On Araniella and Neoscona (Araneae, Araneidae) of the Caucasus, Middle East and Central Asia

Abstract New taxonomic data for species belonging to Araniella Chamberlin & Ivie, 1942 and Neoscona Simon, 1864 occurring in the Caucasus, Middle East and Central Asia are provided. Three species are described as new to science: A. mithrasp. nov. (♂♀, northwestern, central and southwestern Iran), A. villaniisp. nov. (♂♀, southwestern Iran, eastern Kazakhstan and northern India) and N. isatissp. nov. (♂♀, central Iran). Neoscona spasskyi (Brignoli, 1983) comb. nov., stat. res. is removed from the synonymy of N. tedgenica (Bakhvalov, 1978), redescribed and recorded from Iran and Turkmenistan for the first time. New combinations are established for this species, as well as for Araniella nigromaculata (Schenkel, 1963) comb. nov. (♀, north-central China) (both ex. Araneus). Two new synonymies are proposed: Araniella tbilisiensis Mcheidze, 1997 syn. nov. is synonymized with A. opisthographa (Kulczyński, 1905), and Neoscona sodom Levy, 1998 syn. nov. is synonymized with N. theisi (Walckenaer, 1841); the latter is recorded from Iran, Georgia, and Russia (Northern Caucasus) for the first time.


Introduction
Araneidae Clerck, 1757 with 3072 valid species (WSC 2019) is the third largest family of spiders. At least in the Palaearctic, it is the best-studied family of spiders due to numerous publications dealing with the survey of regional fauna, or revisions of Euro-pean and Far East (China, Japan, Korea) species. However, the Central Palaearctic is not well studied in comparison to other parts. Several species described by Bakhvalov (1970Bakhvalov ( , 1974Bakhvalov ( , 1978Bakhvalov ( , 1981 remain known only from the original publications supplied with very schematic figures and brief descriptions. In order to fill this gap, we decided to study all available material from Iran and Central Asian countries and provide step by step reviews of different genera. Among material examined, we recognized two new species of Araniella Chamberlin & Ivie, 1942 and one new species of Neoscona Simon, 1864. While comparing new species with species occurring in the region, we recognized two new synonyms and two new combinations in both genera. The goals of this paper are to provide illustrated descriptions of new species and redescriptions of poorly known species, along with new combinations, synonymies, and distribution records.

Materials 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 JEOL JSM-5200 scanning electron microscope (SEM) at the Zoological Museum of University of Turku, Finland. 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 are provided for leg I only (IV, if missing) and listed as: total length (femur, patella, tibia, metatarsus, tarsus). All measurements are given in millimeters.
Abbreviations not explained in the text: ALE -anterior lateral eye, AME -anterior median eye, PLE -posterior lateral eye, PME -posterior median eye.
Comments. Currently, this genus includes 12 species distributed exclusively in the Holarctic (WSC 2019). Only two species, the generotype and A. proxima (Kulczyński, 1885), are known in both parts of the realm (Palaearctic and Nearctic); all other spe-cies are restricted to the Palaearctic. Although the genus has never been the subject of a global revision, it is well studied, and all species are known by both sexes, with the exception of A. tbilisiensis (Mcheidze, 1997). This species was described on the basis of both sexes, but the male palp has never been illustrated.
Diagnosis. The genus well differs from all Holarctic genera of Araneidae by large (as long as embolus and terminal apophysis), claw-or spine-like median apophysis directed mesally (vs. not claw-or spine-like but having at least 2 arms). Etymology. The specific epithet is a noun in apposition, and refers to Mithra, the god of light in ancient Indo-Iranian mythology.

