New data on zodariid spiders (Arachnida, Araneae) in the Middle East and Eastern Mediterranean

1 European Journal of Taxonomy 1016: 1–139 https://doi.org/10.5852/ejt.2025.1016.3063 europeanjournaloftaxonomy.eu ISSN 2118-9773 2025 · Shafaie S. et al. This work is licensed under a Creative Commons Attribution License (CC BY 4.0) Received: 4 April 2024 • Accepted: 9 June 2025 • Published: 18 September 2025 Topic editor: Magalie Castelin • Section editor: Arnaud Henrard • Desk editor: Kristiaan Hoedemakers Monograph urn:lsid:zoobank.org:pub:971686D8-3C1F-4F8F-9AA1-FBB8BC97E175 New data on zodariid spiders (Arachnida, Araneae) in the Middle East and Eastern Mediterranean Sepideh SHAFAIE 1 , Stano PEKÁR 2,* , Osman SEYYAR 3 1,2 Department of Botany and Zoology, Faculty of Sciences, Masaryk University, Kotlářská 2, 61137 Brno, Czech Republic. 3 Department of Biology, Faculty of Art & Sciences, Niğde Ömer Halisdemir University, TR-51240 Niğde, Turkey. * Corresponding author: [email protected] 1 Email: [email protected] 3 Email: [email protected] Abstract. Although taxonomic research on zodariid spiders has a long history, the identification of many species has remained challenging due to insufficient documentation. In this study, we revisit 50 zodariid species from the Middle East and Eastern Mediterranean. For each taxon examined, we provide detailed photographic documentation of somatic and copulatory structures to facilitate future taxonomic investigations. Seven new species from three different genera are described: Lachesana gavishae Shafaie & Pekár sp. nov. (♂♀, Israel), L. lubinae Shafaie & Pekár sp. nov. (♂♀, Israel), Pax akilae Shafaie & Pekár sp. nov. (♂♀, Syria), Zodarion jakubi Shafaie & Pekár sp. nov. (♂♀, Oman), Z. levyi Shafaie & Pekár sp. nov. (♂♀, Israel and Syria), Z. milani Shafaie & Pekár sp. nov. (♂, Syria), and Z. zonsteini Shafaie & Pekár sp. nov. (♂, Cyprus and Israel). Re-examination of the Z. nicki type specimen confirms its distinction from Z. nitidum, and we provide redescriptions of both sexes of Z. nitidum. Additionally, the male of Z. nicki and the female of Z. namrun are described here for the first time. We revalidate Z. tunetiacum and designate a neotype based on newly examined material. Two new synonymies are also proposed: Acanthinozodium crateriferum syn. nov. is synonymized with A. tibesti, and Z. kunti syn. nov. is synonymized with A. odem. Furthermore, we report ten new country records: A. odem (Syria), A. tibesti (Egypt), Z. abantense (Greece), Z. granulatum (Syria), Z. karpathos (Turkey), Z. morosum (Syria), Z. reticulatum (Turkey), Z. rhodiense (Cyprus, Syria), Z. thoni (Iran, Greece), and Z. tunetiacum (Egypt). Keywords. Ant-eating spiders, Central Asia, North Africa, new species. Shafaie S., Pekár S. & Seyyar O. 2025. New data on zodariid spiders (Arachnida, Araneae) in the Middle East and Eastern Mediterranean. European Journal of Taxonomy 1016: 1–139. https://doi.org/10.5852/ejt.2025.1016.3063 European Journal of Taxonomy 1016: 1–139 (2025) 2 Introduction Zodariidae Thorell, 1881, with a total number of 1307 species, including extant (World Spider Catalog 2025) and fossil (Dunlop et al. 2023) representatives, is amongst the most speciose spider families. The family has a wide distribution, with the highest species diversity occurring in tropical and subtropical regions of the Palearctic, Australian, African, and Neotropical realms. Despite remarkable progress in revealing the biodiversity of zodariids over the past decade (Jocqué & Henrard 2015; Bosmans et al. 2019; Coşar & Danişman 2021; Coşar et al. 2021, 2022; Lecigne & Henrard, 2022; Zamani & Marusik 2021, 2022), a large number of species remain poorly documented. For instance, over 41% of currently described species are known only from a single sex or juvenile specimens. In addition, 10 of the 90 recognized genera are monotypic (World Spider Catalog 2025). The Middle East and Eastern Mediterranean lie at the intersection of five major biodiversity hotspots (Myers et al. 2000), suggesting a high potential for species richness (Krupp et al. 2009). Despite this, current records document only 113 species from 10 genera across four subfamilies – Cryptothelinae C.L. Koch, 1872; Lachesaninae Jocqué, 1991; Storeninae Simon, 1893; Zodariinae Simon, 1890 – in this vast region (World Spider Catalog 2025), suggesting that the actual biodiversity may be significantly underestimated. Notably, a considerable number of zodariid species have been reported from countries such as Greece, Turkey, Iran, and Israel. In contrast, many other countries in the region – including Lebanon, Egypt and Yemen – report fewer than nine species, while some – such as Oman and Kuwait – lack any formal occurrence records. This disparity highlights significant gaps in our understanding of zodariid distribution and emphasizes the need for intensified sampling and taxonomic research throughout the region. While species-level identification in zodariid spiders traditionally relies on differences in the copulatory organ of both sexes, comparative examinations and habitus-based illustrations remain largely unavailable for most described species. This gap underscores a fundamental obstacle in zodariid taxonomy. For instance, Zodarion extraneum Denis, 1935, originally described from a single female specimen, was synonymized by Bosmans (1994) due to the absence of distinct copulatory organ differences between its holotype and specimens of Z. styliferum (Simon, 1870). Later, Pekár et al. (2003) elevated it to a subspecies, Z. styliferum extraneum, based on differences in chromosome number, chromosome systems and body coloration. In subsequent work, Pekár et al. (2012) noted that, despite partial reproductive isolation and the absence of mitochondrial DNA divergence between Z. s. extraneum (with the lightest coloration) and Z. s. styliferum (with the darkest coloration), the two populations displayed distinct ecological preferences, including