A survey of East Palaearctic Hersiliola Thorell , 1870 ( Araneae , Hersiliidae ) , with a description of three new genera

Th ree new genera and eight new species of Hersiliidae are described from the East Palaearctic (Afganistan, Iran, Kazakhstan, Kyrgyzstan, Pakistan, Tajikistan, Turkmenistan, and Uzbekistan). Th e genus Hersiliola Th orell, 1870 (Araneae: Hersiliidae) is revised, and four new species are described. Th e genus includes nine species: H. afghanica Roewer, 1960 (Afghanistan); H. esyunini sp. n. (Uzbekistan); H. foordi sp. n. (Iran); H. lindbergi sp. n. (Afghanistan); H. macullulata (Dufour, 1831) (type species; from Spain and Algeria to Israel and Yemen); H. simoni (O.P.-Cambridge, 1872) (from Spain and Morocco to Israel); H. sternbergsi sp. n. (Turkmenistan, Uzbekistan); H. versicolor (Blackwall, 1865) (Cape Verde); and H. xinjiangenis (Liang & Wang, 1989) (Xinjiang, China). A new genus Duninia gen. n. is described, with two new species, Duninia baehrae sp. n. (type species; Turkmenistan) and D. rheimsae sp. n. (Iran). A new genus Deltshevia gen. n. is described, with two new species, Deltshevia danovi sp. n. (type species; Turkmenistan, Kazakhstan) and D. gromovi sp. n. (Uzbekistan, Kazakhstan). Th e widely ranging Central Asian Hersiliola pallida Kroneberg, 1875 (Kazakhstan, Kyrgyzstan, Pakistan, Tajikistan, Turkmenistan, Uzbekistan) is transferred to a new monotypic genus, Ovtsharenkoia gen. n.


Introduction
Hersiliidae is a globally distributed entelegyne spider family that currently includes 159 species belonging to 12 genera (Platnick 2009;Marusik 2009).Most hersiliid species are found in tropical and subtropical regions.Ranges of only a few species extend north of 40°N.Hersiliids are easily recognizable due to their very long posterior lateral spinnerets.All species of the family are ecribellate.Hersiliidae, together with Oecobiidae, traditionally formed the superfamily Oecobioidea (Lehtinen 1967).Some araneologists also considered Eresidae as a family related to Oecobiidae and Hersiliidae, and placed these three families in Eresoidea (Coddington and Levi 1991).
During the last two decades, this family was subject to extensive studies, which resulted in exhaustive revisions of its Australian, Oriental, Neotropical, and Afrotropical faunas (Baehr and Baehr 1987, 1993, 1998;Rheims and Brescovit 2004;Rheims et al. 2004;Foord andDippenaar-Schoeman 2005, 2006;Foord 2008), as well as a revision of fossil taxa (Penney 2006).Less attention has been devoted to the small Palaearctic genus Hersiliola Th orell, 1870.
Th e published range of the genus in Asia is quite large, and extends southwards to the Karakoram Mts (northeastern Pakistan ;Caporiacco 1935;H. pallida) and eastwards to Xingiang (northwestern China;Marusik 2009;H. xinjiangenis).Hersiliola afghanica was reported as occurring widely across Afghanistan (Denis 1958, Roewer 1960).From Turkey, H. macullulata was reported (Hatay Province; Yağmur et al. 2008), but the family was not included in the recent checklist (Bayram et al. 2008).
Published records of Hersiliola from fi ve Central Asian republics of the former USSR (now the independent countries of Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan) have been also sporadic, although spiders of this genus are very common in deserts and dry mountains, found under stones, in rodent burrows (Krivokhatsky andFet 1982, Krivokhatsky 1987), and in pitfall traps (Kuznetsov and Fet 1984).Taxonomic placement of Central Asian species so far has been unclear; they were usually identifi ed tentatively as either H. macullulata (often misspelled as maculata; see below for discussion of correct spelling), H. pallida, or H. afghanica.No illustrations of male palps from Central Asian Hersiliola have been published.
Here, based on extensive new material, we reappraise all hersiliids found in Afghanistan, Kazakhstan, Kyrgyzstan, Tajikistan, Turkmenistan, and Uzbekistan, with a description of six new species and three new genera (Duninia gen.n., Deltshevia gen.n., and Ovtsharenkoia gen.n.).In addition, we describe two new species from Iran (the fi rst record of Hersiliidae for this country).

Material and methods
Th is paper is based on 238 specimens including 35 collected by the authors (28 specimens collected by VF during his resident work in Turkmenistan in 1975-1984, and 7 specimens collected by YM in Iran in 2000), as well as all available museum material (203 specimens, among them 31 collected by A.V. Gromov during a joint expedition with VF to Uzbekistan and Turkmenistan in 2002).Numerous new specimens from Central Asia reached us, due to the eff orts of Alexander Gromov and Dmitri Logunov, when this paper was already fi nalized; therefore a few descriptions are based on older material (Ovtsharenkoia gen.n.) or on a subadult male with a well developed palp (Deltshevia gromovi sp.n.).