Araniella mithra
Diagnosis. Male palp and epigyne resemble those of A. opisthographa, but the two species can be differentiated by the following characters: 1) the embolus is slimmer in A. mithra sp. nov., vs. triangular-shaped and with a wider base in A. opisthographa; 2) the terminal apophysis in A. mithra sp. nov. is almost as wide over its entire length, vs. wider near the peak in A. opisthographa; 3) the conductor in A. mithra sp. nov. has three distinct spikes, vs. one spike and one more rounded process in A. opisthographa; 4) the tegulum in A. mithra sp. nov. is higher with a short pointed tip, vs. the slender tegulum with a longer tip in A. opisthographa; 5) male carapace unicolor in A. mithra sp. nov., vs. presence of broad dark marginal bands in A. opisthographa; 6) epigyne with slightly longer scape, and the sclerotized bulges are rounded around the base of scape in A. mithra sp. nov., vs. more incised triangular bulges in A. opisthographa.
Palp as in Figs 2A, B; 4A, B; 6A, B. Tegulum with low round ridge and terminally with short pointed tip; terminal apophysis with blunt end and almost equally wide along its length; embolus pointed, sickle-shaped bent; median apophysis sickleshaped bent upwards, covered by small denticles (less visible via stereomicroscope), with pointed tip ended near base of embolus; conductor with three distinct spikes.
Phenology. Adult males and females were collected in mid and late June. Distribution. Known only from the type localities in northwestern, central and southwestern Iran. It is possible that some of the previous Iranian records of A. opisthographa refer to this species. Etymology. This species is named after French mathematician Cédric Villani (born 5.10.1973), winner of the Fields Medal in 2010 and the former director of Sorbonne University's Henri Poincaré Institute, for his "mysterious love" for spiders.
Diagnosis. Male palp and epigyne resemble those of A. proxima and A. opisthographa. Both species, compared to A. villanii sp. nov., have similar shape of embolus, and ter- minal apophysis is identical to that of A. proxima and conductor is identical to that of A. opisthographa. However, the new species can be diagnosed by the following characteristics: 1) the tegulum in A. villanii sp. nov. is markedly shorter, higher, protruding and rounded, vs. more compact non-protruding tegulum with distinctly higher ridge in A. proxima, and slender with pointed tip in A. opisthographa; 2) the terminal apophysis in A. villanii sp. nov. is almost as wide throughout its length, vs. wider at the tip in A. opisthographa; 3) the conductor in A. villanii sp. nov. has one spike and one more rounded process connected to each other, vs. two independent spikes in A. proxima; 4) the median apophysis in A. villanii sp. nov. is longer in comparison to both mentioned species; 5) epigyne of A. villanii sp. nov. has a distinctly broader scape, vs. slender in A. proxima and A. opisthographa; 6) the median plate is narrower and more rectangular in the new species, vs. wider and rounded plate in A. opisthographa and triangular plate in A. proxima; 7) receptacles and entrance ducts in A. villanii sp. nov. do not touch Description (colors and pattern seem faded). Male (holotype). Habitus as in Fig. 1B. Total length 4.37. Carapace 1.91 long, 1.69 wide in pars thoracica, 0.76 in pars cephalica. Eye sizes and interdistances: AME: 0.08, ALE: 0.07, PME: 0.09, PLE: 0.09, AME-AME: 0.12, PME-PME: 0.11. Carapace, sternum, labium, chelicerae, and maxillae reddish brown, lighter ventrally, carapace with two broad dark marginal bands. Legs lighter in color than the carapace, distally with dark broad annulations. Abdomen pale (stored in alcohol, most probably green in live specimens) dorsally, dark gray ventrally, posterodorsally with three pairs of black lateral spots. Spinnerets light brown, apical segment lighter. Leg I measurements: 6.43 (1.97, 0.82, 1.50, 1.46, 0.68). Palp as in Figs 3A, B; 4C, D; 5A, B. Tegulum terminally blunt with round ridge; terminal apophysis with blunt end and almost equally wide along its length; embolus triangular-shaped, with wider base; median apophysis sickle-shaped bent upwards with pointed tip ending near base of embolus and covered by many small denticles; conductor with one distinct spike and one more rounded process.
Epigyne as in Figs 7A, 8A, 9A, 10A. Scape wider in the middle, extending beyond epigynal plate. Copulatory ducts not clearly visible through epigyne cuticle. Oval receptacles are about half their diameter apart; entrance ducts a similar distance apart. Median plate (posterior view), between lateral sclerotized copulatory bulges, slender, slightly wider in the middle. Comments. Araneus tbilisiensis was described based on one male and four females from the environs of Tbilisi, Georgia. There is no indication which specimen/sex was selected as the holotype. Mcheidze (1997) provided figures of male and female habitus, as well as epigyne, but the male palp was not illustrated. Judging from the figure of epigyne and distribution, it is most likely a junior synonym of A. opisthographa, which is already known from the surroundings of Tbilisi (Otto 2019). We tried to obtain the type material for this study, but we have been informed that the single male specimen is most probably lost (V. Pkhakadze, pers. comm.). (Schenkel, 1963), comb. nov.
Comments. The female holotype was collected in southern Gansu (ca. 33°40'N, 104°20'E), north-central China. Figures of Yin et al. (1997) and Song et al. (1999) are reproduced after Schenkel (1963). The holotype (in Muséum National d'Histoire Naturelle, Paris) was examined in 1980 by Yuri Marusik and illustrated, but no data have been copied from the label. Abdominal pattern and shape of epigyne indicates its belonging to Araniella and therefore we provide a new combination.
Diagnosis. The new species is similar to N. theisi and N. spasskyi in having a white median band on sternum (Fig. 13A, C, E), but well differs by having a broad white median band on the venter of abdomen (vs. venter with lateral white band, and dark median band). Males of N. isatis sp. nov. can be easily distinguished from the species occurring in the region by numerous small spines on tibia II (Fig. 14F) lacking in other species (Fig. 14D-E) and median apophysis lacking prolateral extension (Me) (vs. present). Epigyne of this species well differs from other species occurring in Central Asia by having prominent lateral extensions (Le) as long as wide and long scape (Sc) almost 2 times longer than wide (vs. lateral extensions absent or poorly developed and scape almost as wide as long, cf. Fig. 17A, D, G).
Epigyne as in Figs 13F, 17G-I. Long, with scape (Sc) as long as base; lateral extensions (Le) prominent, as long as wide, originates dorsally; scape almost twice longer than wide.
Distribution. Known only from the type locality in Yazd Province, central Iran.
Neoscona spasskyi (Brignoli, 1983), comb. nov., stat. res. Diagnosis. Neoscona spasskyi differs from the similar N. theisi by having a thinner dark median band on the carapace and wider white lateral bands (cf. Figs 11C and 11A, B, D, E). Some specimens of this species have a pyramid-type pattern (Fig. 12A, B) lacking in other species. Males of this species differ from the congeners known in the region by having about 40 ventral spines on tibia II (vs. ca. 90, 20 or 10). Neoscona spasskyi differs from N. isatis sp. nov. by having prolateral extension of median apophysis. Epigyne of this species has the scape almost as wide as long vs. about twice longer than wide in N. isatis sp. nov. It differs from those in N. theisi by having distinct constriction (vs. lacking).
Epigyne as in Figs 13D, 17D-F. Epigyne with distinct constriction; lateral extensions distinct, wider than long; scape almost as wide as long.
Comments. Types of this species have not been found among the Spassky's collection in the Zoological Museum, St. Petersburg (Nekhaeva, pers. comm.). Spassky (1952) described this species as Araneus cruciferoides, a name preoccupied by Tullgren (1910) on the basis of both sexes. Later, a replacement name, Araneus spasskyi, was provided by Brignoli (1983). Marusik et al. (1991) erroneously synonymized it with Neoscona tedgenica (Bakhvalov, 1978), a species known only from a female and a juvenile specimen collected in Turkmenistan (Bakhvalov 1978), and transferred to Aculepeira by Brignoli (1983). Comparing available figures in Spassky (1952) and Bakhvalov (1978) and the newly studied material, these two species differ in the shape of the posterior scape (rounded vs. triangulate) and the dorsal abdominal pattern (white "true" folium on a dark background in N. tedgenica, vs. dark "incomplete" folium on a light background in the other species). For these reasons, we now revalidate the name 'spasskyi' and establish a new combination for it: Neoscona spasskyi (Brignoli, 1972) comb. nov.
Distribution. Tajikistan, Kyrgyzstan (Spassky 1952), Turkmenistan, Iran (first records for both). Diagnosis. This species well differs from other species occurring in the region by the abdomen being as wide as long in the female and with small horns in the male (vs. abdomen longer than wide and lacking horns). Description. See above-cited literature. Distribution. Entire Africa, Mediterranean (Grasshoff 1986) to Turkmenistan (Mikhailov 2013). Fig. 18 Araneus tedgenicus Bakhvalov, 1978: 790, figs 1-4 (♀). Aculepeira tedgenica : Brignoli 1983: 255. Diagnosis. Neoscona tedgenica differs from the closely similar N. spasskyi in the shape of the posterior area of the scape (triangulate vs. rounded) and the dorsal abdominal pattern (white "true" folium on a dark background in N. tedgenica, vs. dark "incomplete" folium on a light background in N. spasskyi).

