Revision of the spider family Zodariidae (Arachnida, Araneae) in Iran and Turkmenistan, with seventeen new species

Abstract Species of the spider family Zodariidae occurring in Iran and Turkmenistan are reviewed. Seventeen species of three subfamilies are described as new to science: Lachesana kavirensissp. nov. (♂, Qom; northern Iran), L. perseussp. nov. (♂, Alborz; northern Iran) (Lachesaninae), Pax ellipitasp. nov. (♂♀, Kermanshah and Lorestan; western Iran), P. leilasp. nov. (♂♀, Fars; southwestern Iran) (Storeninae), Acanthinozodium armitasp. nov. (♂, Tehran; northern Iran), A. atrisasp. nov. (♂♀, Qazvin and Tehran; northern Iran), A. diarasp. nov. (♂, Ilam and Lorestan; western Iran), A. dorsasp. nov. (♂♀, Fars; southern Iran), A. elburzicumsp. nov. (♂♀, Tehran; northern Iran), A. kianasp. nov. (♂, Kurdistan; western Iran), A. masasp. nov. (♂, Kermanshah; western Iran), A. niushasp. nov. (♂♀, Fars, Isfahan and Markazi; central Iran), A. ovtchinnikovisp. nov. (♂, Mary; southeastern Turkmenistan), A. parmidasp. nov. (♂, Isfahan; central Iran), A. parysatissp. nov. (♂♀, Ardabil and Qazvin; northern and northwestern Iran), A. soranisp. nov. (♂, East Azerbaijan and Kurdistan; northwestern and western Iran) and Trygetus susianussp. nov. (♀, Khuzestan; southwestern Iran) (Zodariinae). Zodariellum Andreeva & Tyshchenko, 1968, currently comprising only the type species (Z. surprisum Andreeva & Tyshchenko, 1968) is rediagnosed, with the following species being (re)transferred to it: Z. asiaticum (Tyshchenko, 1970) comb. res., Z. bactrianum (Kroneberg, 1875) comb. nov., Z. bekuzini (Nenilin, 1985) comb. res., Z. chaoyangense (Zhu & Zhu, 1983) comb. res., Z. continentalis (Andreeva & Tyshchenko, 1968) comb. res., Z. furcum (Zhu, 1988) comb. res., Z. mongolicum Marusik & Koponen, 2001 comb. res., Z. proszynskii (Nenilin & Fet, 1985) comb. res., Z. nenilini (Eskov, 1996) comb. res., Z. surprisum Andreeva & Tyshchenko, 1968 comb. res., Z. schmidti Marusik & Koponen, 2001 comb. res., Z. sytchevskajae (Nenilin & Fet, 1985) comb. res. and Z. volgouralensis Ponomarev, 2007 comb. res. (all ex. Zodarion); out of these, Z. proszynskii Nenilin & Fet, 1985, previously known only from the type locality in Turkmenistan, is recorded from northeastern Iran for the first time. This paper raises the number of zodariids known from Iran to 22 species from seven genera (including the first Iranian records of Acanthinozodium Denis, 1966, Pax Levy, 1990 and Zodariellum) and those known from Turkmenistan to five species from five genera (including the first record of Acanthinozodium from this country). Regional distribution records of all species are mapped.


Introduction
The spider family Zodariidae Thorell, 1881 comprises 1186 extant species in 87 genera and five subfamilies globally, as well as 11 species in nine genera known from fossils (Jocqué 1991;Dunlop et al. 2020;WSC 2021). Most of the species occur in the tropical and subtropical regions, with a few genera recorded from the Palaearctic (Jocqué and Dippenaar-Schoeman 2006). Despite their relatively high species richness, this family remains one of the most poorly documented groups of spiders in Iran and Turkmenistan. The first record of Zodariidae in Iran was provided by Ovtchinnikov et al. (2009), and currently, there are only six species belonging to four genera of this family known from this country (Zamani et al. 2021). Four species are known from Turkmenistan, three of which are endemics (Mikhailov 2013;WSC 2021). Recently, we had the opportunity to examine a relatively large collection of Iranian and Turkmen zodariids, in which 17 species are new to science, and two genera and one species in Iran and one genus in Turkmenistan are recorded for the first time. All of the species occurring in this region are surveyed, their distributions are mapped, and the new species are described and illustrated herein.