Specimens were photographed using an Olympus SZX12 stereomicroscope and Olympus Camedia C-5050 camera in the Zoological Museum, University of Turku, Finland.Th e images were montaged using CombineZM image stacking software.
Whole specimens, palps and unmacerated epigynes were photographed in deep cups with a wax bottom.Depressions of diff erent shape and size were made in wax for fi xing specimens in the correct position.Epigynes were macerated using either KOH or lactic acid.In some cases, to make weakly sclerotized structures within epigynes more visible, we used Amido Black 10B (Amidoschwartz, naphthol blue black) amido acid.All measurements are in mm.
Terminology.Th ere are some contradictions in applying terminology for epigynal structures in Hersiliidae.Describing a hersiliid epigyne, Rheims and Brescovit (2004) and Foord and Dippenaar-Schoeman (2005) indicated both S (=spermatheca) and SR (=seminal receptacle).Th e same two terms were used in their character matrix.In fact, these two terms, derived from either Greek ("spermatheca") or Latin/English ("seminal receptacle"), refer to the same function: depository of sperm into the female epigyne.Ubick et al. (2005) treated these terms as synonyms.In our opinion, such confusion is caused by a specifi c structure of epigyne, which in most hersiliids contains accessory globular or digitiform structures near the terminal part of fertilization duct.We call these structures "accessory glands" (Ag), although their function is unknown.
Abbreviations: Morphological terms used in fi gures: Ag, accessory gland; Diagnosis.Hersiliola can be easily distinguished from other Palaearctic hersiliid genera by short spinnerets (shorter than abdomen length) and the shape of copulatory organs: discoid tegulum; long, whip-like coiled embolus; a small tegular apophysis perpendicular to the axis of the palp; insemination ducts coiled around fertilization ducts and uncoiled upper loop.Th e other three Central Asian genera of Hersiliidae (described here) have either a globular tegulum or have more than one tegular apophysis and uncoiled insemination ducts.From habitually similar African genera (Tama Simon, 1882 andTyrotama Foord &Dippenaar-Schoeman, 2005), Hersiliola can be distinguished by a digitate cymbium; presence of a hook-like median tegular apophysis; fl attened bulbus of the male palp [=discoid tegulum]; a fi liform, elongate, spirally coiled embolus; elongate, coiled copulatory ducts; small [relatively smaller] seminal receptacles (Foord and Dippenaar-Schoeman 2005); and some somatic characters.See Foord and Dippenaar-Schoeman (2005) for a detailed redescription of the genus and the type species.See also below for the studied material of the type species and comments on its distribution.
Comments.For a long time this genus, as well as the Afrotropical Tama, was diagnosed as a hersiliid with short spinnerets (shorter than abdomen length).All species with short spinnerets were placed in these two genera until Foord and Dippenaar-Schoeman (2005) recognized that species from southern Africa have copulatory organs and some somatic characters very diff erent from Hersiliola and Tama, and described a new genus Tyrotama.
Composition.Here, we assign nine species to Hersiliola, including four new species: H. afghanica Roewer, 1960    Distinguishing characters.Species of Hersiliola can be distinguished by the shape of the copulatory organs.Th e most important diagnostic characters in the male palp are as follows: (1) position of the embolic base; (2) number of embolic coils; (3) shape and position of tegular apophysis; and (4) size of cymbial terminal part.In females, the most important characters are: (1) shape of median plate; (2) number of coils of insemination duct; (3) shape and size of spermathecae; and (4) position of insemination duct attachment to the spermathecae.
Comments.Th is species was described from Spain, and later reported from Algeria, Israel and Burkina Faso.Its types appear to be lost (Levy 2003).It is very likely that that specimens from Burkina Faso (males) illustrated by Foord and Dippenaar-Schoeman (2005) are not conspecifi c with specimens from Algeria (females).Males from Burkina Faso have their embolus base in another position (at 04 hrs) compared to specimens from Spain or Israel (at 05 hrs).African males may belong to H. versicolor Blackwall, 1865, a species known from Cape Verde from females only.Another possibility is that specimen from Burkina Faso had a partially expanded bulbus, and its tegulum was slightly displaced (turned).
Note.Th is name has been commonly listed as "H.maculata" although Dufour's original name is Aranea macullulata.Th e name was misspelled as "H.macululata" by Simon (1885: 28).Karsch (1881: 7) 'corrected' the name to "maculata" which was widely used.Bonnet (1957Bonnet ( : 2180; footnote) suggested that macullulata is poor Latin, being derived from a non-existent word "macullula" ("a small spot").Th e consistent usage for over 100 years still does not validate Karsch's emendation (ICZN Article 32b-c).In fact, Dufour's original description (1831: 360) shows that his intention was to document a pattern of very small spots; and "macullula" is acceptable as an invented Latin word, parallel to the existing terms "medullula" or "ampullula" (H.D. Cameron, pers. comm. 1995).
Diagnosis.Hersiliola simoni can be recognized by the position of the embolic base (ca.10:30 hrs), a relatively short embolus forming only one coil, and the shape of epigyne.From H. sternbergsi sp.n., it diff ers by a fl atter palp with a smaller number of loops, a diff erent position of the embolic base, and a diff erent tegular apophysis.