Neoscona tedgenica (Bakhvalov, 1978)
Comments. See under Neoscona spasskyi (Brignoli, 1983). Types of this species are lost along with the rest of the private collection of Bakhvalov.
Distribution. Turkmenistan (Bakhvalov 1978).  Diagnosis. Neoscona theisi differs from the congeners occurring in the region by the presence of a wide black median band on the venter of abdomen and thin white lateral stripes (Fig. 12D). Males of this species have tibia II with fewer ventral spines (ca. 20) than N. spasskyi (ca. 40) and N. isatis sp. nov. (ca. 90) and more than in N. adianta (ca. 10). Males of N. theisi can be recognized also by the palp with pointed dorsal extension/projection of the tibia (Fig. 14A) (vs. absent), distinct ventral conical projection of the tegulum (Te) lacking in other species, broad and well sclerotized lamella and wide conductor (vs. lamella thin and weakly sclerotized, conductor clublike), and long prolateral extension of median apophysis, longer than spur of median apophysis (vs. extension absent or as long as spur). The epigyne of N. theisi differs from those of N. isatis sp. nov. and N. spasskyi by the lack of constriction. Females of N. theisi well differ from those of N. adianta by having a white median band on carapace, darker abdominal pattern and the epigyne being almost twice longer than wide (vs. white band absent, epigyne almost as wide as long).

Comments.
Neoscona theisi is a widely distributed species, with a current natural range covering Pakistan to Japan. Levy (1998) described N. sodom on the basis of both sexes from Israel. Judging by the figures provided in the original description, there are no significant differences in the copulatory organs and habitus of N. sodom and N. theisi. Therefore, the former name is synonymized with the latter.
Distribution. Pakistan, India, Philippines, China to Indonesia, Japan. Introduced to Seychelles, Pacific Is. (WSC 2019). The westernmost localities of this species (sub N. sodom) are Cyprus (Bosmans et al. 2019) and Israel (Levy 1998). New records for Iran, Georgia, and Russia.