The spider family Oecobiidae (Arachnida: Araneae) in Iran, Afghanistan and Turkmenistan

The taxonomic and faunistic status of the spider family Oecobiidae in Iran, Afghanistan and Turkmenistan is revised. A new species, namely Uroctea gambronica sp. nov. (♂) is described from southern Iran, and the male of U. grossa Roewer, 1960 is described and illustrated for the fi rst time. Additionally, new faunistic data are provided, including the fi rst records of Oecobius putus O. PickardCambridge, 1876 and U. grossa in Afghanistan and Turkmenistan, respectively, and the re-evaluation of previously misidentifi ed and questionable records of this family in the region. The known distribution ranges of all species are mapped for these three countries.


Introduction
The spider family Oecobiidae Blackwall, 1862 is globally represented by 119 extant species in six genera, two of which occur in the Palaearctic: Oecobius Lucas, 1846 and Uroctea Dufour, 1820 (World Spider Catalog 2020). The latter genus was formerly classified as the sole member of the family Urocteidae Thorell, 1869 until Lehtinen (1967) moved it into Oecobiidae, as the single representative of the ecribellate subfamily Urocteinae Thorell, 1869. Two other subfamilies include the cribellate Oecobiinae Blackwall, 1862 and the ecribellate Uroecobiinae Kullmann & Zimmermann, 1976(Yang et al. 2019. In the Middle East and Central Asia, the family has been relatively poorly studied. In his taxonomic and biogeographic revision of the species of Oecobius occurring in the Mediterranean Region and the Arabian Peninsula, Wunderlich (1995) described several new species and provided many new distribution records of the genus. Also, country level revisions have been provided for Egypt (Hassan 1953), Iran (Zamani et al. 2017a) and Turkey (Demir et al. 2009), and more scattered contributions have been made by Zamani et al. (2016), El-Hennawy (2016), Zamani & Marusik (2018) and Boukan et al. (2018). The genus Uroctea has received far less attention in the region, with few revisionary works provided for Yemen (Rheims et al. 2007) and Turkey (Kunt et al. 2009), and scattered descriptions and records published by Roewer (1960), Zamani et al. (2015) and Fomichev & Marusik (2020). In this paper, we aim to revise the status of this family in Iran, Afghanistan and Turkmenistan, based on museum material and other recently collected specimens.

Material and methods
Specimens were studied using a Euromex MIC465 and an Olympus SZX9 stereo microscopes. Illustrations of internal genitalia were made after digestion of soft tissues in trypsin (Sigma) for 24 hours at room temperature and clearing in methyl salicylate. Lengths of leg segments were measured on the dorsal side. Leg spination is illustrated in a schematic representation where prolateral, dorsal, retrolateral and ventral sides of leg articles are flattened as a folding net (Dürer 1525). Measurements of legs are listed as: total length (femur, patella, tibia, metatarsus, tarsus). All measurements are given in millimeters. The maps (Figs 1-2) were created using the webpage SimpleMappr (online at http://www.simplemappr.net/). Geographic coordinates, if not available from the labels, were georeferenced using Google Earth (online at https://www.google.com/earth/).

Distribution
Egypt, Sudan to Iran, India. Introduced into USA, Mexico (World Spider Catalog 2020). Newly recorded from Afghanistan.

Records in Turkmenistan
Ahal and Balkan Regions (Mikhailov & Fet 1994). These records have been considered as doubtful and potentially referring to another species by Marusik et al. (2015).

Emended diagnosis
Uroctea grossa is closest to U. thaleri Rheims, Santos & van Harten, 2007 and U. gambronica sp. nov., but differs from them and from other species of Uroctea by its larger size, its broad, V-shaped MAb2 making a sharp angle (also present in U. gambronica sp. nov.) and having a wide, fan-shaped terminal section (also present in U. thaleri, cf. Fig. 7A-G), by its gently curved MAb1, its relatively long, stout, curved TL (Fig. 7C, also present in U. thaleri), its short and stout Bd and a Cd that is coiled over 360° and directed outwards anteriorly (vs uncoiled in U. limbata (C.L. Koch, 1843) and inwardly bent in U. thaleri; cf.