Distribution.Hersiliola simoni was reported widely from Spain, Morocco, Niger, Cape Verde, Tunisia, Egypt, Libya, Lebanon, Syria, and Israel, although some records are doubtful or incorrect.Records of this species from Cape Verde should, in our opinion, refer to H. versicolor, the only species described and properly recorded from these islands (cf.Rheims et al. 2004).Th e record and illustrations of H. simoni from Yemen (Rheims et al. 2004) refer, in our opinion, to H. macullulata.Diagnosis.H. versicolor is similar to H. simoni, but diff ers from it by the shape of the median plate of the epigyne, which is much higher and has a wider septum.
Comments.Th is species is known from females only.Th e shape of the median plate illustrated by Foord and Dippenaar-Schoeman (2005: fi g. 4A) appears to be misinterpreted, judging from the fi gure of the vulva, and from the specimen (a slide of the epigyne) examined by us.Black dots on their fi g. 4A and on our Fig.7.1 seem to correspond to accessory glands.On the slide, glands seem to be displaced due to pressure from the cover glass.
Distribution.Known only from the Cape Verde Islands.Comments.Unlike other species from the Middle East (Israel), this specimen has only one coil of the insemination duct, and it can be expected that its male should have a correspondingly shorter embolus.It could belong to H. lucasi (O.P.-Cambridge, 1876), which was described from Egypt and is currently synonymous with H. simoni.Conformation of the epigyne is similar to those illustrated by Benoit (1974) from Tunisia and identifi ed as H. simoni.Benoit (1974) compared his specimen with those from Egypt.Roewer, 1960Figs 1.4, 7.3-4, 8.1 H. a. Roewer, 1960: 48, f. 16a-d (D♀).
Note.Roewer (1960) also listed 19 additional specimens ("paratypes") across a wide range in Afghanistan (Mazari-Sharif, Kabul, Jalalabad, etc.), but all of them were indicated as juveniles.However, in the SMF we found a single adult female, clearly designated as a paratype (allegedly of H. afghanica), with the following label: "1♀ Paratype, SMF 12996, No. 15, Afghanistan".Holotype (GNM) and paratype (SMF) labels are written by the same hand, presumably by Roewer.Study of this SMF female demonstrated that it belongs to another species, which is described below as H. lindbergi sp.n.
Diagnosis.Hersiliola afghanica is most similar to H. sternbergsi sp.n., from which it can be distinguished by larger spermathecae, shape of the upper loop, and insemination duct terminating on dorsal wall of the spermatheca.
Description.Male unknown.Female: Total length 5.5.Carapace 2.0 long, 2.15 wide, femur I 3.25 long, femur I/carapace length ratio 1.63.General colouration light brown.Carapace in poor condition, pattern is not visible.Abdomen with well developed pattern composed by a heart spot with dark margins; posterior part of abdomen with three transverse stripes.Legs with distinct annulations, light and dark rings equal in width.Coxae IV separated by less than one diameter.Chelicerae and maxillae with strong erect hairs.Epigyne as in Figs 7.3-4, 8.1.Septum as wide as lateral arms of median plate; windows distinct.Insemination duct relatively short; upper loop of coil located over the spermatheca; accessory gland digitiform.

Distribution. Afghanistan (Kabul Province).
Comments.Although this species was reported from a number of localities in Afghanistan, all records except for the holotype ♀ from Kabul are based on juveniles that could be misidentifi ed since other hersiliids (H.lindbergi sp.n. or Ovtsharenkoia pallida) could occur in this area.Any of these species could also have been reported by Denis (1958) from Pirzada (Afghanistan) as H. simoni.
All records of H. afghanica from Turkmenistan are misidentifi cations and refer either to Duninia baehrae gen.n. sp.n. or to Deltshevia danovi gen.n. sp.n.Etymology.Th e species name is a patronym honoring its collector, Dr. Knut Lindberg  of Lund, Sweden, who conducted an important expedition to Afghanistan between 1957 and 1960.

Hersiliola lindbergi
Note.Roewer (1960) in his description of H. afghanica indicated only one adult specimen, the holotype female.All paratypes were juveniles.Labels of the holotype of H. afghanica (GNM) and the SMF 12996 "paratype" were written by the same hand, probably by Roewer.
Description.Male unknown.Female: Total length 3.6.Carapace 1.5 long, 1.7 wide, femur I 2.5 long, femur I/carapace length ratio 1.66.General colouration light yellowbrown.Carapace without pattern but with dark margins.Abdomen with a wide heart spot and one transverse stripe; sides with brown bands.Legs without distinct annulations.Chelicerae and maxillae without strong setae.Coxae IV separated by less than one diameter.
Epigyne as in Figs 7.5-6, 8.3, with septum as wide as epigyne openings; median plate not separated from the epigynal plate; translucent spermathecae small and round, close to each other, their diameter approximately equals width of epigyne openings, windows absent; spermathecae separated from the opening by three diameters; insemination ducts long, making six coils around fertilization duct; the upper loop welldeveloped; accessory glands small, digitiform.