Material and methods
Specimens were photographed using an Olympus Camedia E-520 camera attached to an Olympus SZX16 stereomicroscope or to the eye piece of an Olympus BH2 transmission microscope, and a SEM JEOL JSM-5200 scanning electron microscope at the Zoological Museum of the University of Turku. Digital images were prepared using CombineZP image stacking software. Illustrations of internal genitalia were made after clearing them in a 10% KOH aqueous solution. Lengths of leg segments were measured on the dorsal side. Measurements of legs are listed as: total length (femur, patella, tibia, metatarsus, tarsus). All measurements are given in millimetres. Jocqué, 1991 Fig. 32  Etymology. The specific epithet refers to the legendary founder of Mycenae and of the Perseid dynasty in Greek mythology. Noun in apposition.

Lachesana insensibilis
Diagnosis. The new species differs from the similar L. kavirensis sp. nov. by the lack of spines in the proximal mesal part of the cymbium (vs. present; cf. Fig. 3C and 3H) as well as by the RTA with the terminal half almost as wide as the tibia (vs. 1.5 times thinner). The shape of the RTA readily distinguishes this species from congeners in the region (stalk abruptly bent basally and tip gently curved; cf. Fig. 4J and 4F-I).
Palp as in Figs 3F-J, 4J. Tibial apophysis more than 1.2 times longer than tibia, widest part over 2 times wider than stalk; cymbium lacking spines in proximal mesal part; bulb elongate, 1.3 times longer than wide; base of embolus large, 1/3 of the tegulum's length; embolus filamentous, lying in chute formed by conductor.
Female. Unknown. Distribution. Known only from the type locality in Alborz Province, northern Iran (Fig. 32).
Subfamily Storeninae Simon, 1893 Genus Pax Levy, 1990 Type species. Habronestes libani Simon, 1873 from Lebanon. Comments. This is a small genus with seven species distributed exclusively in the Middle East from Turkey and Israel to Iran (including the two new species described below, representing the first record of this genus in this country). Members of this genus can be easily distinguished from all other zodariids found in the study area by the ovoid carapace lacking a distinct separation between the cephalic and thoracic parts and also by males having a modified cymbium.   Kermanshah, 34°28'N, 47°00'E, 18.06.1975 (A. Senglet).
Etymology. The specific epithet refers to Ellipi, an ancient kingdom located on the western side of the Zagros Mountains, between Babylonia at the west, Media at the north-east, Mannae at the north and Elam at the south.
Diagnosis. The new species differs from P. leila sp. nov., the only other Pax species known from Iran, by the cymbium lacking a horn-like outgrowth (vs. present), by having a deep lateral fold of the cymbium (vs. lacking) and the epigyne with a rectangular median plate (vs. triangular). The male of P. ellipita sp. nov. differs from the rest of the species by having the RTA shorter than the tibia (vs. longer), while the female differs by the trilobate posterior margin of the epigyne (vs. solid) and copulatory openings located posteriorly (vs. anteriorly).
Comments. Because the female specimen was not collected together with the male, its assignment to this species is tentative and shall be confirmed when both sexes are collected together.
Distribution. Known from the listed localities in Kermanshah and Lorestan provinces, western Iran (Fig. 32).