Description
Male (measurements based on specimen from Sistan and Baluchistan Province, Iran) Total length 9.80. Carapace broadly oval, yellowish brown with thin grey margin, faint black radial pattern and dark median band from eye region to fovea. Carapace length 4.00, width 5.05. Fovea deep triangular pit, situated 2.15 from front, length 0.30, width 0.35. Eyes surrounded by black pigment. AER 1.03, PER 1.00. AME larger than ALE, separated by slightly less than AME diameter. PLE similar in size to ALE, PME smaller, separated by four times their own diameter (= 0.5). PME and PLE touching. MOQ anterior width 0.65, posterior width 0.75, depth 0.45. Clypeus 0.60. No chilum. Chelicerae long and thin, parallel-sided, yellowish brown, length 1.35, width 0.25, no teeth. Sternum yellow with orange-brown border, smooth, heart-shaped. Sternum length 2.50, width 2.45. Labium free, subtriangular, length 0.60, width 0.65. Endites subrectangular, no serrula, no apical hair tuft. PSP long and narrow, parallel-sided, fused with sternum. Abdomen dorsally either entirely black with concentric pattern of yellowish wrinkles or bearing longitudinal oval central black mark surrounded by broad yellow zone and thin black border (Fig. 6D). Three pairs of orange-brown sigilla are present on dorsal side of abdomen. ALS subcylindrical with short apical segment. PMS small, conical. PLS with long and curved apical segment. Colulus large and covered in stiff setae. Legs greyish brown with orange brown  Palp as illustrated in Fig. 7A-C, with short, pointed and curved E connected to narrow, membranous and spoon-shaped fC, long and thin, gently curved and hooked MAb1, large, sharply angled, V-shaped MAb2 with fan-shaped tip ending in three blunt terminal lobes and with large, blunt and slightly curved TL.
Female (measurements based on holotype, GNM) Total length 13.15. Carapace broadly oval, yellowish brown, margin mottled with black (Fig. 5). Carapace length 4.15, width 5.35. Fovea deep oval pit, situated 2.35 from front, length 0.20, width 0.35. Eyes faintly ringed with black. AER 1.13, PER 1.15. AME larger than ALE, separated by slightly less than AME diameter. PLE similar in size to ALE, PME smaller, separated by 2.5 times their own diameter. Median and lateral eyes touching in both eye rows. MOQ anterior width 0.70, posterior width 0.80, depth 0.50. Clypeus 0.90. No chilum. Chelicerae long and thin, parallel-sided, yellowish brown, length 1.65, width 0.45, no teeth. Sternum light yellow brown with dark reddish brown border, smooth, heart-shaped. Sternum length 2.60, width 2.85. Labium free, subtriangular, length 0.70, width 0.85. Endites subrectangular, no serrula, no apical hair tuft. PSP long and narrow, parallel-sided, fused with sternum. Abdomen of holotype dorsally dark grey with light yellow border and thin, dark grey silky hairs (Fig. 5). Three pairs of orange-brown sigilla present on dorsal side of abdomen. ALS subcylindrical with short apical segment. PMS small, conical. PLS with long and curved apical segment. Colulus large and covered in stiff setae. Legs reddish brown, mottled with grey. Coxa I with flattened, very pale rh, length 0.20, width 0.13. Patellar indentation narrow, one third of length of pa. Tarsus IV straight. Three tarsal claws, two large ones with fine teeth. No tenent hairs.  (Fig. 9C-D) that tends to detach in some vulva preparations (Fig. 9). St small and sclerotized, connected to short and blunt Bd and adjacent to large, thin-walled bursae.

Diagnosis
Uroctea gambronica sp. nov. is closest to U. grossa and U. thaleri. It differs from both species by its short, straight, subconical TL of the male palp (vs long, curved and subcylindrical), the basal part of its MAb1 making a 135° hook (Fig. 7F, vs gently curved in U. grossa and straight in U. thaleri), its smaller, subrectangular terminal section of MAb2 (vs large and fan-shaped), and its sharply toothed retrolateral lobe of MAb2 (vs bluntly toothed). It further differs from U. grossa by its smaller size and its lower number of ti spines (Fig. 4H vs Fig. 4F). It also differs from U. thaleri by its sharply angled MAb2 (vs smoothly bent, Fig. 7D vs Fig. 7G).

Etymology
The new species is named after Gambron, the former English traders' name for Bandar Abbas, the type locality of the species.

