﻿A survey of the spider genus Dysdera Latreille, 1804 (Araneae, Dysderidae) in Iran, with fourteen new species and notes on two fossil genera

﻿Abstract The taxonomy of the Iranian species of the dysderid spider genus Dysdera Latreille, 1804 is revised. Currently, the only species of this genus known from Iran is D.pococki Dunin, 1985, albeit on the basis of a doubtful record. The following 14 species are described as new to science in this paper: D.achaemenessp. nov. (♀; Fars), D.bakhtiarisp. nov. (♂; Chaharmahal & Bakhtiari), D.damavandicasp. nov. (♂; Mazandaran), D.genoensissp. nov. (♂♀; Hormozgan), D.hormuzensissp. nov. (♀; Hormozgan), D.iranicasp. nov. (♂♀; Fars, Hormozgan), D.isfahanicasp. nov. (♂♀; Isfahan), D.mazerunisp. nov. (♀; Mazandaran), D.medessp. nov. (♂; Tehran), D.persicasp. nov. (♂♀; Golestan, Mazandaran), D.sagartiasp. nov. (♂♀; Tehran), D.tapuriasp. nov. (♂♀; Mazandaran), D.verkanasp. nov. (♂; Golestan), and D.xerxesisp. nov. (♂; Bushehr). Distribution records of all species are mapped. Also, the taxonomy of Mistura Petrunkevitch, 1971 and Segistriites Straus, 1967, two fossil genera currently considered in Dysderidae, is discussed and the latter is transferred to Segestriidae.


Introduction
The spider family Dysderidae C.L. Koch, 1837 comprises 591 extant species in 25 genera distributed in the West Palaearctic (WSC 2022). Most species have limited dispersal abilities and very small ranges; one exception is Dysdera crocata C.L. Koch, 1838 which has a cosmopolitan distribution due to anthropogenic transportations (Jocqué and Dippenaar-Schoeman 2006).
Although the first record of this family in Iran dates back to late 19 th century (Pocock 1889), the dysderid fauna of this country remains almost completely unknown. Currently, there are only three species of Dysderidae known from Iran: Dysdera pococki Dunin, 1985, Dysderella transcaspica (Dunin & Fet, 1985), and Harpactea parthica Brignoli, 1980(Zamani et al. 2022a. The Iranian records of several species (i.e., Dysdera aculeata Kroneberg, 1875, D. asiatica Nosek, 1905, D. erythrina (Walckenaer, 1802, Harpactea babori (Nosek, 1905), H. dobati Alicata, 1974, andTedia oxygnatha Simon, 1882) were recently considered as misidentifications and subsequently these species were rejected from the checklist of Iranian spiders (Zamani et al. 2017(Zamani et al. , 2022b. Recently, we had the opportunity to examine a collection of Iranian specimens of Dysdera Latreille, 1804, in which 14 species new to science were detected. In this paper, all species of this genus occurring in Iran are surveyed, their distributions are mapped, and those new to science are described and illustrated. Additionally, the taxonomy of two fossil genera currently considered in Dysderidae is discussed, and one of them is herein transferred to Segestriidae.

Materials and methods
Photographs of specimens and their copulatory organs were obtained using a Nikon D300S DSLR camera attached to a Nikon S800 stereomicroscope, a Tucsen TrueChrome Metrics microscope camera attached to a Nikon Eclipse E200 compound microscope, and an Olympus Camedia E-520 camera attached to an Olympus SZX16 stereomicroscope or to the eye piece of an Olympus BH2 transmission microscope. Digital images of different focal planes were stacked with Helicon Fo-cus™ 8.1.1. Illustrations of internal genitalia were made after digesting tissues off with Neo PanPur commercial pancreatic enzyme cocktail pill, clearing the structures in wintergreen oil (methyl-salicylate), then mounting them on a temperate slide preparation (Coddington 1983). Body measurements exclude the chelicerae and spinnerets. 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. Geographic coordinates of collection localities were obtained from the labels or georeferenced using Google Earth. Measurements and characters of the palp used in the diagnoses are based on the retrolateral view, unless otherwise indicated.
Although Dysderidae appears to be a monophyletic family often considered restricted to the Palaearctic, it is in fact distributed only in the West Palaearctic (from Canary Islands to west Xinjiang) and polyphyletic with its current generic composition. Eleven species of five genera are known from fossils (Dunlop et al. 2020): Dasumiana Wunderlich, 2004, Dysdera (1 sp.), Harpactea Bristowe, 1939 (5 spp.), Segistriites Straus, 1967 (1 sp.), and Mistura Petrunkevitch, 1971 (1 sp.). Judging by the position of the legs (i.e., legs I-III directed forwards) and the overall somatic features of Segistriites cromei Straus, 1967, this monotypic Neogene fossil genus is herein transferred to Segestriidae Simon, 1893. At the time of the description of Segistriites, Segestriidae was not a separate family but rather a subfamily (i.e., Segestriinae Simon, 1893) of Dysderidae. Furthermore, Strauss (1967) explicitly mentions the close affinity of this genus to Segestria Latreille, 1804, the type genus of Segestriidae. The monotypic Neogene fossil genus Mistura also appears to be misplaced in Dysderidae: the holotype specimen of Mistura perplexa Petrunkevitch, 1971 has an unknown arrangement of eyes and several characters different from Dysderidae, including a lack of claw tufts, the presence of an onychium, and long spinnerets (Petrunkevitch 1971).