Etymology. The specific epithet is a feminine given name in the Persian language, meaning "daughter of the night". Noun in apposition.
Diagnosis. The new species differs from all congeners by having a pair of spinelike cymbial outgrowths (Figs 8, 9C, E) (vs. lacking) and a triangular epigynal median plate (vs. absent or rectangular).
Comments. Although this species is rather similar in general appearance to the generotype and the other species known from Iran, the bulb and cymbium conformation is very different from those of other species considered in Pax and most likely belongs to an undescribed genus.
Distribution. Known only from the type locality in Fars Province, southwestern Iran (Fig. 32). This is the easternmost record of the genus Pax.
Comments. Twelve species are currently considered in this genus which are known from the Maghreb and adjacent countries in the south and in Socotra; previously, it was not recorded from Iran (WSC 2021). Although the male of the type species is unknown, Acanthinozodium comprises species having a unique gland located in the conical pit of the cymbium. The 11 species occurring in Iran and the one from Turkmenistan have the same conical pit and are thus placed in this genus; however, the male palpal tibia and particularly the bulbs are different from those of the species occurring in the Maghreb and surrounding countries. Further revisions are needed to clarify the taxonomy and composition of this group, which are beyond the scope of this paper. Etymology. The specific epithet is a Persian feminine name meaning "queen of fire". Noun in apposition.
Diagnosis. The new species has an abdominal pattern similar to that of A. parysatis sp. nov. (dark abdomen with posterior median white stripe; less distinct in males) but differs by having a dark pars cephalica (vs. pale). The male palps of the two species differ by the RTA being thin and more than twice as long as the tibia in A. atrisa sp. nov. (vs. broad and as long as the tibia). Judging by the shape of the male palp, A. atrisa sp. nov. is probably closely related to A. sorani sp. nov., as both have a long RTA, a similarly shaped median apophysis, a posterior tegular process and an embolus with an anterior process. The two species differ by the shape of the embolic process (Ep), wider than the embolus and gently bent at the tip in A. atrisa sp. nov. (vs. spine-like). The epigyne of A. atrisa sp. nov. is most similar to that of A. parysatis sp. nov. in having a small anterior hood and a thin furrow between the lateral lobes but differs by the furrow being shorter than the bursa copulatrix (vs. the same length as bursa). Palp as in Figs 12A-C, 14A-C. RTA (Ra) long (almost as long as bulb) and thin (7 times longer than wide), slightly bent; tegulum with posterior process, sperm duct almost straight retrolaterally, and gently bent along prolateral side; median apophysis wider than long; embolus (Em) broad basally, originating at about the 7 o'clock position, bifurcated terminally, with broad anterior process; embolus proper thin and straight, with small tubercle. Epigyne as in Fig. 16A-D. Fovea triangular, and small anterior hood present; bursae subtriangular, wider anteriorly, separated by less than one of their radii; receptacles smaller than bursae, ovoid, separated by more than 3 diameters.
Distribution. Known only from the listed localities in Tehran and Qazvin provinces, northern Iran (Fig. 32). Etymology. The specific epithet is a Persian feminine name meaning "good listener". Noun in apposition.