differences in microhabitat and diet. Ortiz et al. (2021) expanded on these findings by conducting a large-scale phylogeographic study of the Z. styliferum complex across the Iberian Peninsula, revealing two allopatric species – one primarily in Portugal and the other in Spain. They explained the previously observed lack of mitochondrial divergence by identifying both lighter and darker morphs within each species. Most recently, Shafaie et al. (2025), after examining a large number of specimens, confirmed that the two species identified by Ortiz et al. (2021) could be distinguished by coloration of femora III and differences in male palp, thereby reinstating Z. extraneum as a valid species, predominantly distributed in western Iberia, especially in central-southern Portugal. Similarly, a comparative study of Z. nitidum Audouin, 1826 by Pekár et al. (2022) identified marked ecological and behavioral differences between the black and yellow morphotypes, including variation in circadian activity, prey preferences, and venom composition. These distinctions were further supported by evidence of partial to complete pre-reproductive isolation, suggesting a probable case of recent sympatric speciation. Complementing these findings, Ortiz et al. (2024) conducted a genomic analysis that strongly supports the interpretation that the yellow and black morphs represent distinct and evolutionarily independent lineages, with no detectable recent gene flow. Notably, phylogenetic SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 3 reconstructions indicate that the yellow lineage is more closely related to Z. luctuosum (O. PickardCambridge, 1872), a morphologically distinct species, than to the black lineage. These studies collectively support the hypothesis that additional species complexes may exist within the genus Zodarion, warranting detailed scrutiny of populations across the genus to detect consistent morphological differences. Such complexes often comprise taxa that exhibit pronounced genetic divergence yet display little morphological differentiation. Recognizing that coloration may hold taxonomic value comparable to other morphological traits – and considering that such features have often been insufficiently documented – we aim to expand the current taxonomic dataset by including habitus photographs where such information is lacking. Our objective is to provide a comprehensive taxonomic revision of zodariid species from the Middle East and Eastern Mediterranean regions. Given the high species richness of this family in the region, our findings will be presented across multiple volumes to support and facilitate future taxonomic work. Material and methods All descriptions were obtained from specimens immersed in ethanol. The vulva was dissected and treated for 5–11 min by gentle boiling in 20% KOH. The specimens were photographed using a KEYENCE VHX-5000 digital microscope, and the images were enhanced using Adobe Photoshop CS2 Extended. All measurements are in millimeters. The lengths of the palp and leg segments are measured on the dorsal side and are given as: total (femur, patella, tibia, metatarsus [not present in palp], and tarsus). The spination of legs I–IV is reported only for species of Lachesana. The terminology used follows that of Jocqué (1991), Levy (1992), and Bosmans (1994). Institutional abbreviations and curators CMU = Collection of Masaryk University, Brno, Czech Republic (S. Pekár) HNHM = Hungarian Natural History Museum, Budapest, Hungary (W. Wins) HUJ = Hebrew University of Jerusalem, Jerusalem, Israel (E. Gavish-Regev) MMBC = Moravské Zemské Muzeum, Brno, Czech Republic (J. Mitáček) MNHN = National Museum of Natural History, Paris, France (C. Rolland) MSNB = Museo Civico di Scienze Naturali “E. Caffi“, Bergamo, Italy (P. Pantini) NMBE = Natural History Museum in Bern, Bern, Switzerland (C. Kropf) NMP = National Museum in Prague, Prague, Czech Republic (P. Dolejš) OUMNH = Oxford University Museum of Natural History, Oxford, United Kingdom (Z. Simmons) SMF = Natural History Museum of Senckenberg, Frankfurt am Main, Germany (P. Jäger) ZMFUM = Zoological Museum of the Ferdowsi University of Mashhad, Mashhad, Iran (O. Mirshamsi) ZMH = Zoological Museum of Nature, Hamburg, Germany (D. Harms) ZMMU = Zoological Museum of the Moscow State University, Moscow, Russia (K. G. Mikhailov) ZMUT = Zoological Museum of the University of Turku, Turku, Finland (V. Vahtera) Abbreviations for morphological terms ALE = anterior lateral eye AME = anterior median eye Ap = anterior pocket Br = base of retrolateral tibial apophysis Co = conductor Cp = conductoral prong Cx = coxa d = dorsal Em = embolus FD = fertilization duct European Journal of Taxonomy 1016: 1–139 (2025) 4 Fe = femur juv. = juvenile(s) MA = median apophysis Ml = lamellar extension of the median apophysis Mp = median platelet Ms = median scape Mt = metatarsus p = prolateral Pa = patella PLE = posterior lateral eye PMA = prolateral process of median apophysis PME = posterior median eye PVS = posterior ventral spines r = retrolateral RTA = retrolateral tibial apophysis Sh = spermathecal head Sm = spermathecal medial tube Ss = spermathecal stalk Ta = tarsus Ti = tibia Ts = tibial spur v = ventral Va = ventro-retrolateral apophysis VTP = ventral transparent protrusion of the palpal tibia Results Class Arachnida Cuvier, 1812 Order Araneae Clerck, 1757 Family Zodariidae Thorell, 1881 Subfamily Lachesaninae Jocqué, 1991 Genus Lachesana Strand, 1932 Type species Lachesis perversa Audouin, 1826. Diagnosis For genus diagnosis, refer to Jocqué (1991). Lachesana gavishae Shafaie & Pekár sp. nov. urn:lsid:zoobank.org:act:E314B50D-8742-4003-BC4B-13F516A87F71 Figs 1–3, 106; Tables 1–2 Diagnosis The male of L. gavishae sp. nov. resembles those of L. perseus Zamani & Marusik, 2021 in the general shape of the retrolateral tibial apophysis, and L. insensibilis Jocqué, 1991 in the bulb details and the mesoproximal spines on the cymbium. However, the new species