Subfamily Dysderinae C.L. Koch, 1837
Diagnosis. This subfamily can be diagnosed from other dysderids by the edge of sternum-labium joint ca. 2.5-3× longer than the edge of the maxilla-sternum joint, all tarsi bearing claw tufts, posterior metatarsi bearing scopulae, and the spineless anterior tibiae and metatarsi. Furthermore, the bulb of dysderines does not bear a free embolus (with the exception of Harpactocrates Simon, 1914), and the posterior diverticulum of endogyne is large and wide (Deeleman-Reinhold and Deeleman 1988;Le Peru 2011;Kunt et al. 2019).
Diagnosis. Dysdera can be diagnosed from other dysderine genera by the interdistance of PLE and PME less than half of their diameter, three or four cheliceral teeth in one series, punctiform (= highly reduced) fovea, and femur I at least twice as long as coxa I. The bulb is cylindrical, bearing a broad posterior apophysis and a distal psembolus. The endogyne is composed by an anterior diverticulum bearing a dorsal arch and a ventral arch, a transverse receptacle and a posterior diverticulum bearing a transverse bar (Deeleman-Reinhold and Deeleman 1988; Le Peru 2011).
Comments. Deeleman-Reinhold and Deeleman (1988) proposed nine species groups within Dysdera: aculeata, asiatica, crocata, erythrina, festai, lata, longirostris, ninnii, and punctata. Charitonov (1956) and Fomichev and Marusik (2021) proposed an additional cylindrica group composed of Central Asian species considered within asiatica group by Deeleman-Reinhold and Deeleman (1988), mostly based on their disjunct distribution. Here, we tentatively treat these species within aculeata group, primarily on the basis of the conformation of male palp and considering that the discovery of similar species in Iran fills this distributional gap. Furthermore, characters based on spination used by Deeleman-Reinhold and Deeleman (1988) in definition of the species groups are not followed here as they appear to be variable; assignment of the species treated here to their respective groups is primarily based on the conformation of male palp.

aculeata species group
Diagnosis. This group can be diagnosed by a combination of the following characters: the carapace elongated and hexagonal, and the psembolus longer than the tegulum, with an anterior (= median) crest and an acuminate apex (Deeleman-Reinhold and Deeleman 1988). Comments. Currently, there is no clear distinction between the aculeata and asiatica groups, both of which are in serious need of a thorough revision (see Dimitrov 2021). Etymology. The specific epithet is a noun in apposition, referring to the apical ancestor of the Achaemenid dynasty of rulers of Persia.