Diagnosis.H. lindbergi sp.n. clearly diff ers from other species by the very long insemination ducts making six coils and an apical loop.It can be also distinguished by small round spermathecae, the shape of the septum and the epigyne openings.
Comments.Th ere is a certain possibility that H. lindbergi sp.n. belongs to another genus.Unlike in H. maculullata, H. afganica, H. simoni, H. sternbergsi sp.n., H. versicolor, and Duninia baehrae sp.n., the median plate in this species is not separated from the epigynal plates and has no accessory sclerites or windows.Th e position of H. lindbergi sp.n. cannot be confi rmed until a male is found.
Distribution.Afghanistan, without a precise locality.Epigyne as in Figs 13-14, 8.5, with septum as wide as lateral arms of median plate; median plate not well separated from epigynal plate; translucent spermathecae oval, separated by their length (height) from the epigynal opening; insemination duct short, with only one loop; fertilization duct with a long appendix-like accessory gland extending along the duct.
Diagnosis.Hersiliola foordi sp.n. can be easily distinguished from all other congeners by its very short and wide insemination ducts, unseparated median plate, and lack of windows.
Comments.Th is species, like H. lindbergi sp.n., could belong to a separate genus.Its median plate is not separated from the rest of the epigynal plate.It also has no accessory sclerites and no windows.
Distribution.Th e species is known only from two nearby localities near Shiraz, southern Iran.Etymology.Th e species name is a patronym honoring our late colleague and friend Maris Šternbergs , of Riga, Latvia, the only Latvian araneologist of his generation, who also instructed VF in spider studies during his visit to Badghyz (Turkmenistan) in April 1977.

Hersiliola sternbergsi
Diagnosis.Hersiliola sternbergsi sp.n. is most similar to H. simoni, from which it can be distinguished by a longer terminal portion of the cymbium, thicker seminal ducts, lower position of the tegular ridge, and position of the base of the embolus (at almost 12 hrs).Females of H. sternbergsi sp.n. diff er by a wider septum, thicker lateral arms, and only one coil of the insemination duct around the fertilization duct.Th e epigyne of H. sternbergsi sp.n. is also similar to that of H. afghanica.Th ese two species have an almost identical septum shape, but in H. sternbergsi sp.n. the epigyne is smaller in size, has smaller spermathecae, and fertilization ducts terminating on the ventral wall of the spermatheca (on the dorsal wall in H. afghanica).
Female.Total length 5.0-5.2.Carapace 1.6-1.95long, 1.8-2.05(wider than long), femur I 2.5-2.85,femur I/carapace length ratio 1.46-1.56.Pattern distinct, composed by wide dark marginal bands, radial spots, transverse spot on median grove and V- Comments.Th is species has been collected widely in the lowland deserts of Central Asia (Turkmenistan and Uzbekistan), but constantly misidentifi ed (and misspelled) as "H.maculata" or H. pallida.Spassky (1952), who did not list "H.maculata", men- tioned a wide distribution of "H.pallida" in the Turanian Zoogeographical Province, thus obviously assuming this name for the population from Ashgabat identifi ed by Vlassov and Sytchevskaya (1937) as "H.maculata".Our study shows that this lowland desert Central Asian taxon is a new species, diff erent from H. macullulata (see above on H. macullulata).
Hersiliola sternbergsi sp.n. is widespread in lowland Turkmenistan on various types of desert substrates from clay and gypsum to sand.Krivokhatsky and Fet (1982) provide details on its phenology and ecology as a bothrophile in rodent burrows in the sand desert of East Karakum (Repetek Reserve), where it was most numerous in the entrances of burrows of the gerbil, Rhombomys opimus Licht.In Repetek, adult females of H. Etymology.Th e species name is a patronym honoring our friend and colleague Sergei Esyunin of Perm, Russia.
Diagnosis.H. esyunini sp.n. is closely related to H. xinjiangenis, from which it can be distinguished by its smaller size, diff erently shaped tegular apophysis and the median plate of the epigyne.From all congeners H. esyunini sp.n. can be easily distin- guished by a relatively shorter tip of the cymbium, undivided tegular apophysis and a median plate of the epigyne with turned up lateral edges of the median plate.Description.Male.Total length 3.4-3.65.Carapace 1.4-1.5 long, 1.5-1.6 wide, femur I 3.0-3.1 long, femur/carapace length ratio 2.1-2.14.Body yellow-light brown with pattern formed by brown hairs and spots.Carapace with brownish eye area and posterior cephalic part, and brown vertical stripe on clypeus (Figs 1.2, 2.2).Margins of carapace brown.Abdomen with indistinct pattern.Legs with broad, dark annulations, dark rings wider than light ones.Femur I almost entirely dark.Coxae IV separated by one diameter.Palp as in Figs 3.1, 4.4, 5.4, 6.6-7, with femur, patella+tibia and cymbium subequal in length, tibia in terminal part slightly wider than femur.Tegulum discoid, with long fi liform embolus and tegular apophysis.Embolus starts at about 01 hrs, makes a loop of more than 270° and terminates around 10 hrs.Tegular apophysis perpendicular to tegulum.Apical part of the bulb slightly fl attened (embolus straight, not rounded).
