3B,. as in the prosoma and legs, abdomen covered with ciliate hairs only, plumose hairs absent. male palp. Relatively short compared to other filistatines, although 1.3-1.8 longer than carapace. The shortest palp in comparison to the carapace is in the type species (palp/carapace length = 1.3). Femur unmodified, cylindrical; equal in length to or longer than tibia. Patella unmodified, short, about one third of femur length. tibia long and swollen, 1.5-2 times thicker than femur; ventromedially with patch of dense, suberect setae.
Cymbium. Conical at base, and the rest is cylindrical. only Z. logunovi sp. nov. has a conical dorsal outgrowth in the terminal one-third ( Fig. 1K-L). Tip of cymbium without dorsoapical excavation, its margin varies from strongly slanting (some specimens of Z. schmitzi, Fig. 7A-D) to almost straight (perpendicular to axis of cymbium). Dorsum of cymbium with brush of dense, long suberect hairs; some hairs as long as cymbium. one species, Z. logunovi sp. nov., with set of thick hairs along ventral and lateral edges of cymbium (Fig. 1J). Terminal part of dorsum hairless, semitransparent.

Species grouping
To assist with identifications, the species treated here are assigned to seven informal species groups. these assignments are preliminary, considering that males in many species are unknown, and not based on a monophyletic grouping, though some of the groups may actually reflect phylogenetic relationships.

Distribution
the genus is known from Crete to eastern kazakhstan, south to the Sinai Peninsula in egypt, Fars and kerman Provinces of iran and Southeastern afghanistan, and north to northwestern and northeastern Kazakhstan .

Ecology
Most species inhabit more or less arid habitats: deserts, semi-deserts, steppes, maquis or deciduous shrubland, where spiders may occur under rocks and in crevices of clay or rocky escarps where they build small tubular webs (see Fig. 50). Adult males were collected manually and by pitfall traps, generally during late spring or summer. only one species, Z. wunderlichi sp. nov., was found in a humid dense broad-leaved and mixed mountain forest dominated by walnut (Juglans regia). Concluding from the known data, Zaitunia avoid the true sandy deserts, with the only exception being Z. psammodroma sp. nov., known currently only from males. Contrary to all other congeners, males of Z. psammodroma sp. nov. possess short and distally dilated metatarsi iV with numerous ventrodistal spines, a clear adaptation to moving on loose sandy substrate. Zaitunia species are found from the seashore to altitudes over 2000 m (Z. beshkentica).

Etymology
The specific name refers to the type locality: Aleppo (Haleb). Colour. Carapace brownish-yellow with rich darker brownish pattern occupying clypeus, postocular area, radial grooves and, partially, the surface between those grooves; eye tubercle and narrow bands along margins dark brown; labium and sternum pale brownish-yellow; chelicerae, palps and legs light brownish-yellow with darker diffuse brownish spots and fasciae; abdomen uniformly yellowish-brown.  Fig. 6C-D). Median receptacles clublike, almost 3 times longer than globular lateral receptacles, and separated by one diameter of their heads.

Diagnosis
Males differ from those of all other congeners by a long and large cigar-shaped tegulum, a very thick spermophore occupying the whole bulb, a strongly slanting tip of the cymbium and by the extremely short and sideward-directed embolus (vs smaller coniform and longer forward-directed ones in other species). Females are easily distinguished by their intense dark coloration (darker than that in any other species) as well as their low and small globular receptacles, with outer lobe only slightly exceeding the median one in size ( Fig. 7E-K).
palp 43a). Femur equal in length to tibia; tibia thick, uniformly swollen. Cymbium moderately long, with or without small dorsodistal projection. Broad, blunt-tipped tegulum with thick spermophore forming 4 complete coils closely adjoining each other. Embolus short, lacking a neck, flattened and bent retrolaterally. Colour. as in male, but carapace, palps and legs slightly lighter; in contrast, fasciae on palps and legs somewhat darker than in male.

Variation
Carapace length in males varies from 1.03 to 1.30, in females from 1.37 to 2.00; ground colouration varies from medium yellowish-brown with a darker brown abdomen to an almost uniform dark brown.
Ecology the species inhabits many types of habitats, from deserts through steppes and maquis to low oak evergreen forest; in all these habitats, it occurs mainly under rocks (see Fig. 50A).