Diagnosis.
The new species differs from its congeners occurring in the region by the very long receptacle (Re), longer than the posterior margin of the dorsal arch (Da) (vs. shorter).
Male. Unknown. Distribution. Known only from the type locality in Fars Province, southern Iran (Fig. 35). Etymology. The specific epithet is a noun in apposition, referring to an Iranian tribe primarily inhabiting Chaharmahal & Bakhtiari, Khuzestan, Lorestan, Bushehr, and Isfahan provinces.  Diagnosis. This species can be distinguished from other species of the aculeata group occurring in the region by having a wider psembolus (i.e., 1.5× wider than the tegulum).
Diagnosis. The new species differs from all Dysdera species occurring in the region by the receptacle divided into two chambers (vs. undivided), and the indistinct dorsal arch (vs. distinct).   Description. Female. Habitus as in Fig. 5A-C. Total length 8.11. Carapace 3.26 long, 2.56 wide. Eye diameters: AME 0.13, PME 0.13, PLE 0.14. Carapace, sternum, chelicerae, labium, and maxillae orangish. Legs pale orange. Abdomen pale creamcoloured, without any pattern. Spinnerets uniformly pale cream-coloured. Endogyne as in Fig. 6A, B; length/width ratio ca. 2.6; receptacle 2× longer than wide, divided in two chambers, with an anterior median concavity; anterior angles located almost at mid-part of each chamber and directed anteriorly, approximately as long as wide; dorsal arch indistinct; transverse bar ca. 4.6× longer than wide, mid-part similar to an inverted trapezoid, anterior part 1.3× longer than posterior part; lateral edges directed postero-laterally, clearly separated from transverse bar by an incision; posterior diverticulum elongated horizontally.
Male. Unknown. Comments. The species group (or even generic) assignment of this species is tentative pending the collection of the corresponding male.
Distribution. Known only from the type locality in Hormuz Island, the Persian Gulf (Fig. 35).  Etymology. The specific epithet is an adjective and refers to the country from where the specimens of the new species were collected.
Diagnosis. The male of the new species is somewhat similar to that of D. arabica Deeleman-Reinhold, 1988 from Oman (cf. Fig. 8A-D and Deeleman-Reinhold and Deeleman 1988: figs 309-310), but differs by the small, claw-like posterior apophysis (vs. broad and rounded) and keel-like median crest (vs. rounded). The male of D. iranica sp. nov. differs from those of its congeners occurring in Iran by the elongate keel-like median crest (vs. rounded or triangular). The female of D. iranica sp. nov. is most similar to that of D. tapuria sp. nov. by having a very wide receptacle (i.e., > 2× wider than the transverse bar), but differs by having a triangular extension (Te) in the anterior margin of the receptacle (vs. absent).
Distribution. Known from the listed localities in Fars and Hormozgan provinces, south-central and southern Iran (Fig. 35). Etymology. The specific epithet is an adjective, referring to the type locality of the species.

Dysdera isfahanica
Diagnosis. The male of this species differs from those of the other species of the aculeata group occurring in the region by the very long psembolus (i.e., length of psembolus/length of tegulum = 1.85 in the new species, vs. 1.6 or less in most other species), rounded arch-like ridge (Ar), presence of the notch of posterior apophysis (vs. absent), and median position of the posterior apophysis on the psembolus (vs. close to tegulum). Dysdera persica sp. nov. also bears a long psembolus (i.e., length of psembolus/length of tegulum = 2), but differs from D. isfahanica sp. nov. in the shape of the posterior apophysis. The female of this species can be recognized by its long anterolaterally stretched angles of the receptacle.
Comments. The material of this species and D. mazeruni sp. nov. (i.e., one male and two females in total) were reported by Roewer (1955); although he indicated that the females were collected in two different localities, they were found preserved in the same vial. The paratype female of D. isfahanica sp. nov. is matched with the holotype male due to their similar spination pattern and colouration.
Distribution. Known only from the type locality in Isfahan Province, central Iran (Fig. 35). Etymology. The specific epithet is a noun in apposition, named after an Iranian language of the northwestern branch spoken by the Mazandarani people.
Diagnosis. The new species is similar to D. isfahanica sp. nov., but differs by the arched anterior margin of receptacle (vs. almost straight), almost square-shaped dorsal arch (vs. distinctly trapezoidal), and shorter anterior angles (vs. longer, cf.    Fig. 13A Endogyne as in Fig. 14E-G; length/width ratio ca. 2.2; receptacle with slightly arched anterior margin, ca. 5× longer than wide, anterior angles slightly rounded; dorsal arch almost square-shaped, posterior margin 1.2× longer than anterior margin; transverse bar straight, 1.7× longer than receptacle; lateral edges broad, wider than receptacle; posterior diverticulum narrowing posteriorly.

Description. Female (Holotype). Habitus as in
Male. Unknown. Comments. As for the previous species. Distribution. Known only from the listed localities in Mazandaran Province, northern Iran (Fig. 35). Etymology. The specific epithet is an adjective, referring to the historical region of the Middle East, located in eastern Mesopotamia, which is now Iran.
Diagnosis. The male of this species differs from those of the other species of the aculeata group occurring in Iran by the extremely long bulb (especially psembolus, i.e., twice longer than tegulum), and by the tegulum with posterior margin 1.3× longer than anterior margin (vs. equal or shorter in length). The female of D. persica sp. nov. differs from those of its congeners by having the broadest dorsal arch, bearing almost angled anterior corners (vs. rounded).   Fig. 15D-F. Total length 9.06. Carapace 4.19 long, 3.20 wide. Eye diameters: AME 0.14, PME 0.16, PLE 0.17. Carapace, sternum, chelicerae, labium, and maxillae reddish brown. Legs orange. Abdomen greyish, without any pattern. Spinnerets uniformly dark yellowish. Measurements of  Palp as in Fig. 16A-D; bulb ca. 2.3× longer than wide; tegulum bell-shaped, almost as long as wide; psembolus 2× longer than tegulum; median crest triangular, ca. 2× shorter than length of psembolus, ca. 2.4× wider than high; posterior apophysis broad; incision between tegulum and psembolus present; retrolateral crest gradually rounded.