Acanthinozodium niusha
Diagnosis. The male of the new species is most similar to A. parysatis sp. nov. from which it differs by a sharply tapering RTA, shorter tip of the cymbium (cf. Fig. 13A and 13F), the angle of embolus and the claw of the median apophysis. Female of A. niusha sp. nov. is similar to A. atrisa sp. nov. by having a similar anterior hood but differs by the longer furrow leading to the hood (as long as receptacle vs. 1.5 times shorter) and less separated receptacles.
Distribution. Known only from the listed localities in Fars, Isfahan and Markazi provinces, central Iran (Fig. 32).    Etymology. The new species is named after our late colleague Sergei V. Ovtchinnikov who made important contributions to the study of Central Asian spiders.
Diagnosis. The new species differs from the congeners in the region (except A. parmida sp. nov.) by the small size of the palp (cymbium 0.28 long, vs. > 0.6) and the unique embolus which has a long furrow with a serrate ventral margin (Fig. 22E). It is closely related to A. parmida sp. nov. from central Iran, from which it can be differentiated by having a longer than wide bulb (vs. almost as long as wide), relatively longer cymbium (length/width ratio 1.86 vs. 1.4), and a different shape of median apophysis (anterior portion larger than posterior one, vs. opposite; cf. Figs 22G and 19B).
Description. Male. Body missing. It is assumed to be a very small zodariid based on the size of the palp.
Palp as in Fig. 22D, E, G-I. Ventral tibial apophysis lacking; RTA almost triangular, slightly longer than tibia; cymbium longer than wide; sperm duct tracking margin of tegulum, lacking any turns; median apophysis longer than wide, with anterior part larger than posterior part; embolus originates at about 8:00 o'clock position with its terminal 2/3 having a longitudinal furrow (Er) with a finely serrated ventral margin.
Female. Unknown. Note. The sample collected by Gromov contained three males. The palp of one specimen was dissected and imaged with a SEM in 2005. Then, all three specimens were given to Ovtchinnikov who was planning to revise this group. After his death, his collection was transferred to Almaty, Kazakhstan by Alexander Gromov and is now inaccessible.
Distribution. Known only from the type locality in Mary Region, southeastern Turkmenistan (Fig. 32). Etymology. The specific epithet is an ancient Persian feminine name, meaning "fairy-like". Noun in apposition.

Acanthinozodium parysatis
Diagnosis. The male palp of A. parysatis sp. nov. is similar to that of A. niusha sp. nov. by the overall shape of the median apophysis and RTA but differs in lacking a retrolateral extension on the median apophysis (vs. present) and by having a shorter, stouter RTA (vs. longer and tapering). The epigyne of the new species is most similar to that of A. atrisa sp. nov. in having a small anterior hood and thin furrow between the lateral lobes but differs by the relative length of the anterior part of the fovea being shorter than the bursae in A. parysatis sp. nov. (vs. as long as bursae).
Distribution. Known only from the listed localities in Ardabil and Qazvin provinces, northern and northwestern Iran (Fig. 32). Etymology. The specific epithet refers to a dialect or a language of the Kurdish languages that is spoken in Iraq, mainly in Iraqi Kurdistan, as well as the Kurdistan Province, Kermanshah Province, and West Azerbaijan Province of western Iran. Noun in apposition.

Acanthinozodium sorani
Diagnosis. The new species is most similar to A. atrisa sp. nov. but differs by embolic anterior process which is spine-like and thinner than the embolus proper in A. sorani sp. nov. (vs. broad) and also by the relative length of the RTA, longer than bulb in A. sorani sp. nov. (vs. shorter).
Female. Unknown. Distribution. Known only from the listed localities in Kurdistan and East Azerbaijan provinces, western and northwestern Iran (Fig. 32). Etymology. The specific epithet is a Persian feminine name meaning "righteous", "virtuous", and "good". Noun in apposition.
Diagnosis. The new species differs from all congeners in the region by the short RTA having 2 claw-like outgrowths on the tip (vs. tip tapering and lacking 2 claws) and also by the modified embolus, widening near the tip (vs. unmodified).
Female. Unknown. Distribution. Known from the type locality in Tehran Province, northern Iran, and another locality (Haji Abad) which cannot be georeferenced because there are many places with this name in Iran (Fig. 33). Etymology. The specific epithet is a Persian feminine name meaning "motherland". Noun in apposition.
Diagnosis. Acanthinozodium diara sp. nov. is very similar to A. masa sp. nov. by the shape of the RTA, the ventral tibial apophysis and the curvature of the embolus but differs by the shape of the median apophysis (cf. Fig. 20D and 20J) and the conductor having a small retrolateral indentation apically (vs. without indentation).  Fig. 18B. Total length 4.70. Carapace 2.18 long, 0.92 wide at pars cephalica, 1.52 wide at pars thoracica. Eye sizes and interdistance of PMEs: AME: 0.19, ALE: 0.12, PME: 0.10, PLE: 0.12, PME-PME: 0.25. Carapace dark brown, with irregular dark patches and lines. Sternum, labium and maxillae light brown. Chelicera dark brown, with one retromarginal tooth. Legs light brown, without annulations. Abdomen black, without any pattern; slightly lighter ventrally. Spinnerets pale, uniform in color. Measurements of legs Palp as in Figs 20D-F, 23D-F. RTA long and conical, with a small projection apically (Fig. 20E); tegulum with posterior process; ventral apophysis small and conical; median apophysis almost as long as wide, with outgrowths on all four sides, posterior one largest; embolus broad basally, originating at about the 7 o'clock position; embolus proper thin and steadily curving.