Zaitunia annulipes-group
Remarks this group unites moderately dark-coloured species. the clypeus, the eye tubercle and the carapace margins are noticeably darker than the general background colour. the abdomen is uniformly darkcoloured or with a weak, darker dorsal pattern. Legs i-iV are lighter than the body and provided with relatively narrow darker fasciae. the male palpal tibia is moderately short. the conical tegulum is much shorter than the cymbium. the embolus is a relatively long to very long apically twisted lamina. keel of embolus absent. the vulva has two pairs of receptacles. three species are included: Zaitunia annulipes (Kulczyński, 1908) from Cyprus, Zaitunia kunti sp. nov. from Cyprus and turkey, and Zaitunia minoica sp. nov. from Crete.

Diagnosis
Zaitunia annulipes differs from the congroupers by males possessing a shorter embolus gently curved in the subapical part and by females having the receptacles with the median units considerably larger than the outer ones (in congroupers embolus is considerably longer and more sharply curved subapically, and the median and outer receptacles are subequal in size -cf. Figs 10i, 11F-G, 12G-i, 13C-D).

Paralectotypes
CYPRUS: 2 juvs, with same collecting data. Colour. Carapace and legs mostly light brownish-yellow; cephalic part including eye tubercle and leg fasciae diffuse medium brown; clypeus and lateral margins of carapace more intensive brown to dark brown; labium, maxillae, sternum and leg coxae pale brownish-yellow; abdomen uniformly medium brown.
palp 43b). all segments except cymbium with dark pigmentation; femur longer than tibia. tibia almost 2 times wider than femur. Cymbium longer than bulb; bulb conical, twice as long as diameter of tegulum. tip of embolus twisted.

Note
Male of this species is described for the first time. Although this species was already transferred to Zaitunia by Brignoli (1982), this nomenclatorial act was overlooked by Brignoli (1983) himself and by subsequent catalogs (Platnick 1989(Platnick , 1993(Platnick , 2015World Spider Catalog 2015).

Ecology
According to the label data, the species was found in the maquis (Mediterranean shrub) litter.

Diagnosis
Zaitunia kunti sp. nov. differs from related species by having the embolus more sharply curved in the subapical part and receptacles subequal in size and clearly separated from each other (in Z. annulipes, the embolus is clearly shorter and more gently curved, and the median receptacles are considerably larger than the outer ones; whereas in Z. minoica sp. nov., a longer embolus is noticeably less sharply curved in the subapical part, and the receptacles, although subequal in size, almost touch each other (cf. Figs 9, 12G-I, 13C-D).

Etymology
The specific epithet is given after our good friend and colleague, Turkish arachnologist Kadir Boğaç kunt.

Variation
Carapace length in female paratype from Cyprus 1.68; this specimen has body and leg pattern slightly more contrast than those in female paratype from Turkey ( Fig. 11A-C), but the relative size and arrangement of darker marks do not differ sufficiently.
Ecology the type locality in turkey belongs to the coastal maquis zone; the female paratype from Cyprus was collected in the highlands (1700 m).

Diagnosis
Zaitunia minoica sp. nov. differs from other species of its group by having a longer embolus gradually curved in the subapical part, and by the receptacles subequal in size with widely spaced (more than  2 diameters) median pair (in Z. annulipes, the embolus is noticeably shorter and more gently curved, and the median receptacles are considerably larger than the outer ones; whereas in Z. kunti sp. nov., the slightly shorter embolus is more sharply curved subapically, and the receptacles are close together (about 1.5 of their diameter) -cf. Figs 9, 10F-G, I).

Etymology
The specific epithet is given after the Minoan civilisation, which flourished during the Bronze Age in Crete. Colour. Carapace dark ochre-yellow, with brownish clypeus, narrow marginal bands, postocular area and radial grooves; eye tubercle brownish-black; labium, sternum and maxillae yellow; chelicerae, palps and legs yellow with diffuse brownish spots and fasciae; abdomen uniformly dark brown.
palp 43d). Femur as long as tibia; tibia 1.5 times thicker than femur. Cymbium moderately long and slightly curved, without dorsal hump. tegulum globular. embolus long, gradually tapering near base and with parallel margins in mid-part, spermophore in embolus straight, tip of embolus screwshaped.
Colour. Whole body and legs light brownish-yellow, carapace with darker brown clypeus (however, with lighter yellow median stripe), long but narrow median spot dilated anteriorly; eye tubercle and narrow bands along carapace margins dark brown; chelicerae with narrow median brownish stripes; palpal and leg femora with narrow and incomplete light brown fasciae; abdomen without dorsal pattern.

Remarks
During the present study, we found that Roewer (1962) erroneously assigned two different species of Zaitunia, almost indistinguishable from each other by habitus, to the type series. Hence, we redescribe the holotype, but the paratype of Filistata afghana, previously redescribed by Zonstein et al. (2013), is considered to represent a new species, Z. huberi sp. nov., listed and described below.