Description. Male (Holotype). Habitus as in
Female. Habitus as in Fig. 15A Endogyne as in Fig. 17A, B; length/width ratio ca. 1.9; receptacle 3.5× longer than wide, 1.2× wider than transverse bar; anterior angles rounded; dorsal arch trapezoidal; transverse bar straight, ca. 1.8× longer than receptacle; lateral edges with almost horizontal anterior margins, as long as width of receptacle; posterior diverticulum rectangular.
Distribution. Known only from listed localities in Golestan and Mazandaran provinces, northern Iran (Fig. 35). Comments. The Iranian record of this species is doubtful. Dunin (1985) described D. pococki on the basis of a male specimen from Turkmenistan, and the female of the species remains undescribed. Without providing any illustrations, Pocock (1889) reported a single female specimen from northeastern Iran which he tentatively identified as D. concinna L. Koch, 1878; this record was later attributed to D. pococki by Deeleman-Reinhold and Deeleman (1988), due to their close collection localities and without an examination of the Iranian material. This matter should be revisited once the female of D. pococki is described and the specimen reported by Pocock from Iran is studied and illustrated.

Dysdera sagartia
Diagnosis. The male of this species differs from those of the other species of the aculeata group occurring in Iran by the strong dorsal incision between tegulum and   psembolus, and the posterior apophysis bent on right angle (vs. no or small incision, and posterior apophysis not bent on right angle); the most similar species is D. mikhailovi Fomichev & Marusik, 2021 from Tajikistan, from which the new species differs by having a dorsal incision between tegulum and psembolus, the parallel dorsal sides of tegulum and psembolus (vs. dorsal margin of psembolus inclined), and smaller retrolateral crest angled at distal 1/3 of psembolus (vs. larger and angled at mid-part). The female of D. sagartia sp. nov. differs from those of its congeners occurring in Iran by having an arched anterior margin of receptacle in combination with a lack of anterior angles (vs. species with arched receptacle have anterior angles), and the almost semiround dorsal arch (vs. trapezoidal).
Distribution. Known only from the listed localities in Tehran Province, northern Iran (Fig. 35). Etymology. The specific epithet is a noun in apposition, referring to an Old Persian word for the Gorgan region, meaning "land of wolves".

Dysdera verkana
Diagnosis. The male of the new species is most similar to that of D. sagartia sp. nov., but differs by the more rounded median crest, the posterior apophysis not bent on right angle (cf. Fig. 22A and Fig. 19A : 1d,2pl;Ti: 4pl,2rl,5v;Mt: 3pl,2rl,6v. IV: Fe: 6d,1pl;Ti: 4pl,4rl,5v;Mt: 4pl,2rl,6v. Palp as in Fig. 22A-D; bulb ca. 2.2× longer than wide; tegulum bell-shaped, almost as long as wide; psembolus 1.66× longer than tegulum; median crest rounded, ca. 2.3× shorter than length of psembolus, ca. 2.7× wider than high; posterior apophysis broad; incision between tegulum and psembolus present; retrolateral crest roundly bent, forming right angle. Female. Unknown. Distribution. Known only from the type locality in Golestan Province, northern Iran (Fig. 35). Diagnosis. This group can be diagnosed by a combination of the following characters: the chelicerae straight or anteriorly convergent and longer than half of the length of the carapace, carapace broad and flat, and bulb with small or no lateral projection (Deeleman-Reinhold and Deeleman 1988). Etymology. The new species is named after Xerxes I, the fourth King of Kings of the Achaemenid Empire, ruling from 486 to 465 BC; adjective.
Diagnosis. The new species differs from all of its congeners occurring in the region by having a stylus (St), rounded median crest (Mc) and wide posterior apophysis (Ap); none of the other species has a rounded median crest, and those with a stylus, have a small posterior apophysis.
Female. Unknown. Distribution. Known only from the type locality in Bushehr Province, southern Iran (Fig. 35).

longirostris species group
Diagnosis. This group can be diagnosed by a combination of the following characters: cheliceral fang as long as the basal segment, carapace broad, flat and anteriorly convergent, and bulb with lateral projection smaller than the apex (Deeleman-Reinhold and Deeleman 1988). Etymology. The specific epithet is an adjective, referring to the type locality of the species.