can be distinguished from the former species by (1) the presence of spines on the mesoproximal part of cymbium (vs absent, cf. Fig. 2A, D SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 5 Fig. 1. Lachesana gavishae Shafaie & Pekár sp. nov. A–B, E–F. Holotype, ♂ (HUJINV-Ar 21312). C–D. Paratype, ♀ (HUJINV-Ar 21313). A. Habitus, dorsal view. B. Habitus, ventral view. C. Habitus, dorsal view. D. Habitus, ventral view. E. Carapace, ventral view. F. Leg I, lateral view. Scale bars: A–E = 2 mm; F = 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 6 and Zamani & Marusik 2021: fig. 3h, j); and (2) the uniformly pale and patternless abdomen (vs with distinct pattern, cf. Fig. 1A and Zamani & Marusik 2021: fig. 1e). The new species can be distinguished from L. insensibilis by having at least 1.5 times as many spines on each leg (e.g., leg IV, 53 in L. gavishae vs 20 in L. insensibilis) and the absence of a scutum (Fig. 1A). In addition, the new species might be distinguished from all other species of Lachesana by the smallest body size (7.8 mm vs < 9.8 mm) and by the retrolateral tibial apophysis with a rather short and pointed tip (Fig. 2C, G). The female can be distinguished from those of other congeners by a heart-shaped epigynal plate with large lobes that are strongly chitinized anterolaterally (Fig. 3A, C) (vs not heart-shaped), as well as by a unique form of spermathecae, comprising a slightly divergent, upward-sloping straight spermathecal stalk ending in a convergent, globular spermathecal head (vs a cylindrical head in L. rufiventris Simon, 1873; exclusively divergent in L. tarabaevi Zonstein & Ovtchinnikov, 1999; cf. Fig. 3B, and Levy 1990: fig. 19 and Zonstein & Ovtchinnikov 1990: fig. 4, respectively). Etymology The new species is named in honor of our Israeli colleague, Efrat Gavish-Regev. Fig. 2. Male palp of Lachesana gavishae Shafaie & Pekár sp. nov., holotype (HUJINV-Ar 21312). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Dorsal view. E. Prolateral view. F. Ventral view. G. Retrolateral view. Abbreviations: see Material and methods. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 7 Type material Holotype ISRAEL • ♂; Negev Desert, Mashabim; 31.007° N, 34.739° E; 27–30 Mar. 2007; S. Pekár and S. Hénriques leg.; sand; HUJINV-Ar 21312. Paratypes ISRAEL • 2 ♀♀; same collection data as for holotype; 20 Mar. 2007; HUJINV-Ar 21313, HUJINV-Ar 21314. Other material examined ISRAEL • 1 ♀, 9 juv.; same collection data as for paratypes; CMU • 1 juv.; same collection data as for paratypes; 27–30 Mar. 2007; S. Pekár and S. Hénriques leg.; CMU • 3 juv.; same collection data as for paratypes.; 4 Apr. 2009; J. Král leg.; CMU. Description Male Habitus (Fig. 1A–B, E–F). Carapace yellow; chelicerae, mouthparts, sternum, and spinnerets pale; labium arrow-shaped and considerably longer than wide; gnathocoxae 1.5 times as long as labium. Chelicerae mesally densely covered with brown setae. Legs entirely dirty yellow except for ventrally pale coxae; legs with 214 spines (Table 1). Abdomen dirty yellow on both sides, without scutum. Fig. 3. Female copulatory organ of Lachesana gavishae Shafaie & Pekár sp. nov., paratype (HUJINVAr 21313). A. Intact epigyne, ventral view. B. Vulva, ventral view. C. Epigyne, ventral view. D. Vulva, ventral view. Abbreviations: see Material and methods. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 8 MeasureMents. Total length 7.85; carapace 4.4 long, 2 wide at pars cephalica, 3.15 wide at pars thoracica. Eye sizes and interdistances: AME: 0.19, ALE: 0.12, PME: 0.15, PLE: 0.17, AME–AME: 0.04, AME– ALE: 0.06, ALE–ALE: 0.37, PME–PME: 0.06, PME–PLE: 0.19. Legs: I 13.55 (4.25, 1.56, 3.12, 2.81, 1.81), II 14.68 (3.81, 1.5, 3.06, 4.06, 2.25), III 13.33 (4.06, 0.09, 1.94, 4.87, 2.37), IV 16.47 (4.68, 1.75, 2.93, 5.68, 1.43). Palp: 4.1 (1.6, 0.7, 0.5, 1.3). PalP (Fig. 2A–G). RTA distally oblique, both pro/retrolateral edges parallel, dorsal part tapering bladelike; RTA about 0.6 times as long as tibia, widest part over 3 times as wide as stalk. Cymbium with spines on mesoproximal side; bulb 1.5 times as long as wide. Embolus filamentous, embolar base large, about half the length of tegulum. Female (paratype HUJINV-Ar 21313) Habitus (Fig. 1C–D). Carapace yellow anteriorly, pale posteriorly; chelicerae yellowish-orange covered by black setae. Mouthparts, sternum and abdomen similar to those in male; gnathocoxae triangular shield-shaped, yellow proximally and pale distally. Carapace and abdomen without pattern. Spinnerets pale proximally and dirty yellow distally. Legs entirely yellow, with 97 spines (Table 2). MeasureMents. Total length 10.95; carapace 5.4 long, 2.4 wide at pars cephalica, 3.52 wide at pars thoracica. Eye sizes and interdistances: AME: 0.15, ALE: 0.12, PME: 0.12, PLE: 0.13, AME–AME: 0.06, AME–ALE: 0.09, ALE–ALE: 0.46, PME–PME: 0.09, PME–PLE: 0.34. Legs: I 12.8 (4.1, 1.9, 2.9, 2.4, 1.5), II 12.5 (4.7, 1.8, 2.1, 2.5, 1.4), III 11.3 (3.3, 1.8, 1.5, 3, 1.7), IV 14.1 (3.9, 1.8, 2.5, 3.9, 2). ePigyne (Fig. 3A–D). Epigynal plate about 1.3 times as wide as long, including heart-shaped plate, both large lobes darkly chitinized anterolaterally; two divergent copulatory ducts extended laterally. Anterior pockets strongly sinuous, touching each other. Spermathecae situated along epigynal plate and confining edges of epigynal pockets, including rather long, divergent, upward-sloping straight spermathecal stalks ending in globular and convergent spermathecal heads; spermathecal heads separated by about their diameter. Size variation Total body length 10.95–13.77, carapace length/width 5.4–6.87/2.4–3.5 in females (N = 3). Distribution The species is currently only known from the type locality in Israel. Table 1. Male leg spination of Lachesana gavishae Shafaie & Pekár sp. nov. Abbreviations: d = dorsal; Fe = femur; Mt = metatarsus; p = prolateral; Pa = patella; r = retrolateral; Ti = tibia; v = ventral. Fe Pa Ti Mt Total Id4 p5 r6 v1 p1 r1 p4 r2 v10 p2 r5 v9 