Comments.Earlier, Marusik (2009) confused this species with the related H. xinjiangensis due to the similarity of their male palps, i.e. same shape of the apical part of the tegulum (character unknown in other Hersiliola species), same position of the embolic tip, and same shape of the tegular apophysis.Misidentifi cation was made before we started this revision of East Palaearctic Hersiliola.
Distribution.Th e species is known only from two nearby localities in eastern Uzbekistan.
Diagnosis.Th is species can be easily distinguished from all congeners, except for H. esyunini sp.n., by a relatively shorter tip of the cymbium, fl attened apical part of the tegulum, and undivided tegular apophysis.From the sibling species H. esyunini sp.n., H. xinjiangensis can be distinguished by its larger size, diff erent shape of the tegular apophysis and median plate of the epigyne.
Description (translated from original description)."Female.Total length 5.8, body fl at.Cephalic part of carapace swollen, the position of PME highest.Cervical groove deep.Th orax low and fl at.Eye area brown, 8 eyes in 2 rows.Both AER, PER curved, AER curved stronger than PER.ALE white, others 6 eyes black, AME largest, located in the front of head; ALE small, located below in front of PLE. 4 eyes of PER near same size.Chelicera small, yellowish brown, with one promarginal tooth and no retromarginal.Pedipalps and legs yellowish brown, each segments of pedipalps and legs with blackish brown annuli in the median and the distal.Tarsus with 3 claws.Upper claw with a single tooth.Leg formula 2143.Opisthosoma dorsally with yellowish brown scales and grayish brown spots.Heart spot black, with three pairs of muscular depressions.Th e venter yellow, without marking.Anterior spinnerets robust, their distal segment small.Median spinnerets thin, nearly the same length as anterior ones.Rear spinnerets located on the sides of median ones.Th e distal segment same length as opisthosoma, smaller than the basal one.Colulus present.Epigyne brown, septum inverse T-shaped.
Male.Total length 5.00.Habitus, colour and markings style as in female.
Th is species inhabits crevices and holes of walls.It is a common species.Its colouration is cryptic and therefore it is not easy to fi nd specimens." Comments.Type specimens (2♂, 2♀) were supposedly deposited in the Department of Plant Protection, Xinjiang August 1 st Agricultural College, Urumqi, Xinjiang, China (Liang and Wang 1989).At our request, Shuqiang Li tried to fi nd these types but failed.It seems that after the retirement of Tie Liang these specimens were lost or transferred.Th e embolus base was not depicted by Liang and Wang (1989).Judging from the conformation of the epigyne in its sibling, H. esyunini sp.n., it is likely that median plate of the epigyne and vulva (endogyne) have been somewhat misinterpreted.
Distribution.H. xinjiangensis is known only from central Xinjiang (China).It is the northernmost species of the genus and of the entire family Hersiliidae.In Europe (Iberian Peninsula), the northernmost locality of Tama edwardsi (Lucas, 1846) (Ribera et al. 1988) is in northeastern Portugal ca.41°N, while the type locality of H. xinjiangensis lies north of 44°N.
Etymology.Th e genus name is a patronym honoring our late colleague, friend, and a prominent araneologist Pyotr Dunin  who lived and worked in Baku (Azerbaijan) and Togliatti (Russia).Gender: feminine.
Diagnosis.Duninia gen.n. can be easily distinguished from other genera of Hersiliidae by its globular tegulum, thick seminal duct, non-screw-shaped embolus that is thicker than the tegular apophysis, round spermathecae, and a heavily sclerotized copulatory opening below the epigynal plate.A globular tegulum is also present in the Central Asian genus Deltshevia gen.n., but this genus has a screw-shaped embolus and a thinner tegular apophysis.Females of Ovtsharenkoia pallida also seem to have copulatory openings below the epigynal plate, but they are not heavily sclerotized as in Duninia gen.n.; also, O. pallida does not have round spermathecae.
Description.Body length ca. 5 mm, carapace as long as wide from 1.75 to 2.25.Pattern as in Hersiliola.Palp cymbium with a long tip, tegulum globular, seminal duct wide, embolus wide and not screw-shaped, tegular apophysis claw-like, with its claw part thinner than embolus.Epigynal plate with a septum-like structure, but without windows and openings, median plate without distinct margins.Copulatory opening is located below the epigynal plate in an epigastral fold; spermathecae round, coiled ducts absent.
Comments.Conformation of the epigyne in this genus is, in some respects, unique among entelegyne spiders.Th e epigynal plate has no openings (furrows or fovea) but possesses two pairs of fovea on the vertical posterior wall located inside the epigastral fold.Th e lateral pair of fovea is shallow and elongate.Another pair of fovea is closer to the median part and located deeper; they have a more heavily sclerotized wall and are very deep.It seems that these deep pockets have copulatory openings inside the anterior part.Lateral pockets seem to match the tegular apophysis of the male.