Ecology
according to the label data, the holotype was collected in a cave; other information is not available.

Diagnosis
Females of Z. akhanii resemble those of Z. persica Brignoli, 1982 by having sinuous tube-like receptacles, but the receptacles of Z. akhanii have two loops, whereas those of Z. persica have four. Colour. Light yellowish, with distinct pattern on carapace and legs: clypeus entirely dark, wide dark median band terminating near fovea; femora i-iii with median-ventral spot. abdomen uniformly yellowish-grey, without darker pattern.

Variation
Total body length 4.80-7.20. Pale specimens may have light clypeus and may lack spots on femora.

Ecology
Specimens were found in large, dusty cribellate webs made around human dwellings.

Diagnosis
By structure of the endogyne, Z. alexandri resembles Z. medica, but differs from it and all other congeners by their entire, relatively short and robust sac-shaped receptacles (Fig. 16A-B). Colour. Carapace, chelicerae, labium and sternum pale yellowish-brown, carapace with darker brownish margins, clypeus and postocular area; eye tubercle dark brown; palps and legs with weak, diffuse darker brown spots; abdomen uniformly greyish-yellow.

Distribution
Known only from the type locality (Fig. 47).

Etymology
The specific epithet is given after the prominent Italian arachnologist Paolo Marcello Brignoli (1942Brignoli ( -1986 who described three species of Zaitunia from iran.
Colour. Carapace, chelicerae and legs mostly medium brownish-yellow; abdomen dorsally, diffuse leg fasciae, cephalic area and reticulate clypeal pattern medium violet-brown; eye tubercle and carapace margins dark violet-brown; sternum light yellow with brownish margins; abdomen ventrally and spinnerets light violet-brown.

Diagnosis
in structure of the vulva, females of Z. medica resemble those of Z. alexandri, but differ from them and all other congeners by their unpaired, very short and robust knob-shaped (transverse oval) receptacles (Fig. 17D, F-H).

Note
It is worth noting that Brignoli (1982) illustrated a more elongate receptacle with a stem that had distinct pores on the top, whereas the examined holotype has receptacles lacking a stem and covered with less distinct pores.

Diagnosis
Females of Z. persica resemble those of Z. akhanii by having sinuous, tube-like receptacles, but Z. persica has four loops (or bends), whereas Z. akhanii has only two loops.
Colour. Carapace and chelicerae brownish-yellow; carapace with brown to dark brown central area, clypeus and margins; eye tubercle brownish-black; maxillae, labium, sternum and leg coxae pale brownish-yellow; palps and legs medium brownish-yellow with incomplete brownish fasciae; abdomen uniformly pale yellowish-grey without darker pattern.

Distribution
Known only from the type locality (Fig. 47).

Zaitunia logunovi-group
Remarks this group includes only one light-coloured species. the dorsal body and legs with very weak darker pattern. Cymbium with brush of thick and dense setae on ventral and lateral edges (lacking in other groups). The cap-shaped tegulum is much shorter than the relatively long cymbium. The embolus is very short and twisted. the endogyne has two pairs of receptacles. the only member of this group, Z. logunovi sp. nov., is found in kazakhstan and kyrgyzstan.
Zaitunia logunovi sp. nov. urn:lsid:zoobank.org:act:116a2332-4326-4aae-9878-0827BC1CB330 43e Diagnosis Males of Z. logunovi sp. nov. differ from other congeners by a very short embolus directed forward (only Z. schmitzi possesses a similarly short embolus, but in the latter case the embolus is sidewarddirected, cf. Fig. 7A-D). The ventral brush of setae on the cymbium adjoining the bulb is more welldeveloped than in any other congener (cf. Figs 24C, 28G-I, 30I). Females are easily distinguished from all other female congeners except Z. annulipes and Z. halepensis sp. nov. by having long, non-divergent median receptacles considerably exceeding the small and globular lateral pair in size (Fig. 21). Females of Z. logunovi sp. nov. differ from those of Z. annulipes and Z. halepensis sp. nov. by the shape of their digitiform (not clublike) median receptacles, which are considerably thinner than in the two latter species (cf. Figs 5C-D, 9D-F).
Etymology this species is named after our friend and colleague Dmitri Logunov (university of Manchester, Manchester, UK).   palp (Figs 20,43e; paratype from the type locality). relatively long and slender; pale, without pigmentation; femur slightly longer than tibia; diameter of tibia 1.33 times wider than in femur; cymbium with distinct dorsal hump, about 2 times longer than bulb, retro-and proventral edge of cymbium with brush of thick and dense setae; bulb hemispherical; embolus short and screw-shaped, without a neck. Colour. as in male, but darker brownish pattern of carapace is more developed; dorsal abdominal pattern weaker; legs with a few incomplete darker fasciae.
eyeS. aMe 0.07, aLe 0.17, PLe 0.15, PMe 0.10, aMe-aMe 0.05. endogyne (Fig. 21). Both pairs of receptacles cylindrical and with corrugated stems; median receptacles twice as long as laterals and about 1.3 times thinner, median receptacles with wide base and, in anterior view (Fig. 21C), both pairs of same diameter; gland pores cover only tips of receptacles.