50 II d4 p7 r5 v1 d1 p2 r1 p4 r3 v12 p4 r2 v11 57 III d3 p6 r4 v5 d1 p2 r1 d2 p3 r4 v6 d3 p3 r2 v9 54 IV d4 p6 r4 v3 d2 p1 r1 d2 p3 r4 v6 d3 p5 r2 v7 53 SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 9 Lachesana lubinae Shafaie & Pekár sp. nov. urn:lsid:zoobank.org:act:EF76D45C-001C-4947-B30C-D43AD30FF63C Figs 4–6, 106; Tables 3–4 Diagnosis This new species is most similar to L. gavishae sp. nov. in terms of body coloration and overall palp morphology. The male of the new species can be recognized by a wider retrolateral tibial apophysis apical width/basal width ratio (3.8 vs 3.1); and different number of spines on leg segments and cymbium (i.e., metatarsus spines 89 vs 57 and cymbium spines 9 vs 16). The female can be distinguished by a remarkably spatulated, mushroom-shaped epigynal plate (vs heart-shaped, cf. Fig. 6A, C and Fig. 3A, C), and convergent spermathecal stalks ending in kidney-shaped spermathecal heads (vs parallel spermathecal stalks, with globular spermathecal heads, cf. Fig. 6B, D and Fig. 3B, D). Furthermore, the total number of leg spines can separate both adult and juvenile specimens of L. lubinae sp. nov. from those of L. gavishae (> 130 spines vs < 97). Etymology This species is named in honor of our colleague, the Israeli arachnologist, Yael Lubin. Type material Holotype ISRAEL • ♂; Negev Desert, Mashabim; 31.007° N, 34.739° E; 19 Oct. 2004; J. Král leg.; sands; HUJINV-Ar 21315. Paratype ISRAEL • 1 ♀; same collection data as for holotype; 20 Mar. 2007; S. Pekár leg.; HUJINV-Ar 21316. Other material examined ISRAEL • 6 juv.; same collection data as for paratype; CMU • 7 juv.; same collection data as for paratype; 27–30 Mar. 2007; S. Pekár and S. Hénriques leg.; CMU • 2 juv.; same collection data as for paratype; 4 Apr. 2009; J. Král leg.; CMU. Description Male Habitus (Fig. 4A–B, E, G). Carapace dark yellow, with several pale stripes radiating from fovea; chelicerae, gnathocoxae, and arrow-shaped labium pale, labium as long as wide; chelicerae mesally densely covered with thin, brown setae; gnathocoxae 1.6 times as long as labium. Sternum pale. Fe, Pa, Table 2. Female leg spination of Lachesana gavishae Shafaie & Pekár sp. nov. Abbreviations: d = dorsal; Fe = femur; Mt = metatarsus; p = prolateral; Pa = patella; r = retrolateral; Ti = tibia; v = ventral. Fe Pa Ti Mt Total Id4 p1 r1 v0 p2 r1 p0 r0 v4 p0 r0 v10 23 II d4 p1 r0 v0 P1 r0 p2 r0 v2 p1 r0 v10 21 III d4 p1 r0 v0 p3 r2 d0 p3 r4 v0 d1 p3 r2 v6 29 IV d3 p0 r0 v0 p3 r1 d0 p3 r2 v0 d1 p4 r2 v5 24 European Journal of Taxonomy 1016: 1–139 (2025) 16 MeasureMents. Total length 7.56; carapace 3.95 long, 1.01 wide at pars cephalica, 2.65 wide at pars thoracica, scutum length 2.59. Eye sizes and interdistances: AME 0.33, ALE 0.39, PME 0.34, PLE 0.39, AME–AME 0.17, AME–ALE 0.28, ALE–ALE 1.29, PME–PME 0.22, PME–PLE 0.73. Legs: I 8.44 (2.37, 1.12, 1.8, 1.74, 1.41), II 7.99 (2.59, 1.12, 1.52, 1.58, 1.18), III 7.98 (2.3, 1.12, 1.41, 2.03, 1.12), IV 10.29 (2.7, 1.18, 2.24, 2.59, 1.58). Palp: 4.11 (1.58, 0.34, 0.28, 1.91). PalP (Fig. 8A–F). RTA with an oblong extension, comprising a rather long, forward pointed tibial spur and a robust hump. Median platelet extended mesally and anteriorly; conductoral prong finger-shaped; median apophysis rectangular basally and narrowed apically, with a folded tip. Embolus very long, originating at a 4 o’clock position, narrowed slightly and folded around bulb. Female (paratype NMP P6A 7515) Habitus (Fig. 7C–D). Carapace, chelicerae, sternum, mouthparts similar to those in male. Legs yellow except for slightly darker Cx, greyish Fe and light grey tips of Mt; spinnerets pale, palp segments greyish, with darker Ta. Abdomen blackish-brown dorsally, with no pattern, dark brown ventrally, with three subparallel yellow stripes in the middle, lateral stripes wider than middle stripe. MeasureMents. Total length 8.29; carapace 3.5 long, 1.24 wide at pars cephalica, 2.37 wide at pars thoracica. Eye sizes and interdistances: AME 0.33, ALE 0.39, PME 0.45, PLE 0.39, AME–AME 0.11, AME–ALE 0.33, ALE–ALE 0.99, PME–PME 0.17, PME–PLE 0.56. Legs: I 8.44 (2.37, 1.12, 1.8, 1.74, 1.41), II 7.99 (2.59, 1.12, 1.52, 1.58, 1.18), III 7.71 (2.03, 1.12, 1.41, 2.03, 1.12), IV 10.29 (2.7, 1.18, 2.24, 2.59, 1.58). Palp: 3.03 (1.01, 0.45, 0.45, 1.12). Fig. 9. Female copulatory organ of Pax akilae Shafaie & Pekár sp. nov., paratype (NMP P6A 7515). A. Intact epigyne, ventral view. B. Epigyne, ventral view. C. Vulva, ventral view. D. Vulva, ventral view. Abbreviation: see Material and methods. Scale bars: 0.4 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 17 ePigyne (Fig. 9A–D). Epigyne about 1.6 times as wide as long, comprising three longitudinal tubes: one dark medial and two light, sub-lateral tubes accompanied by a pair of dark marginal tubes. Spermathecal tubes parallel comprising a pair of elongated, yellow structures, situated medially, externally attached to a pair of reddishbrown, larger structures ending in twisted coils. Size variation Total body length 6.5–7.62, carapace length/width 3.5–3.95/2.01–2.75 in males (N = 3). Total body length 7.12–8.29, carapace length/width 3–3.5/2.24–2.3 in females (N = 5). Distribution The species is currently only known from the type locality in Syria. Pax engediensis Levy, 1990 Fig. 10 Pax engediensis Levy, 1990: 356, figs 64–67. Diagnosis Refer to Levy (1990). Type material examined Holotype ISRAEL • ♀; En Gedi; 26 Mar. 1957; A. Shulov leg.; HUJ 14216. Description For a detailed description of female, refer to Levy (1990). Distribution Israel. Pax islamita (Simon, 1873) Figs 11–12 Habronestes islamita Simon, 1873: 75. Storena islamita – Simon 1893: 247. Pax islamita – Levy 1990: 358, figs 68–71. — Seyyar et al. 2008: 52, figs 1–11. Diagnosis Refer to Levy (1990). Material examined ISRAEL • 3 ♀♀, 9 juv.; Adullam; 31.641° N, 34.931° E; 25 Mar. 2001; S. Pekár, S. Hénriques and M. Řezáč leg.; CMU • 2 ♂♂, 6 ♀♀, 2 juv.; same collection data as for preceding; 5 Apr. 2017; S. Pekár