Dysdera damavandica
Diagnosis. The male of the new species is most similar to that of D. concinna L. Koch, 1878 from Azerbaijan, but differs by longer bulb (i.e., bulb length/tegulum width =3.1, vs. 2.7), relatively shorter median crest, and longer stylus (cf. Fig. 26A and Dunin 1982: fig. B). Dysdera damavandica sp. nov. is also similar to D. tapuria sp. nov. but differs by the median crest higher than wide (vs. wider than high), relatively longer stylus (cf. Fig. 26A and Fig. 30A) and posterior apophysis located at distal half of the bulb (vs. located at mid-part).

Female. Unknown.
Distribution. Known only from the type locality in Mazandaran Province, northern Iran (Fig. 35). Etymology. The specific epithet is a noun in apposition, referring to an ancient Iranian people who inhabited an area known as Media between western and northern Iran.

Dysdera medes
Diagnosis. The male of the new species is similar to that of D. granulata Kulczyński, 1897 from Italy and the Balkan Peninsula, but differs by the shape of the tegulum (i.e., almost as wide as long, vs. 1.5× longer than wide), and by thinner psembolus (as wide as tegulum, vs. wider than tegulum). The male of D. medes sp. nov. differs from those of its congeners occurring in Iran by the very long median crest (i.e., longer than half of psembolus, vs. shorter), abrupt tip of psembolus in ventral and dorsal views (Fig. 28C, D) (vs. not abrupt), and posterior apophysis with two teeth (vs. one).
Female. Unknown. Distribution. Known only from the type locality in Tehran Province, northern Iran (Fig. 35). Etymology. The specific epithet is a noun in apposition, referring to the term applied to a mountainous region located in the Caspian coast of northern Iran.
Diagnosis. The male of the new species is most similar to that of D. concinna, but differs by longer bulb (i.e., bulb length/tegulum width = 3.1, vs. 2.7), median crest wider than high (vs. higher than wide), and shorter stylus (cf. Fig. 30B and  is also similar to that of D. damavandica sp. nov., but differs by the median crest wider than high (vs. higher than wide) and relatively shorter stylus (cf. Fig. 30A and Fig. 26A). The female of this species differs from those of its congeners occurring in the region by the very wide lateral edges of the receptacle (i.e., approximately half of the receptacle's width, vs. less than half ).

Female. Habitus as in
Distribution. Known only from the listed localities in Mazandaran Province, northern Iran (Fig. 35).

ninnii species group
Diagnosis. This group can be diagnosed by a combination of the following characters: chelicerae shorter than the width of carapace, carapace relatively short with anteriorly converging lateral margins, and bulb with simple crest, simple apex bearing a long subapical tooth, and a crescent-shaped lateral projection (Deeleman-Reinhold and Deeleman 1988). Etymology. The specific epithet is an adjective, referring to the type locality of the species.    Diagnosis. The male of the new species differs from those of its congeners by having weakly sclerotized bent stylus (St) (Fig. 32C, F) (vs. stylus, if present, not bent). The female of this species is very similar to that of D. iranica sp. nov. by having a wide receptacle (i.e., > 2× wider than transverse bar), but differs by the relatively wider receptacle lacking an anterior triangular projection, and the presence of a median concavity on the transverse bar (vs. receptacle with triangular projection, transverse bar without concavity).
Distribution. Known only from the type locality in Hormozgan Province, southern Iran (Fig. 35).

Discussion
Considering the results of this paper, there are 17 species of three genera of Dysderidae known from Iran. This number is considerably lower than what is known for the neighbouring Turkey (i.e., 69 species in seven genera; Danışman et al. 2022) and the Caucasus (i.e., 65 species in seven genera; Otto 2022). The material treated here, although comprising a relatively large number of new species, were collected in a few localities primarily in northern parts of the country (Fig. 35). Considering the small distribution ranges of most dysderids and the presence of several mountainous regions and biodiversity hotspots in Iran (e.g., Alborz and Zagros mountain ranges; Farashi and Shariati 2017), it can be assumed that any new material from this region could potentially comprise further undescribed species. It is possible that the true diversity of Dysderidae in Iran could range between 40 to 60 species, if not higher; interestingly, the widespread and cosmopolitan D. crocata has not yet been recorded from this country.
Furthermore, the diversity of Dysderidae in Central Asia is relatively low (i.e., 21 species of three genera; Mikhailov 2022). Most of the species (18) belong to Dysdera, and Dysderella and Harpactea are known only from western Turkmenistan (Dunin 1992;Zamani et al. 2017). The eastern boundary of the family is westernmost Xinjiang, and it appears that their diversity gradually decreases east of the Caucasus (Fomichev and Marusik 2021).