Variations
Carapace length in males varies from 1.15 to 1.40, in females from 1.47 to 1.85. Darker pattern and markings in some specimens are almost indistinct (Fig. 19A).
Ecology the species was found in steppe habitats in foothills and low mountains.

Zaitunia spinimana-group
Remarks this group includes relatively pale-coloured species. only the clypeus, post-ocular area and dorsal abdominal pattern are slightly darker than the pale background colour. Legs i-iV mostly lacking darker fasciae and are concolorous with the prosoma. the conical tegulum is much shorter than the cymbium. the embolus is relatively long to very long. the endogyne with two pairs of receptacles. three species are included: Z. martynovae (Andreeva & Tyshchenko, 1969) from Tajikistan, Z. spinimana sp. nov. from Turkmenistan and Kazakhstan, and (tentatively, because of the unknown male) Zaitunia inderensis Ponomarev, 2005 from western kazakhstan.

Diagnosis
By the structure of the vulva, females of Z. inderensis resemble Z. spinimana sp. nov., but differ from it and all other congeners by the enlarged lateral receptacles which are considerably larger and more robust than the median pair (Fig. 22E).

Variation
Carapace length in female paratypes varies from 1.19 to 1.40; no distinction in the colouration is evident.

Ecology
According to label data and the original description (Ponomarev 2005), the species was found in a low, semi-desert karst plateau.

Distribution
Known only from the type locality (Fig. 48).

Diagnosis
By the structure of the bulb, Z. martynovae resembles Z. spinimana sp. nov., but differs from it and all other male congeners by the long and flattened corkscrew-shaped embolus 43F). Females are easily distinguished from all other congeners due to their large and swollen median receptacles (Fig. 26).
palp (Figs 25a-d, 43F). relatively long and slender, 1.78 times longer than carapace; tibia 1.3 times wider than femur, dorso-distally with brush of strong setae; cymbium shorter than bulb; bulb sinuous in lateral view; spermophore sinuous in embolic neck; embolus longer than neck and equal in length to tegulum + neck, embolus bent twice.    Colour. as in male but abdomen slightly darker dorsally, and ventrally with darker X-shaped spot in genital area. eyeS. aMe 0.07, aLe 0.13, PLe 0.11, PMe 0.09, aMe-aMe 0.06. endogyne (Fig. 26A-C). With large and wide conical median receptacles and small globular lateral receptacles; median receptacles with corrugated stem, their bases separated by less than the diameter of their heads, heads separated by 1.6 of their diameter; heads of both pairs densely covered with gland pores.

Variations
Carapace length in females varies from 1.50 to 1.75. Variation of coloration and structure of the endogyne in females from Tajikistan and Turkmenistan is shown in Figs 23A-B, D-F, and 26D-O, respectively. the body and leg pattern, including the X-shaped spot in the female genital area, may be well-developed or indistinct.
Ecology this species is found in different habitats from foothill steppes and shrubland to open Juniperus forest in the middle mountain belt. The spiders occur under stones (Fig. 50B), and in crevices in clay escarps (Fig. 50C) and rock outcrops (Fig. 50D) where they build small webs below or near the cavity entrance (Fig. 50E, a close-up view of such a web is also shown in Fig. 50F).

Diagnosis
By structure of the bulb, Z. spinimana sp. nov. resembles Z. martynovae but differs from the latter by its shorter and narrower corkscrew-shaped embolus (cf. Figs 24A-D, 43F, 27C-E, 44A). Additionally, it differs from Z. martynovae and all other congeners by having a femur, tibia and metatarsus i with unusually numerous, long spines (Figs 27A). By structure of the vulva, Z. spinimana sp. nov. is similar to Z. inderensis; they differ by the receptacles, which are subequal in size (Fig. 27H-I).