Th ere are few taxa among entelegyne spiders that have a copulatory opening on the posterior vertical wall of the epigyne located in the epigastral fold.Besides some Erigoninae, we know of only one such genus, Paratus (Liocranidae) (cf.Marusik et al. 2008).Paratus has epigynal plate without furrows and fovea, and a small fovea hidden in the epigastral wall that leads to two closely separated copulatory openings.
Th e male palp of Duninia gen.n. is unique among Hersiliidae because of its short, broad embolus and tegular apophysis extended along the cymbial axis.
Diagnosis.From its sibling D. rheimsae sp.n., females can be easily distinguished by a larger body and epigyne, diverging spermathecae, and deep pockets longer than the lateral ones.
Description.Male (holotype from Kurkulab, abdomen and most of legs are missing): Carapace 2.12 long, 2.25 wide, femur I 5.0 long, leg I/carapace length ratio 2.36.Palp as in Figs 3.3,4.5,5.5,6.8,bulbus globular, basal portion of seminal duct thick; embolus very massive, fl at and short, twice shorter than the apical portion of the cymbium, opening large and clearly visible, tegular apophysis claw-like, large, located on the apical part of tegulum, extends parallel to cymbial axis, its terminal part much thinner than embolus.
Female: epigyne as in Figs 8.6, 11.1-4, lacks windows and distinct openings; median plate twice as wide as high; translucent spermathecae round, separated from median plate by more than one diameter; lateral sides of epigynal plates with "arches" clearly visible on dissected epigyne; vulva with two round spermathecae separated by one diameter and large lateral wings.Lateral arches lead to small pockets, and large wings correspond to large and deep pocket that seems to correspond to copulatory opening.It is not clear which structures of the vulva correspond to fertilization ducts, and also not clear whether the fertilization duct has accessory glands.
Distribution.Turkmenistan (Bolshoi Balkhan, Central Kopetdagh, Badghyz).Etymology.Th e species name is a patronym honoring our colleague Cristina A. Rheims (São Paulo, Brazil), for her contributions to in-depth modern studies of Hersiliidae.
Diagnosis.D. rheimsae sp.n. can be distinguished from the sibling species D. baehrae sp.n. by its smaller body and epigyne size, converging spermathecae, and less slanting epigynal pockets bearing large, clearly visible hemispheres.
Comments.Judging from the conformation of the male palp and epigyne in its sibling D. baehrae sp.n. and conformation of the epigyne in D. rheimsae sp.n., it is reasonable to suggest that male of this species should have a smaller embolus (matching a small, deep pocket of D. rheimsae sp.n.) and a larger tegular apophysis.
Distribution.Known only from type locality in northern Iran.
Diagnosis.Deltshevia gen.n. diff ers from other genera of Hersiliidae by a globular tegulum; thick and screw-shaped embolus with a tapering, sharp tip; large two-armed tegular apophysis extended upward; large epigyne openings; and thick copulatory ducts.
Comments.A globular tegulum and tegular apophysis extended upward are also found in Duninia gen.n.In the latter genus, however, the tegular apophysis has one clawlike arm; this arm is thinner than the embolus and is extended upward (along the cymbial axis).In Deltshevia gen.n., the tegular apophysis has two arms: an upper arm directed upward, and a claw-like arm perpendicular to the cymbial axis.Females of this genus can be easily recognized by the presence of a (real) septum, large copulatory openings, and a wide (wider than spermathecae) and uncoiled insemination duct.Septum and small copulatory openings are present also in Hersiliola, but insemination ducts in this genus are always coiled and thinner than the spermathecae, or subequal in width to the spermathecae.Description.Small hersiliids 5-7 mm long, with carapace 2.1-2.7 long and wide.Palp globular; embolus thick and screw-shaped; tegular apophysis large, with two arms: a massive vertical arm and a small horizontal arm.Epigyne with large copulatory openings; wide insemination ducts; septum and median plate distinct; spermathecae oval; fertilization duct short; accessory gland globular.
Diagnosis.Deltshevia danovi sp.n. is similar to D. gromovi sp.n., from which it can be easily distinguished by the shape of the embolus, tegular apophysis and epigyne.Deltshevia gromovi sp.n has a shorter and screw-shaped embolus, while D. danovi sp.n. has a longer, non-screw-shaped embolus.Th e apical portion of the tegular apophysis in D. gromovi sp.n. has subparallel margins, while the apical portion of the tegular apophysis in D. danovi sp.n. is triangular, with slanting margins.Th e epigyne of D.  Female.Total length 6.0-7.0.Carapace 2.1-2.3 long, 2.4-2.5 wide (wider than long).Carapace pattern and eye arrangement as in Figs 9.4-5.Abdominal pattern as in Fig. 9.3, dorsal side with a dark heart band and four transverse stripes, sides of abdomen with dark spots, venter without pattern.Epigynal plate 1.00-1.14wide, fovea 0.47-0.57wide, plate/fovea ratio 2-2.1.Epigyne as in Figs 10.1-3, with distinct windows, well-separated septum, median plate anchor-like, epigynal opening distinct (with well-expressed borders), opening diameter larger than septum width and equal to median plate height; translucent spermathecae elongate and located aside of openings.Vulva simple, with large pockets (continuation of epigynal opening), with oval spermathecae separated by more than three times their widths; insemination duct short, accessory gland globular, poorly visible.