Description. Male (holotype). Habitus as in
Female. Unknown. Distribution. Known from the listed localities in Ilam and Lorestan provinces, western Iran (Fig. 33). Etymology. The specific epithet is a Persian feminine name meaning "precious". Noun in apposition.

Acanthinozodium dorsa
Diagnosis. The male of the new species differs from all congeners in the region by the prolateral outgrowth of the median apophysis projecting ventrally (vs. not projecting) and the long and broad RTA, apically twisted and lacking outgrowths (vs. with 1-2 outgrowths). The female is most similar to A. elburzicum sp. nov. but differs by the epigynal hood being longer than wide (vs. wider than long) and less separated receptacles (ca. 2.5 times of their diameter vs. 4).
Female. Habitus as in Fig. 18D. Total length 4.46. Carapace 1.93 long, 0.96 wide at pars cephalica, 1.46 wide at pars thoracica. Eye sizes and interdistance of PMEs: AME: 0.15, ALE: 0.12, PME: 0.09, PLE: 0.10, PME-PME: 0.15. Coloration as in male. Measurements of legs: I: 6.58 (1.39, 0.56, 1.52, 1.88, 1.23 Epigyne as in Fig. 30F-I. Epigynal plate over 3 times wider than long; anterior hood ca. 2 times longer than wide; receptacles subrectangular, separated by about 2.5 times of their widths. Distribution. Known from the type locality in Fars Province, southern Iran (Fig. 33). Diagnosis. The male of the new species is similar to A. diara sp. nov. by the similar shape of the RTA and they both have a small retrolateral indentation apically on the conductor, but differs by the shape of the median apophysis, which is with a distinctly larger prolateral outgrowth. The female of the new species is most similar to A. dorsa sp. nov. but differs by the epigynal hood being wider than long (vs. longer than wide) and receptacles separated by about 4 diameters (vs. ca. 2.5).
Palp as in Figs 21D-F, 23A-C, 29A. RTA (Ra) long and conical, with a small projection apically; ventral apophysis (Va) small and conical; tegulum with posterior process; median apophysis (Ma) wider than long, with outgrowths on all four sides, the triangular prolateral one the largest; embolus (Em) originating at about the 7 o'clock position; embolus proper thin and steadily curving.
Epigyne as in Fig. 30A-E. Epigynal plater over 3 times wider than long, lacking fovea; anterior hood wider than long; receptacles with rounded posterior parts, separated by about 4 diameters.
Etymology. The specific epithet is a Kurdish feminine name meaning "nature". Noun in apposition.
Diagnosis. The new species is very similar to A. masa sp. nov. by the shape of the RTA and ventral tibial apophysis but differs by the shape of the prolateral outgrowth of the median apophysis which is finger-like and triangular, terminally with a small hook (vs. broad and bifurcated).
Female. Unknown. Distribution. Known from the type locality in Kurdistan Province, western Iran (Fig. 33). Etymology. The specific epithet is a Kurdish feminine name, meaning "bright like the moon". Noun in apposition.