Etymology
The specific epithet is derived from the Latin spina-(thorn, spine) and -manus (hand, appendage); the proposed name refers to the spiny legs of the male.  Note the holotype male and the only collected paratype female are very similar in possessing a very similar shape of the carapace (flattened more than usual) and a similar conformation of the eye group. Although the distance between the localities listed above is about 500 km, these slightly hilly and extremely uniform desert landscapes have no significant natural barriers. We prefer currently to treat these specimens as the same species (with no serious objectives against this assumption) rather than consider them representatives of two very close but distinct species.
Colour. Whole body and legs pale yellowish-white; clypeus with weak pale orange area; eye tubercle marked with medium to dark brown; abdomen dorsally with slightly darker and almost indistinct narrow median stripe.
palp 44a). Femur and tibia subequal in length; tibia 1.4 times wider than femur; cymbium slightly shorter than bulb; spermophore relatively wide; neck of embolus shorter than embolus proper; embolus long and arched, slightly screw-shaped.
SpeCial CHaraCterS. Leg i with numerous long spines, located proapically on femur, prolaterally and ventrally on tibia and metatarsus (Fig. 27A) Colour. as in male except for darker brownish abdomen.
eyeS. aMe 0.07, aLe 0.13, PLe 0.10, PMe 0.08, aMe-aMe 0.06. endogyne (Fig. 27H-I). Both pairs of receptacles club-like, lateral receptacles longer than median, with distinct corrugated stem; median receptacles separated by more than 2.5 diameters; gland pores cover entire head of lateral receptacles and only top of median ones.

Ecology
According to Zyuzin & Tarabaev (1994) and the label data, this species occurs in upland desert area; no more details are known.

Diagnosis
Males of Z. beshkentica are similar to those of Z. wunderlichi sp. nov. but differ from them by the shape of the raised embolic keel, which terminates more abruptly (cf. Figs 28G-I and 32D-F), whereas females rather resemble Z. huberi sp. nov. in vulval structure but differ from the latter species by the shape of the median receptacles, which are longer and tubiform (vs shorter and rounded in Z. huberi sp. nov., cf. Figs 29I-K, 30C-E). Color. Whole spider pale greyish-brownish-yellow; eye tubercle brownish-black; diffuse and narrow median line and margins of carapace light brown, as well as a weak and diffuse dorsal abdominal pattern consisting of interrupted median line anteriorly and a few transverse fasciae posteriorly.

Variations
Carapace length in females varies from 1.30 to 1.95. Specimens inhabiting lowland desert areas are almost uniformly pale-yellowish, with darker pattern on the body and legs very weak to absent (sometimes only metatarsi and tarsi are slightly dark, but the eye tubercle is still black). By contrast, specimens collected in foothills have typical though somewhat paler coloration, with darker median lanceolate spots on carapace and abdomen combined with weak and diffuse transverse fasciae on legs and in the posterior part of the abdomen (cf. Fig. 29A-H).

Ecology
Spiders were collected in desert, semi-desert and dry steppe biotopes in piedmont plains and low foothills and in the middle mountain zone with open park woods composed of Juniperus spp. and Acer spp.

Distribution
Known from western Tajikistan and southern Uzbekistan (Fig. 49).

Diagnosis
By structure of the vulva, Z. huberi sp. nov. resemble Z. beshkentica but differ from them by the shape of the median receptacles, which are shorter and round (vs longer and tube-shaped in Z. beshkentica, cf. Figs 29I-K, 30C-E).

Etymology
The specific epithet is given after the prominent German arachnologist Bernhard Huber, who has transferred some misplaced asian genera from Pholcidae to Filistatidae.

Description Female
HabituS. See Fig. 30a-B. body lengtH. 5.35. Colour. Whole body and legs light brownish-yellow, abdomen without dorsal pattern, tarsi of palp and legs i-iV darkened, eye tubercle blackish-brown.
eyeS. aMe 0.09, aLe 0.23, PLe 0.20, PMe 0.20, aMe-aMe 0.11. endogyne (Fig. 30C-E). Both pairs of receptacles globular, median receptacles separated from each other by the same distance as lateral and median receptacles; median receptacles larger, separated by less than one diameter. leg meaSurementS.

Note
See comments in the redescription of the holotype of Z. afghana. Both of these species are very similar in somatic characters, which probably misled Roewer (1962). Nevertheless, they differ so drastically in the structure of the endogyne (cf. Figs 14G-K, 30C-E) that they are assigned here to different species groups.

Distribution
Known only from the type locality (Fig. 49).

Diagnosis
The species differs from all other congeners in having modified metatarsi IV -shortened, distally dilated and flattened, with ventrodistal spines (Fig. 30G, K).