Description Male
Total length 8.85. Carapace broadly oval, yellowish brown, somewhat mottled with grey, with faint black radiae and grey reticulate pattern near border. Carapace length 3.50, width 4.05. Fovea deep triangular pit, situated 1.70 from front, length 0.35, width 0.25. Eyes surrounded by black pigment. AER 0.90, PER 0.88. AME larger than ALE, separated by half of AME diameter. PLE similar in size to ALE, PME smaller, separated by three times their own diameter. PME and PLE touching. MOQ anterior width 0.50, posterior width 0.70, depth 0.40. Clypeus 0.65. No chilum. Chelicerae long and thin, parallelsided, yellowish brown, length 1.15, width 0.20, no teeth. Sternum yellow with light brown border, smooth, heart-shaped. Sternum length 2.20, width 2.05. Labium free, subtriangular, length 0.45, width 0.55. Endites subrectangular, no serrula, no apical hair tuft. PSP long and narrow, parallel-sided, fused with sternum. Abdomen whitish with narrow black border and central, oval, olive green patch with three pairs of olive brown sigilla (Fig. 6E). ALS light brown, blunt and conical, with short apical segment. PMS small, conical. PLS grey, with long and curved apical segment. Colulus large and covered in stiff setae. Legs yellow brown with orange brown mt and ta. Patellar indentation narrow, one third of length of pa. Tarsus  Male palp as in Fig. 7D-F, with short, pointed and curved E connected to membranous, scoop-shaped fC, with long and thin, bent and hooked MAb1, with large, sharply angled, V-shaped MAb2 with relatively short, subrectangular scoop-shaped tip ending in two blunt and one toothed terminal lobe and broad, straight, subconical, blunt TL.

Discussion
The male palp and vulva of Oecobiidae are notoriously complex. For the male palps, Shear (1970), Baum (1972) and Bosselaers (1999) used the terminology of Comstock (1910), without due consideration of homology, in spite of the fact that Comstock's terminology was mainly based on the palpal structure of Linyphiidae Blackwall, 1859 and Araneidae Clerck, 1757. Coddington (1990) highlighted this problem and proposed a palpal terminology specifically applicable to Oecobiidae and Hersiliidae Thorell, 1870. Coddington calls Baum's (1972) "radix" the "oecobiid tegular lobe". This structure is called "tegular lobe" by Rheims et al. (2007) and Kunt et al. (2009). Baum's "terminal apophysis" and "functional conductor" are named "oecobiid terminal apophysis" and "oecobiid embolic apophysis", respectively, by Coddington. Baum's (1972: fig. 62) "median apophysis" is situated proximally to the tegulum and, as a result, is certainly not a median apophysis sensu Comstock (1910: 172, 179), which is a sclerite "Arising within the distal margin of the tegulum", and "joined by a flexible articulation to the tegulum". Coddington (1990: fig. 5) completely misinterprets Baum's median apophysis and therefore has no useful name for it. What Coddington calls "median apophysis" is in fact a basal lobe of the tegulum. As a result of this confusion, Yang et al. (2019: 137) could not "match the median apophysis of Coddington (1990) and Bosselaers (1999) with male palpal structures of Uroctea species here described." The "median apophysis" of Bosselaers (1999) is basically Baum's median apophysis. Yang et al. (2019) developed their own terminology, considering Baum's terminal and subterminal apophyses as one multibranched median apophysis. We consider this interpretation justified and applied it here. Yang's et al. (2019) use of the term conductor, on the other hand, seems unjustified, as this structure is not a separate sclerite in Uroctea, but an appendage of the embolus. Therefore, we used the term "functional conductor". For Baum's "radix", we followed Rheims et al. (2007) and Kunt et al. (2009) in calling it "tegular lobe". For the sake of clarity, the palpal terminology used by different authors is summarized in Table 1.
It is noteworthy that seven species of Uroctea (U. gambronica sp. nov., U. grossa, U. hashemitorum Bosselaers, 1999, U. limbata, U. paivani (Blackwall, 1868, U. sudanensis Benoit, 1966 and U. thaleri) differ from Uroctea durandi, the type species of the genus, by the considerably larger number of prolateral spines on tarsus, metatarsus and tibia ((1)3-9(16) vs 0-3(5), Fig. 4), by the very large, scoopshaped "branch 2" of the median apophysis (vs medium-sized, flat, pointed and subtriangular), the absence of a MAb3 (vs present), the presence of a conspicuous tegular lobe in the male palp (vs very small to absent), and the presence of a blind ending duct in the vulva (vs absent). This group of species might belong to a distinct, currently undescribed genus; this matter shall be investigated in future using integrative taxonomic methods.
Considering the results of this paper, there are now nine species of Oecobiidae known from Iran, and three are known from Afghanistan and Turkmenistan. Nevertheless, the arachnological research in the region is still in the discovery phase, and new sampling efforts, especially in remote and geographically diverse areas (e.g., Zagros, Kopet Dagh and Hindu Kush mountain ranges, Sahand-Bazman Volcanic and Plutonic Belt), could yield more undescribed species and new records.