Description. Male (holotype). Habitus as in
Palp as in Figs 20I-K, 24A, B. RTA long and conical, with a small projection apically; tegulum with posterior process (Tp); ventral apophysis relatively large and conical; median apophysis wider than long, with a large bifurcated outgrowth prolaterally; embolus broad basally, originating at about the 7 o'clock position; embolus proper thin and steadily curving.
Female. Unknown. Distribution. Known only from the type locality in Kermanshah Province, western Iran (Fig. 33). Etymology. The specific epithet refers to a Persian princess, the only daughter of Bardiya (Smerdis), son of Cyrus the Great. Noun in apposition.
Diagnosis. The new species differs from the congeners in the region by its smaller size, by having a dorsal scutum, and by the very short tip (free part) of embolus having a long furrow with a serrate ventral margin. It is closely related to A. ovtchinnikovi sp. nov. from southeastern Turkmenistan, from which it can be differentiated by having an almost as long as wide bulb (vs. longer than wide), shorter cymbium (length/width ratio 1.4 vs. 1.86), a different shape of median apophysis (posterior portion larger than anterior one, vs. opposite) and the different position of embolic base (6:30 o'clock, vs. 8:00 o'clock; cf. Figs 19B and 22G).
Distribution. Known only from the type locality in Turkmenistan. Epigyne as in Fig. 27E-H. Epigyne medially with posteriorly diverging sclerotization, laterally with translucent fertilization ducts; receptacles almost round, separated by less than their diameters, each subdivided into a large, weakly sclerotized part and a smaller, more heavily sclerotized part.
Diagnosis. All Zodariellum spp. are clearly different from the generotype of Zodarion, Z. nitidum (Audouin, 1826), and the species considered in the genus by the following: 1) the cymbium has a tutaculum (vs. lacking in Z. nitidum and other species groups); 2) filamentous embolus starting at the 5 o'clock position (vs. 6 -in Z. nitidum, and many species groups of Zodarion sensu lato lack the filamentous embolus, or they are not at 5 o'clock position); 3) the absence of a ventral tibial apophysis (vs. present in the generotype) and 4) the shape of the RTA: one arm longer than wide, with claw or wart-like outgrowth posteriorly from the tip (vs. wider than long, with 3 branches in the generotype and various shapes in other species groups). Females of the two generotypes have long and twisted copulatory ducts, but they are converging in Zodariellum and diverging in Zodarion.
To illustrate the conformation of the male palp in Zodariellum (male specimens are lacking among the material studied in Iran and Turkmenistan), we have provided figures of Z. bactrianum, a species previously known from the original description only and previously considered in Zodarion.

Records in Turkmenistan.
Balkan (Nenilin and Fet 1985) (Fig. 33). Distribution. Previously known only from the type locality in Turkmenistan. This is a new record for Iran, representing the southernmost record in the known genus range.
Comments. This species is known from the original description only.

Genus Zodarion Walckenaer, 1826
Type species. Enyo nitida Audouin, 1826 from Egypt. Comments. With 177 named species, this is the largest genus within Zodariinae (WSC 2021). Based on the copulatory organs of the species currently considered in this genus, Zodarion does not appear to be monophyletic. There are only two species that are morphologically similar to the generotype: Z. luctuosum (O. Pickard-Cambridge, 1872) and Z. lutipes (O. Pickard-Cambridge, 1872), from the eastern Mediterranean (east of Tunisia to Iran). Comments. Until recently, Z. expers was placed in Ranops Jocqué, 1991, but now it is placed in Zodarion. It differs from the generotype of Ranops, but the copulatory organs are also very different from those of Z. nitidum, and most likely it represents a separate genus. Although it is absent in Iran or Turkmenistan, we have provided figures of this species to illustrate the differences with the generotype of Zodarion. Furthermore, although Z. nitidum has not been recorded in the current study area, because it is the type species of the genus, we have provided illustartions for this species as well, to demonstrate its differences with other Zodariinae genera in Iran and Turkmenistan, as well as with other species currently placed in Zodarion. Fig. 33 Acanthinozodium buettikeri Ono & Jocqué, 1986: 7, f. 1-4 (♂♀). Zodarion buettikeri : Levy 1992: 85;: 69, f. 3E-F (♂).