Etymology
The specific epithet is derived from psammo-(a combining form of the Greek psámmos, "sand") + -droma (a combining form of the Greek drómos meaning "running"), indicating the ability of these spiders to run on a loose, sandy substrate. Colour. Carapace light yellowish-orange, other parts of body and legs uniformly pale greyish-yellow.

Variation
Carapace length in males varies from 1.42 to 1.45; no differences in coloration are evident.

Ecology
the species occurs in an extra-arid sandy desert with Haloxylon shrubs; the males were collected in pitfall traps in rodent colonies. Most likely the spiders inhabit abandoned gerbil burrows.

Distribution
Known only from the type locality (Fig. 49).

Diagnosis
By the structure of the bulb, Z. wunderlichi sp. nov. is similar to Z. beshkentica, but differs by the shape of the raised embolic keel, which terminates more gently (cf. Figs 28G-I, 32D-F), whereas females differ by the structure of the vulva, which is more similar to those of Z. maracandica and Z. ferghanensis sp. nov.; they differ from both these species by the shorter median receptacles (cf. Figs 33a-B, D, 35E-J, 38E-I).
Etymology this species is named after the famous German arachnologist, our friend and colleague Jörg Wunderlich.

Note
although males and the only collected female listed above were found in two separate areas distant from each other by about 100 km, they are very similar in having the same type of body and legs colouration, as well as in possessing a very similar carapace setation and a similar conformation of the eye group. additionally, they all occurred in almost identical humid biotopes. thus, we have no doubt that in both these situations, we are dealing with representatives of the same species.
Colour. Whole spider pale greyish-yellow; eye tubercle dark brown; Y-shaped median spot occupying cephalic portion and extending to clypeus, and margins of carapace light brown; weak and diffuse dorsal abdominal pattern consisting of lancet-shape median spot anteriorly and a few transverse fasciae posteriorly pale brown.
palp 44d). Femur slightly longer than tibia (in prolateral view); tibia 1.5 times wider that femur; cymbium as long as bulb; keel of embolic neck with straight margin (not curved in terminal part); tip of embolus bent downward. Colour. as in male, except without darkened carapace margins.

Variation
Length of the carapace in males varies from 1.15 to 1.23, the body and leg colouration does not vary significantly.
Ecology the species inhabits humid habitats including walnut forest of Juglans regia L. in the middle mountain belt, where it certainly prefers more open slopes, occurring under stones in low forest and shrubs.

Zaitunia maracandica-group
Remarks this group includes pale-coloured species. the clypeus, post-ocular area, dorsal abdominal pattern and often also lateral margins of the carapace are slightly darker than the pale background colour. Legs i-iV with or without darker fasciae. the conical tegulum is much shorter than the cymbium. the relatively long embolus is provided with a low, more or less reduced keel. the vulva has two pairs of receptacles. Five species are included: Z. ferghanensis sp. nov. from kyrgyzstan and uzbekistan, Z. maracandica (Charitonov, 1946) from Uzbekistan and Kazakhstan and (tentatively, because of unknown male characters) Z. feti sp. nov. from turkmenistan, Z. minuta sp. nov. from uzbekistan and Z. zonsteini Fomichev & Marusik, 2013 from kazakhstan. Key to species of the Zaitunia maracandica-group (Males of Z. feti sp. nov., Z. minuta sp. nov. and Z. zonsteini are unknown.)
Colour. Whole spider medium brownish-yellow; eye tubercle dark brown to brownish-black; cephalic part, clypeus and margins of carapace not darkened; abdomen with almost indistinct and slightly interrupted darkened spot dorso-medially.
palp 44e). Femur and tibia subequal in length; tibia about 1.5 times wider than femur, wider in basal part than terminally; cymbium and bulb subequal in length; bulb conical, embolus gently curved and roundly bent subapically.    Colour. as in male but darker brownish pattern on carapace (narrow median longitudinal stripe, two shorter stripes located sideward and reticulated area covering clypeus) better developed; abdomen also a little darker, with almost indistinct brownish median spot. eyeS. aMe 0.09, aLe 0.14, PLe 0.13, PMe 0.10, aMe-aMe 0.05. endogyne ( Fig. 35E-J). Receptacles separated; median receptacles club-like, and lateral receptacles subconical; both pairs with corrugated stems; head of median receptacles separated by 3 diameters; pores present on stem and heads.

Variation
Carapace length in males varies from 1.35 to 1.47, in females from 1.40 to 1.65. Carapace with clypeus and margins slightly or noticeably pigmented; abdomen with or without weak dorsal pattern, consisting of a narrow interrupted median stripe and several pairs of short lateral chevrons (Figs 34,35K).
Ecology this species inhabits semi-desert habitats in piedmont plains and foothills.