Deltshevia gromovi sp.n. urn:lsid:zoobank.org:act:D7FA30A6-E286-4A5A-9C33-D799D815E437Figs 3.5,4.7,5.7,Hersiliola macullulata: Zyuzin et al. 1994: 6 (Kazakhstan).Misidentifi cation.Note: Much new material from Central Asia reached us, due to the eff orts of Alexander Gromov and Dmitri Logunov, when this paper was already fi nalized; therefore the following description and fi gures of a male are based on a subadult with welldeveloped palp.Etymology.Th e genus name is a patronym honoring our friend and colleague, and a prominent araneologist, Vladimir Ovtsharenko of New York, USA.Gender: feminine.
Diagnosis.Ovtsharenkoia gen.n. can be easily distinguished from other genera of Hersiliidae by its short spinnerets and the shape of the copulatory organs.Th e male palp has a unique conformation for the family due to the presence of a complex outgrowth in the basal part of the tegulum.All other genera similar to Hersiliola have only one apophysis (tegular).Females of Ovtsharenkoia gen.n. can be recognized by a small median plate of the epigyne, a transverse translucent fertilization duct, large pale areas next to the median plate, and absence of distinct spermathecae.
Description.Same as for the type species.Diagnosis.Same as for the genus.Description.Male (described here for the fi rst time).Total length 4.8.Carapace 2.0 long, 2.25 wide, femur I 4.25 long, femur/carapace length ratio 2.13.Carapace light brown with dark margins, radial dark stripes and median dark band; cephalic part separated from the thoracic part by a dark V-shaped spot.Abdomen light brown with a brown heart spot, transverse stripes and dark sides.Dorsal pattern variable.Venter of abdomen without pattern.Palp as in Figs 3.6, 4.8, 5.8, 6.9, femur and cymbium equal in length, patella+tibia almost as long as cymbium or femur; tegulum round, basal part extended; embolus whip-like, arched; embolus makes a half loop, base of embolus located on retrolateral side; tegulum with two apophyses: apical (?tegular) and basal.Basal apophysis (outgrowth) complicated, subdivided into three parts: lamellate, digitiform retrolateral, and mesal.Embolus passes below mesal part of basal apophysis.Apical apophysis perpendicular to the cymbial axis.
Female.Total length 4.2-6.0.Carapace 2.0-2.5 long, 2.25-2.75wide, femur I 3.5, femur/carapace length ratio 1.75.Colouration as in male.Epigyne as in Figs 12.1-5, with strongly chitinized small median plate, septum well-developed, its length subequal to length of lateral arms; width of septum variable; windows absent, lateral sides of epigynal plate with small extensions, indicating pockets; aside of median plate epigyne is pale, size of pale part variable; upper part of epigynal plate with a pair of transverse stripes formed by translucent insemination (?) ducts.Distinct spermathecae absent, insemination ducts relatively thin, coils absent; fertilization ducts with globular accessory glands.Insemination ducts have small globular extensions that possibly correspond to spermathecae proper.
Size of epigyne variable, although size of median plate is not variable, as well as position of translucent fertilization ducts.Distance between epigastral fold and ducts is the same in large and small epigynes.
Comments.Details of epigynal structure remain uncertain.It is not clear whether openings on the median plates are real copulatory openings or just fovea, origins of copulatory (insemination) ducts as well as the position of the terminal part of the fertilization ducts.Most probably the insemination duct is weakly sclerotized and originates in pockets lying below the epigynal plate.
We also examined a juvenile female reported as Hersiliola pallida by Caporiac co (1935) from Karakoram Mts.(Pakistan) which tentatively belongs to this species.Th is specimen originates from a single high mountain locality, Chongo in Braldu Valley, Baltistan (in the vicinity of the Baltoro Glacier); see Spoleto (1930) for the 1929 Italian expedition route.Although Caporiacco (1935) reported three adult males, these specimens could not be located in MZUF (L.Bartolozii, pers. comm.).
Earlier records from lowland Turkmenistan were misidentifi cations and refer to Hersiliola sternbergsi sp.n. (Repetek, Badghyz) or Duninia baehrae sp.n. (Badghyz).However, new material collected by S. L. Zonstein in 1993 shows that this species is indeed found in Turkmenistan.It was collected in the very southwest of Badghyz, in the mountainous area bordering Iran (Zyulfagar Range in the watershed of Tejen, which forms an important zoogeographic boundary between the Turkmeno-Khurassan mountains to the west and all Central Asian/Himalayan mountains to the east; see Fet 1994).
Distribution.Kazakhstan, Kyrgyzstan, Pakistan (Baltistan: Karakoram Mts.), Tajikistan, Turkmenistan (south), Uzbekistan.Unlike most species of Hersililola and other habitually similar genera, this species is restricted exclusively to mountainous areas and seems to be the only species of hersiliids in the Palaearctic that penetrates human settlements and buildings (Bishkek).It could be a petrophilous species that inhabits cliff s and walls.