and S. Korenko leg.; CMU • 1 ♂; Mount Carmel; 13 Apr. 2009; J. Král leg.; CMU • 1 ♂; Carmel; 32.733° N, 35.05° E; 6 May 2023; M. Řezáč leg.; CMU. European Journal of Taxonomy 1016: 1–139 (2025) 18 TURKEY • 1 ♂; SE Turkey; 28 Apr. 2004; M. Řezáč leg.; CMU • 1 ♂, 1 ♀; Toprakkale; 37.061° N, 36.148° E; 67 m a.s.l.; 1 May 2009; CMU • 1 ♂; Mersin, 12 km to Anamur, 1 km to Ala Bridge; 36.174° N, 32.894° E; 93 m a.s.l.; 14 May 2009; R. Kaya leg.; ZMUT. Description For a detailed description, refer to Levy (1990). Distribution Turkey, Israel, Syria, Lebanon. Fig. 10. Pax engediensis Levy, 1990, holotype, ♀ (HUJ 14216). A. Habitus, dorsal view. B. Habitus, ventral view. C. Intact epigyne, ventral view. Scale bars: A–B = 1 mm; C = 0.4 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 19 Fig. 11. Habitus of Pax islamita (Simon, 1873) (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 20 Fig. 12. Copulatory organs of Pax islamita (Simon, 1873) (CMU). A–G. Male palp. H–I. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Ventro-retrolateral view. D. Retrolateral view. E. Prolateral view. F. Ventral view. G. Retrolateral view. H. Intact epigyne, ventral view. I. Vulva, ventral view. Scale bars: 0.4 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 21 Pax palmonii Levy, 1990 Figs 13–14 Pax palmonii Levy, 1990: 352, figs 60–63. Diagnosis Refer to Levy (1990). Type material examined Holotype ISRAEL • ♂; Ma’oz Hayyam; 30 May 1943; A. Shulov leg.; HUJ 14215. Paratype ISRAEL • 1 ♀; same collection data as for holotype; HUJ 14215. Description For a detailed description, refer to Levy (1990). Distribution Israel. Subfamily Zodariinae Simon, 1890 Genus Acanthinozodium Denis, 1966 Type species Acanthinozodium spinulosum Denis, 1966. Diagnosis For genus diagnosis, refer to Jocqué & Henrard (2015). Acanthinozodium odem (Levy, 2007) Figs 15–16, 105 Zodarion odem Levy, 2007: 15, figs 38–39. Zodarion kunti Coşar et al., 2021: 49, figs 5a–f, 6a–d, 7a–c, 8a–c, 9a–b, 10a–b. Syn. nov. Acanthinozodium odem – Zonstein 2025: 392, figs 1–21. Diagnosis Refer to Zonstein (2025). Type material examined Holotype ISRAEL • ♀; Odem Forest, Golan Heights (219/290 Is. Gr.); 950 m a.s.l.; May 1996; S. Rakefet leg.; HUJ 15220. European Journal of Taxonomy 1016: 1–139 (2025) 22 Fig. 13. Habitus of Pax palmonii Levy, 1990. A–B. Holotype, ♂ (HUJ 14215). C–D. Paratype, ♀ (HUJ 14215). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 23 Fig. 14. Copulatory organs of Pax palmonii Levy, 1990. A–D. Holotype, male palp (HUJ 14215). E. Paratype, female copulatory organ (HUJ 14215). A. Palp, prolateral view. B. Palp, ventral view. C. Palp, ventro-retrolateral view. D. Palp, retrolateral view. E. Intact epigyne, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 24 Fig. 15. Habitus of Acanthinozodium odem (Levy, 2007). A–B. ♂ (CMU). C–D. ♀ (CMU). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. Fig. 16 (next page). Copulatory organs of Acanthinozodium odem (Levy, 2007). A–G. Male palp (CMU). H–J. Female copulatory organ (CMU), specimens from Syria (H) and Israel (I–J). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Prolateral view. E. Ventral view. F. Retrolateral view, arrow indicates circular pit. G. Embolar tip, ventral view. H. Intact epigyne, ventral view. I. Intact epigyne, ventral view. J. Vulva, ventral view. Abbreviation: see Material and methods. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 25 European Journal of Taxonomy 1016: 1–139 (2025) 32 Fig. 21. Habitus of Palaestina expolita O. Pickard-Cambridge, 1872 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Abbreviation: see Material and methods. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 33 Palaestina eremica Levy, 1992 Figs 23–24 Palaestina eremica Levy, 1992: 74, figs 19–26. Diagnosis Refer to Levy (1992). Type material examined Holotype EGYPT • ♂; Sinai, St Catherine Monastery, (049/775 Isr.grid); 14 Apr. 1987; V.D. Roth leg.; HUJ 14250. Fig. 22. Copulatory organs of Palaestina expolita O. Pickard-Cambridge, 1872 (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Palp with broken median apophysis, ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Abbreviations: see Material and methods. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 34 Fig. 23. Habitus of Palaestina eremica Levy, 1992. A–B. Holotype, ♂ (HUJ 14250). C–D. Paratype, ♀ (HUJ 14251). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 35 Paratype EGYPT • 1 ♀; same collection data as for holotype; HUJ 14251. Description For a detailed description, refer to Levy (1992). Distribution Egypt, Israel. Fig. 24. Copulatory organs of Palaestina eremica Levy, 1992. A–C. Holotype, male palp (HUJ 14250). D. Paratype, female (HUJ 14251). A. Palp, prolateral view. B. Palp, ventral view. C. Palp, retrolateral view. D. Intact epigyne, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 36 Genus Parazodarion Ovtchinnikov, Ahmad & Gurko, 2009 Type species Zodarion raddei Simon, 1889. Diagnosis For genus diagnosis, refer to Ovtchinnikov et al. (2009). Parazodarion raddei (Simon, 1889) Figs 25–26, 106 Zodarion raddei Simon, 1889: 383. Zodarion vlasovi Sytshevskaja in Vlassov & Sytshevskaja, 1937: 249, figs 1–3. Zodarion raddei – Denis 1958: 111, figs 42–44. — Fet 1985: 274. — Marusik & Koponen 2001: 46, fig. 27. Parazodarion raddei – Ovtchinnikov et al. 2009: 471, fig. 1.1–6. — Zamani & Marusik 2021: 182, fig. 26a–c. Diagnosis Refer to Ovtchinnikov et al. (2009). Material examined AFGHANISTAN • 1 ♀; Badghis, Bala Murghab; 18 Jul.–4 Sep. 1964; O. Jakeš leg.; MMBC 7633 (5601/ Ent). IRAN • 5 ♀♀; Taybad, Dowqarun; 8 Nov. 2014; P. Rashidi leg.; ZMFUM-TA 1183. Description For a detailed description, refer to Ovtchinnikov et al. (2009). Remarks The single female specimen from Afghanistan exhibits slight differences compared to the Iranian population, particularly in the height of the spermathecal bodies, which are arranged in descending order (vs more or less equally sized, cf. Fig. 26D and Fig. 26B), and in the lighter proximal parts of Fe I and II (vs entirely dark, cf. Fig. 25C and Fig. 25A). Given the unlikely prospect of obtaining additional material from Afghanistan in the near future, we treated this single specimen as P. raddei. Distribution Afghanistan, Iran, Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, United Arab Emirates. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 37 Fig. 25. Habitus of Parazodarion raddei (Simon, 1889), ♀. A–B. Specimen from Iran (ZMFUM-TA 1183). C–D. Specimen from Afghanistan (MMBC 7633 (5601/Ent). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 38 Genus Trygetus Simon, 1882 Type species Trygetus sexoculatus O. Pickard-Cambridge, 1872. Diagnosis For genus diagnosis, refer to Levy (1992). Trygetus sexoculatus (O. Pickard-Cambridge, 1872) Figs 27–28 Palaestina sexoculata O. Pickard-Cambridge, 1872: 270. Trygetus senoculatus – Simon 1893: 439. Trygetus sexoculatus – Jocqué 1991: 149, figs 378–383. — Levy 1992: 77, figs 27–38. Diagnosis Refer to Jocqué (1991) and Levy (1992). Fig. 26. Female copulatory organs of Parazodarion raddei (Simon, 1889). A–B. Specimen from Iran (ZMFUM-TA 1183). C–D. Specimen from Afghanistan (MMBC 7633 (5601/Ent). A. Epigyne, ventral view. B. Vulva, ventral view. C. Intact epigyne, ventral view. D. Epigyne, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 39 Material examined ISRAEL • 2 ♂♂; Sayert Shaked; 1999; J. Levy leg.; CMU • 1 ♀; Lehavim; 31.364° N, 34.823° E; 12 Apr. 2001; S. Pekár leg.; CMU • 2 ♀♀; same collection data as for preceding; 20 Apr. 2001; S. Pekár leg.; CMU • 1 ♂; Halukim; 30.854° N, 34.768° E; 21 Mar. 2007; S. Pekár leg.; CMU • 8 ♂♂, 3 ♀♀, 1 juv.; same collection data as for preceding; 7 Apr. 2019; S. Pekár leg.; CMU • 2 ♂♂; same collection data as for preceding; 29 Mar. 2023; S. Pekár leg.; CMU • 1 ♀; Wadi Paran; 8 Apr. 2019; S. Pekár leg.; CMU. Description For a detailed description, refer to Levy (1992). Distribution Egypt, Israel. Fig. 27. Habitus of Trygetus sexoculatus (O. Pickard-Cambridge, 1872) (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 40 Genus Zodarion Walckenaer, 1826 Type species Enyo nitida Audouin, 1826. Diagnosis For genus diagnosis, refer to Levy (1992). Fig. 28. Copulatory organs of Trygetus sexoculatus (O. Pickard-Cambridge, 1872) (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 41 Zodarion nitidum Audouin, 1826 Figs 29–31, 87, 106 Enyo nitida Audouin, 1826: 350, pl. 3 fig. 7. Enyo longipes Audouin, 1826: 351, pl. 3 fig. 8. Zodarion cambridgei Denis, 1938: 387. Clotho nitida – Walckenaer 1837: 639. Clotho longipes – Walckenaer 1837: 640. Argus longipes – Walckenaer 1841: 347, pl. 16 fig. 6. Zodarion longipes – Walckenaer 1847: 563. Enyo nitida – Walckenaer 1847: 563. — Thorell 1869: 108. — F.O. Pickard-Cambridge 1876: 559. Enyo luisante – Simon 1864: 160. Zodarium longipes – Thorell 1869: 107. Zodarium nitidum – Simon 1885: 29. Zodarion nitidum – Simon 1893: 432, figs 403–407; 1908: 425. — Denis 1937: 6, pl. 1 fig. 1; 1962: 31, fig. 2. — Levy 1992: 86, figs 53–61. — Zamani & Marusik 2021: 188, fig. 28a–c; 2022: 162, fig. 11a–c. Metargus longipes – F. Pickard-Cambridge 1902: 8. Diagnosis Zodarion nitidum specimens are most similar to Z. nicki in the shape of their copulatory organs, but the former species is distinguished by a brownish carapace and yellow coxae (vs black carapace and black to grey coxae; cf. Fig. 29A, C and Fig. 84A, C). Furthermore, males of Z. nitidum possess uniformly yellow femora (vs entirely black femora in Z. nicki; cf. Fig. 29A and Fig. 84A). Material examined EGYPT • 1 ♀; Alexandria; MNHN-AR2854 • 2 ♀♀; Cairo or Aswan, Lake Mariout; MNHN-AR2847. ISRAEL • 1 ♂; Midreshet Sde Boker; 30.857° N, 34.781° E; 1 Apr. 2001; S. Pekár leg.; HUJINV-Ar 21301 • 1 ♀, same collection data as for preceding; HUJINV-Ar 21302 • 1 juv.; Midreshet Sde Boker; 10 Mar. 2001; S. Pekár leg.; CMU • 1 ♂; Arava Area, 12 km N of Ein Yahav Moshav, Hatzeva Moshav; 30.754° N, 35.269° E; 13 Mar. 2001; S. Pekár leg.; CMU • 1 ♀, 3 juv.; s. loc.; Apr. 2001; S. Pekár leg.; CMU • 12 ♂♂, 9 ♀♀, 50 juv.; same collection data as for preceding; Feb.–Mar. 2018; L. Sentenská leg.; CMU • 1 ♀, 2 juv.; same collection data as for preceding; 6 Apr. 2019; S. Pekár leg.; CMU • 9 ♂♂ 3 ♀♀ 1 juv.; Halukim; 30.854° N, 34.784° E; 16 Apr. 2001; S. Pekár leg.; CMU • 1 ♀; same collection data as for preceding; 27 Mar. 2007; S. Pekár leg.; CMU • 3 ♂♂, 1 juv.; same collection data as for preceding; 18 Mar. 2017; O. Macháč leg.; CMU • 6 ♂♂; same collection data as for preceding; 9 Apr. 2019; S. Pekár leg.; CMU • 1 ♀; Mashabim; 31.000° N, 34.757° E; 4 Sep. 2002; Y. Lubin leg.; CMU • 2 ♀♀; same collection data as for preceding; Sep. 2004; J. Král leg.; CMU • 1 ♂, 1 ♀; same collection data as for preceding; 1 Apr. 2016; S. Korenko leg.; CMU • 1 ♂; same collection data as for preceding; 1 Apr. 2017; S. Korenko leg.; CMU • 1 juv.; same collection data as for preceding; 6 Apr. 2019; S. Pekár leg.; CMU • 1 ♀; same collection data as for preceding; 28 Mar. 2023; S. Pekár and V. Opatová leg.; CMU • 1 ♀; same collection data as for preceding; 29 Mar. 2023; S. Pekár leg.; CMU • 1 ♂; Beersheba, between Beer Sheva and Mitzpe Ramon; 6–29 May 2003; M. Řezáč leg.; CMU • 1 ♀; Meirav; 29 Sep. 2007; J. Král leg.; CMU • 2 ♂♂, 2 juv.; Yeroham; 30.971° N, 34.97° E; 3 Mar. 2016; S. Pekár, S. Korenko and E. Líznarová leg.; CMU • 18 ♂♂, 11 ♀♀; same collection data as for preceding; 5 Apr. 2017; S. Pekár and S. Korenko leg.; CMU • 2 ♀♀, 8 juv.; same collection data as for preceding; 18 May 2017; O. Macháč leg.; CMU • 3 juv.; same collection data as for preceding; 5 Apr. 2019; S. Pekár leg.; CMU • 1 juv.; Ramon Crater, Nahal Paran; 30.333° N, 34.968° E; 8 Apr. 2019; S. Pekár leg.; CMU • 6 ♀♀, 2 juv.; s. loc.; May 2019; S. Pekár, L. Sentenská and S. Korenko leg.; CMU • 2 juv.; Hatira; 30.872° N, 34.816° E; 28 Mar. 2023; S. Pekár and V. Opatová leg.; CMU • 1 ♂, 1 ♀, 4 juv.; Borot Lotz; 30.511° N, 34.612° E; 29 Mar. 2023; S. Pekár and V. Opatová leg.; CMU. European Journal of Taxonomy 1016: 1–139 (2025) 48 Fig. 33. Zodarion aculeatum Chyzer, 1897, holotype, ♀ (HNHM 1187). A. Prosoma, dorsal view. B. Prosoma, ventral view. C. Abdomen, dorsal view. D. Abdomen, ventral view. E. Epigyne, ventral view. Scale bars: A–D = 1 mm; E = 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 49 Zodarion barbarae Bosmans, 2009 Figs 34–35 Zodarion barbarae Bosmans, 2009: 248, figs 80–83, 138–139. Zodarion barbarae – Bosmans et al. 2019: 32, fig. 20a–g. — Özkütük 2022: 241, figs 1–15. — Coşar & Danışman 2024: 268, figs 21–22. Fig. 34. Habitus of Zodarion barbarae Bosmans, 2009. A–B. Holotype, ♂ (NMBE-AR4147). C–D. Paratype, ♀ (NMBE-AR4147). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 50 Diagnosis Refer to Bosmans (2009). Type material examined Holotype GREECE • ♂; Peloponnisos, Oros Taigetos, Moni Panaghi Giatrissa; 1100 m a.s.l.; 30 Sep. 1991; B. Knoflach and K. Thaler leg.; NMBE-AR4147. Paratypes GREECE • 2 ♀♀; same collection data as for holotype; NMBE-AR4147. Fig. 35. Copulatory organs of Zodarion barbarae Bosmans, 2009. A–C. Holotype, male palp (NMBEAR4147). D–E. Paratype, female copulatory organ (NMBE-AR4147). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 51 Other material examined CYPRUS • 1 ♂, 14 juv.; Nicosia, Lefka Dam; 35.063° N, 32.837° E; 288 m a.s.l.; 25 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU. TURKEY • 1 ♂, 1 ♀ 14 juv.; Konakli, Macchie Hill, 10 km W of Alanya; 20 May 2005; J. Dolanský leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Distribution Greece, Cyprus, Turkey. Zodarion beroni Komnenov & Chatzaki, 2016 Figs 36–37 Zodarion beroni Komnenov & Chatzaki in Komnenov et al., 2016: 41, figs 97–111. Diagnosis Refer to Komnenov et al. (2016). Material examined GREECE • 1 ♂; 20 km after Dadia; 660 m a.s.l.; 22 Jun.–21 Jul. 2015; M. Chatzaki leg.; CMU • 6 ♂♂, 1 ♀; 2 km after Pefka/Retziki Suburb; 121 m a.s.l.; 22 Jun.–21 Jul. 2015; M. Chatzaki leg.; CMU • 2 ♂♂; between Aisymi-Mega Dereio, 500 m W.; 389 m a.s.l.; 22 Jul.–26 Aug. 2015; M. Chatzaki leg.; CMU. Description For a detailed description, refer to Komnenov et al. (2016). Distribution Greece. Zodarion blagoevi Bosmans, 2009 Fig. 38 Zodarion blagoevi Bosmans, 2009: 249, figs 84–87, 140–141. Zodarion blagoevi – Deltshev et al. 2022: 248, figs 2, 6, 10, 14, 20–22, 30, 34, 38, 42, 46, 49, 52, 55. Diagnosis Refer to Bosmans (2009). Material examined GREECE • 10 ♂♂; Aladjagiola Wetland (N of Chrisoupoli); 41.01° N, 24.703° E; 25 Jun. 2006; M. Schröder leg.; CMU. Description For a detailed description, refer to Bosmans (2009). European Journal of Taxonomy 1016: 1–139 (2025) 52 Fig. 36. Habitus of Zodarion beroni Komnenov & Chatzaki, 2016 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 0.4 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 53 Distribution Bulgaria, Greece. Zodarion christae Bosmans, 2009 Figs 39–40 Zodarion christae Bosmans, 2009: 284, figs 186–187, 196–197. Zodarion christae – Coşar et al. 2021: 46, figs 1a–f, 2a–e, 3a–b. Diagnosis Refer to Bosmans (2009). Fig. 37. Copulatory organs of Zodarion beroni Komnenov & Chatzaki, 2016 (CMU). A–C, F. Male palp, with rotated bulb inside. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. F. Detail of embolus, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 54 Fig. 38. Zodarion blagoevi Bosmans, 2009 (CMU), ♂. A. Habitus, dorsal view. B. Habitus, ventral view. C. Palp, prolateral view. D. Palp, ventral view. E. Palp, retrolateral view. Scale bars: A–B = 1 mm; C–E = 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 55 Fig. 39. Habitus of Zodarion christae Bosmans, 2009 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 56 Material examined TURKEY • 1 ♂, 1 ♀; Isparta, Aksu, Dedegöl Mountain; 37.656° N, 31.291° E; 8 Jul. 2003; M. Řezáč leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Remarks Due to damage sustained by the median apophyses in the male palps and the epigyne during the clearing process, it was not possible to include photographs of these structures in this study. Distribution Greece, Turkey. Fig. 40. Copulatory organs of Zodarion christae Bosmans, 2009 (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 57 Zodarion crewsae Coşar, Danışman & Kunt, 2022 Figs 41–42, 106 Zodarion crewsae Coşar et al. 2022: 229, figs 1–16. Diagnosis Refer to Coşar et al. (2022). Material examined TURKEY • 1 ♂; Tunceli, Munzur River; 38.957° N, 39.561° E; 22 Jun. 2002; M. Řezáč leg.; CMU. Description For a detailed description, refer to Coşar et al. (2022). Remarks A figure-based comparison of the single specimen from eastern Turkey with specimen from the southern population revealed that the eastern specimen exibited a significantly shorter distal branch of the median apophysis, and a lighter femur I and carapace. However, to accurately classify this population, additional specimens are needed to evaluate whether it represents a new species. Distribution Turkey. Fig. 41. Habitus of Zodarion crewsae Coşar, Danışman & Kunt, 2022 (CMU), ♂. A. Dorsal view. B. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 64 Zodarion frenatum Simon, 1885 Figs 48–49 Zodarion frenatum Simon, 1885: 336. Zodarion frenatum – Fage 1921: 174, fig. 2. — Denis 1937: 39, pl. 8 figs 65–68. — Wunderlich 1980: 240. — Bosmans 1997: 271, figs 12–13, 85–86; 2009: 286, figs 192–193, 198–199. — Dimitrov 2024: 114, figs 10–14. Zodarion creticum – Roewer 1928: 119, pl. 2 fig. 37. Zodarion rhodiense – Caporiacco 1948: 47, fig. 6. Fig. 48. Habitus of Zodarion frenatum Simon, 1885 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 65 Zodarion rhodiense nigrifemur – Caporiacco 1948: 49. Zodarion ionicum – Brignoli 1984: 314, figs 41–42. Diagnosis Refer to Bosmans (1997). Type material examined Paratype GREECE • 1 juv.; Naxos; E. Keyseling leg.; MNHN-AR2850. Fig. 49. Copulatory organs of Zodarion frenatum Simon, 1885 (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 66 Other material examined GREECE • 1 ♀, 1 juv.; Volo (St.); MNHN-AR2851 • 1 ♀; 1.5 km after Dadia; 660 m a.s.l.; 14–22 May 2015; M. Chatzaki leg.; CMU • 3 ♂♂, 2 juv.; same collection data as for preceding; 22 Jun.