Etymology
The specific epithet is given after our good friend and colleague Victor Fet (Marshall University, USA), the author of many works devoted to the spiders of turkmenistan.
Colour. Body and legs pale brownish-yellow, carapace with contrasting median brown spot and narrow margins; eye tubercle blackish-brown; legs with distinct brown marks and fasciae; abdomen with brown dorsal pattern consisting of contrasting narrow median band combined in posterior third with a few pairs of weak and diffuse lateral chevrons.
eyeS. aMe 0.07, aLe 0.14, PLe 0.12, PMe 0.10, aMe-aMe 0.07.  (Fig. 36D-E). Median receptacles club-like, wide at the base, with heads wider than stems, pore glands cover whole receptacle, receptacles separated by almost 3 diameters of heads. Lateral receptacles globular, with short stems; heads of median and lateral receptacles separated by one diameter. leg meaSurementS. Ecology the holotype was collected in the desert foothill area; other details are unknown.

Diagnosis
in the structure of the bulb, males of Z. maracandica resemble Z. ferghanensis sp. nov. but differ by the shape of the palpal tibia, which is considerably more swollen, with thickened setae, as well as by the shape of the vestigial embolic keel, which terminates more gently than in Z. ferghanensis sp. nov. (cf. Figs 35A-D, 37A-C). In the structure of the endogyne, Z. maracandica is similar to Z. ferghanensis sp. nov. and Z. wunderlichi sp. nov., but differs from these species by the shape of the median receptacles, which are subequal to the lateral pair (by contrast, in Z. ferghanensis sp. nov. the lateral receptacles are largest, cf. Fig. 38E-J) and from Z. wunderlichi sp. nov. by the longer median receptacles (which are shorter in the latter species; cf. Fig. 33A-B, D).
palp 44F). Femur longer than tibia and almost 2 times thinner; tibia with a few strong ventro-retrolateral setae in terminal part; cymbium subequal in length to bulb; bulb conical; embolic part long and straight; ventral keel of embolic neck gradually tapering; tip of embolus bent downward.  Colour. as in male, but abdomen dorsally and carapace margins are somewhat darker.

Variation
Carapace length in females varies from 1.53 to 2.05; spiders may have darker marks and fasciae on the carapace, legs and abdomen, or these marks may be almost completely absent on the pale background (cf. Fig. 38A-D).
Ecology this species occurs in various habitats from foothill deserts, steppes and shrubs to open Juniperus forest in the middle mountain belt.

Etymology
The specific epithet is derived from the Latin minutus ("little, small, minute").  Colour. Body and legs pale sandy-brown, carapace with slightly darker median spot and noticeably darker narrow margins; eye tubercle blackish-brown; abdomen uniformly colored, without dorsal pattern.
Ecology this species has been collected in steppe and semi-desert biotopes.

Remarks
unlike Zaitunia, females of Pholcoides monticola lack the metatarsal crest on leg IV (Fig. 45C), typical for the Filistatinae. However, this species shares relatively long legs, a rounded carapace and sternum (cf. Fig. 45a and Zonstein et al. 2013: figs 1-2, respectively), spinnerets located closer to the posterior tip of the abdomen (Fig. 45B and Zonstein et al. 2013: fig. 3) and a characteristic calamistrum with two posteriorly convergent rows of setae with P. afghana (Fig. 45C and Zonstein et al. 2013: fig. 6). The structure of the endogyne in these two species is also similar, although not identical (cf. Figs 45D-e and Zonstein et al. 2013: figs 11-12), and differs drastically from that in Zaitunia. Based on these reasons, the aforementioned species is transferred from Zaitunia to Pholcoides. its relationship to P. afghana and other congeners (some of which are still undescribed) will be considered in detail in a taxonomic revision of Pholcoides (Zonstein & Marusik, in prep.).