We have no doubt that Oecobiidae are related to Hersiliidae.In addition to clear somatic characters indicated by earlier authors such as eye pattern, an almost round carapace (width equals length), etc., both families have elongated posterior lateral spinnerets and the same way of holding the palps (converging femora, and diverging tibia and terminal joint) in both sexes (Fig. 1.2, 2.1; fi g. 71a in Jocqué and Dippenaar-Schoeman 2006).Similarities between Oecobiidae and Hersiliidae even led Strand (1913) to confusion when he described Oecobius (s.l.) brachyplura (Strand, 1913) in Hersiliola (see Fet 2008).Coddington and Levi (1991) placed Oecobiidae and Hersiliidae in Eresoidea.Although Eresidae, like Oecobiidae and Hersiliidae, are very diff erent from the rest of the entelegyne spiders, we doubt that they should be placed in the same superfamily due to entirely diff erent somatic morphology, morphology of copulatory organs, ecology, and behavior.
One of the additional diagnostic features of Hersiliidae, not indicated by earlier authors, is the presence of "accessory glands" in the epigyne.Th ese structures appear to be present in the majority of hersiliid genera such as Hersilia, Hersiliola, Deltshevia gen.n., Duninia gen.n., Yabisi Rheims & Brescovit, 2004, most of Tyrotama Foord & Dippenaar-Schoeman, 2005, Murricia Simon, 1882, Neotama Baehr & Baehr, 1993, and Tamopsis Baehr & Baehr, 1987.Some Tamopsis species appear to have two pairs of sack-like structures in addition to spermathecae (cf.fi gs 25e, 28e in Baehr and Baehr 1987).Neotropical Iviraiva Rheims & Brescovit 2004 has acinoform spermathecae (not known in other entelegynes), and it is not clear whether they have accessory glands.Interestingly, structures that appear to be homologous to "accessory glands" have been observed in the related family, namely Uroctea limbata (C.L. Koch, 1843) and two unidentifi ed species of Uroctea (Oecobiidae) (cf.fi gs 10-16 in Baum 1972).Baum (1972) called these structures a "blind endender Anhang".Th ey were found in only half the studied species.Unlike in Hersiliidae, "acessory glands" in Uroctea are placed close to the spermathecae, but not close to the terminal part of the fertilization duct.
Structures similar to the "accessory glands" of Hersiliidae are common among entelegyne spiders.Most Hadrotarsinae (Th eridiidae) have one or even two additional pairs of "spermathecae" (function unknown).Some Hahniidae have a second pair of globular structures called secondary receptacula (spermathecae) (cf.Harm 1966).Primary and secondary receptacula (spermathecae) in most of hahniid species are of the same size.Th e agelenid genus Azerithonica Guseinov, Marusik & Koponen, 2006 has a globular accessory gland at the midpoint of the fertilization duct (Guseinov et al. 2006, fi g. 33).Struc-tures similar to accessory glands are present in other agelenid genera too, such as Agelescape and Malthonica.In all above mentioned cases in other families, accessory glands are associated with insemination ducts, while in Hersiliidae and some Oecobiidae they are connected (if present) with fertilization ducts.In Hersiliidae, accessory glands are much smaller than spermathecae (receptacula), with the exception of Deltshevia gromovi sp.n.where accessory glands are only 2 to 2.5 times smaller than the spermathecae.
It is also worth mentioning that, unlike in all other entelegynes, fertilization ducts in Hersiliidae are long and heavily sclerotized (often more heavily than insemination ducts).Relatively long fertilization ducts are known in Oecobiidae, but they are weakly sclerotized.
Judging from the description provided by Baehr and Baehr (1993) for the Oriental genus Hersilia, this genus could be split into several genera.For example, the male of H. kinabaluensis Baehr & Baehr, 1993 has a bulbous tegulum, a screw-shaped embolus (fi liform or stick-like in other species) directed parallel to the cymbium, and lacks a tegular apophysis (which is present in all other species of the genus).Some females of Hersilia have one and some, two pairs of "accessory glands".At the same time, some species of Hersilia lack distinct spermatheca (like Ovtcharenkoia pallida).
Of these, two species, H. foordi sp.n. and H. lindbergi sp.n., known only from females, could in fact belong to other genera.It is very likely that actual diversity of Hersiliola is higher, and it seems that some junior synonyms could be revalidated.Hersiliola pallida Kroneberg, 1875 is transferred to a new genus, Ovtsharenkoia gen.n. (see below).
Th e species name is a patronym honouring our colleague Stefan H. Foord (Th ohoyandou, South Africa), for his contributions to in-depth modern studies of Hersiliidae.Description.Male unknown.Female: Total length 5.01-5.75.Carapace 2.1-2.25 long, 2.2-2.45wide, femur 3.1-3.75long, femur I/carapace length ratio 1.47-1.67.Coxae IV separated only slightly.Abdominal pattern such as in holotype of H. afghanica formed by short fl at hairs of orange-light brown, dark brown, and white colour.