Taxonomic placement of Zaitunia
When Lehtinen (1967) described Zaitunia, he allocated it to a group of three genera along with Filistata Latreille, 1810 and Kukulcania Lehtinen, 1967. He stated that, unlike other family members, all three genera share the presence of spines on leg femora, tarsi and metatarsi, and the ejaculatory duct (spermophore) is tightly coiled. Concurrently, he considered Zaitunia as lacking tarsal spines and possessing the calamistrum composed of "2 rows of 2 strong bristles" (Lehtinen 1967: 300, table 3). Contrary to that statement, during the present revision, we found that all Zaitunia species have few to numerous short tarsal spines on the legs. the calamistrum in Zaitunia was found to be uniseriate,       (Andreeva & Tyshchenko, 1969) in Southern Tajikistan. A. ♀ of Z. schmitzi (Kulczyński, 1911) on the turned stone bottom (Israel, Shoham, image courtesy of Amir Weinstein). B. Surroundings of Gandzhina: the spiders occur under the stones. C. Pyandzh-karatau Mts, one of the co-authors (YM) near the clay escarps inhabited by filistatids. D. Foothills of the salt dome khodzha-Mumin: the spiders inhabit crevices and cavities in the rock outcrops. E. One of the figured escarps, a close-up photo showing the twin entrance leading to the inhabited cavity. F. Captured ♀ of Z. martynovae (Andreeva & Tyshchenko, 1969) weaving a web in the cover of a collecting tube.
although it is certainly composed of two or even three formerly separate but now fused and juxtaposed rows of 6-7 large and 2-6 tiny setae each ( Fig. 2A-E). Brignoli (1982) was more interested in emphasizing distinctive features of Zaitunia than in finding its affinities. He first provided a figure showing the calamistrum of Zaitunia with a characteristic gap between two groups of setae (Op. cit.: fig. 14) like that in Filistata insidiatrix (Op. cit.: fig. 1). However, regarding Z. persica, he presented setae of the calamistrum as having serrate margins and being dilated apically, which has not been confirmed by our examination. We found that the structure of these setae in Z. persica does not differ from that in any other Zaitunia: similarly as with Filistata and Kukulcania, the ribbed and curled lanceolate bristles in Zaitunia are gradually widened in the medial part and then are also gradually narrowed to an acute apex ( Fig. 2B-C, E). It is notable that none of Brignoli's figures presenting the calamistra of Filistata, Zaitunia and Sahastata Benoit, 1968 showed them in the correct position on the elevated process (Op. cit.: figs 1, 14 and 17, respectively). Gray (1995) tentatively assigned Zaitunia to the nominative subfamily which was considered to certainly include Filistata, Kukulcania and Sahastata. However, because of scarce data, Zaitunia was not included in the cladogram. For the same reason,  confirmed the provisional placement of Zaitunia among the Filistatinae, but did not include this genus in their phylogenetic reanalysis of the filistatid genera. Zonstein (2009b) also noted a close similarity between Zaitunia, Filistata and Kukulcania because they "share a relatively broad cribellum and a very short uniseriate calamistrum in the female, as well as curved pseudosegmented tarsi in the male" (Op. cit.: 126). Having re-examined these characters, we found the two latter ones to be significant, whereas the former appears to be confusing. Although Zonstein et al. (2013) and later Marusik & Zamani (2015) in these regional works listed Zaitunia in the Filistatinae, they have entirely focused on the characters distinguishing Zaitunia from the similarlooking Filistata, without considering the taxonomic position of Zaitunia in detail. thus, the above-listed suggestions concerning the taxonomic position of Zaitunia should not be recognized as sufficiently substantiated. An attempt to clarify this question as far as possible has been one of the targets of this study.
During this revision we found that all species assigned to Zaitunia share the following characters (listed according to the description format accepted here): 1) body and legs are covered only with ciliate hairs (Fig. 1D-I); plumose hairs (like those showed by Gray 1995: fig. 10, andLise et al. 2010: figs 12, 20) are absent 2) clypeus short, broad-oval, and steeply inclined 3) clypeus with a comb of long, thick and dense reclined setae (see Fig. 42), more developed in males (Fig. 42A, D, F-G) 4) eye tubercle low or moderately high (probably associated with character 2) 5) thoracic fovea is absent 6) labium usually wider than long (maximum as wide as long); this character is also probably associated with character 2 7) male palpal tibia moderately short and more or less swollen 8) cymbium subcylindrical in shape 9) cymbium relatively short (at least shorter than that in most filistatines; probably associated with character 7) 10) spermophore tightly coiled 11) leg femora, tibiae and metatarsi I-IV spinose 12) calamistrum consists of setae located on a metatarsal crest distant from each other (see Marusik et al. 2014: fig. 25). The state of these characters in Microfilistata is unclear.
in total, 16 of 24 considered states in Zaitunia are shared with Filistata; 14 with Kukulcania; 11 with Microfilistata; 10 with Sahastata, and only a few, including absence of the thoracic fovea, whose taxonomic significance is not completely evident, with genera of the Prithinae. We thus conclude that Zaitunia should be placed in the nominative subfamily Filistatinae s. str. it will be possible to consider further details concerning its allocation within the subfamily and intergeneric relationships when the filistatine genera Filistata and Sahastata are revised (both genera are currently under study; Zonstein & Marusik, in prep.).