–21 Jul. 2015; M. Chatzaki leg.; CMU • 2 ♂♂, 1 ♀, 1 juv.; 2 km after Pefka/Retziki Suburb; 121 m a.s.l.; 16 May–22 Jun. 2015; M. Chatzaki leg.; CMU • 1 ♀; same collection data as for preceding; 22 Jul.–24 Aug. 2015; M. Chatzaki leg.; CMU • 1 ♂; 1.5 km N of Polia; 50 m a.s.l.; 22 Jul.–26 Aug. 2015; M. Chatzaki leg.; CMU. Description For a detailed description, refer to Bosmans (1997). Remarks Even though Bosmans (1997) did not examine the type of Z. frenatum, he reported that the holotype tube (MNHN-AR2850) contained one male and one juvenile. However, our examination of the tube revealed only a single juvenile specimen, suggesting that the holotype male is probably lost. Furthermore, while Fage (1921) and Bosmans (1997) recorded a single female in tube MNHN-AR2851, we identified two specimens, including one female and one juvenile. Distribution Italy, Albania, North Macedonia, Bulgaria, Greece, Turkey. Zodarion geshur Levy, 2007 Figs 50–51 Zodarion geshur Levy 2007: 13, figs 33–37. Diagnosis Refer to Levy (2007). Type material examined Holotype ISRAEL • ♂; Geshur, S Golan (217/240 .3. J); May 1998; R. Sharon leg.; HUJ 15215. Paratype ISRAEL • 1 ♀; same collection data as for holotype; HUJ 15215. Description For a detailed description, refer to Levy (2007). Remarks Although Levy (2007) noted that the type specimens were collected in May 1996, the label indicates May 1998. Distribution Israel. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 67 Fig. 50. Habitus of Zodarion geshur Levy, 2007. A–B. Holotype, ♂ (HUJ 15215). C–D. Paratype, ♀ (HUJ 15215). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 68 Fig. 51. Copulatory organs of Zodarion geshur Levy, 2007. A–C. Holotype, male palp (HUJ 15215). D. Paratype, female intact epigyne (HUJ 15215). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 69 Zodarion granulatum Kulczyński, 1908 Figs 52–53, 105 Zodarion granulatum Kulczyński, 1908: 59, pl. 2 figs 4, 10. Zodarion granulatum – Levy 1992: 92, figs 72–84. — Bosmans 2009: 275, figs 170–171, 178–179. — Lecigne & Henrard 2022: 37, fig. 5e, h. — Bosmans & Gavalas 2023: 33, fig. 21a–e. — Dimitrov 2024: 114, fig. 18. Zodarion parashi – Wunderlich 2022: 48, figs 105–106. Diagnosis Refer to Bosmans (2009) and Lecigne & Henrard (2022). Type material examined Holotype (Zodarion parashi) GREECE • ♂; Eastern Aegean Island Naxos; 25.57° N, 37.0.37° E; 20 Jul. 1982; L. Parashi leg.; ZMH A0014216. Material examined CYPRUS • 1 ♂ (subadult); Nicosia, Olympus, Moutoulas Mountain; 34.977° N, 32.828° E; 800 m a.s.l.; 25 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU. GREECE • 1 ♂; Aladjagiola Wetland (N Chrisoupoli); 28 Jun. 2008; M. Schröder leg.; CMU. SYRIA • 1 ♂; Aln Al-Zarah, near Idlib, Canyon of Oronthes; 18 Mar. 2001; M. Řezáč leg.; CMU • 1 ♀; Mashta Al-Helu, Odem Forest; 3 Apr. 2001; M. Řezáč leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Remarks The length and arrangement of the PVS differ between males and females. Male specimens possess significantly longer PVS, which are arranged in a somewhat radiating pattern (Fig. 52B), whereas females have shorter PVS arranged parallel to each other (Fig. 52D). Moreover, the single male specimen from Syria exhibits a smaller PMA compared to the specimen from Greece, along with a more slender embolar tip. Distribution This species was previously recorded in Greece, Cyprus, Turkey, Lebanon, and Israel (World Spider Catalog 2025). This is the first report of this species from Syria European Journal of Taxonomy 1016: 1–139 (2025) 70 Fig. 52. Habitus of Zodarion granulatum Kulczyński, 1908 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view, arrow indicates granulations. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 71 Zodarion hauseri Brignoli, 1984 Figs 54–55 Zodarion hauseri Brignoli, 1984: 312, fig. 40. Zodarion hauseri – Stefanovska et al. 2008: 37, figs 5–9. — Bosmans 2009: 251, figs 92–95, 144–145. — Naumova et al. 2017: 480, figs 28–30. Diagnosis Refer to Bosmans (2009). Material examined GREECE • 1 ♂; Aladjagiola Wetland (N of Chrisoupoli); 25 May 2008; M. Schröder leg.; CMU • 5 ♂♂; Ergatikes; 37.721° N, 24.04° E; 18 May 2023; D. Ortiz and S. Pekár leg.; CMU • 1 ♂; Oulintzi; 37.946° N, 23.834° E; 18 May 2023; D. Ortiz and S. Pekár leg.; CMU • 3 ♂♂, 3 ♀♀; Ano Vakalopoulo; Fig. 53. Copulatory organs of Zodarion granulatum Kulczyński, 1908 (CMU). A–C, F. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. F. Detail of embolus, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 72 37.996° N, 23.922° E; 18 May 2023; D. Ortiz and S. Pekár leg.; CMU • 2 ♂♂, 1 ♀; Livadia; 38.448° N, 22.866° E; 24 May 2023; D. Ortiz and S. Pekár leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Fig. 54. Habitus of Zodarion hauseri Brignoli, 1984 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 73 Distribution North Macedonia, Bulgaria, Greece. Fig. 55. Copulatory organs of Zodarion hauseri Brignoli, 1984 (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 80 Remarks The overall body size, morphology, and male palp structure of Z. judaeorum closely resemble those observed in the genus Palaestina. However, Z. judaeorum lacks the coarse cheliceral dentition that serves as the distinguishing feature of Palaestina. Further investigation is necessary to clarify the phylogenetic relationship between Z. judaeorum and Palaestina. Distribution Israel. Fig. 61. Copulatory organs of Zodarion judaeorum Levy, 1992. A–C. Holotype, male palp (HUJ 14565). D. Paratype, female intact epigyne (HUJ 14564). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 81 Zodarion karpathos Bosmans, 2009 Figs 62, 105 Zodarion karpathos Bosmans, 2009: 253, figs 96–99. Diagnosis Refer to Bosmans (2009). Type material examined Holotype GREECE • ♂; Dodekanisa, Karpathos, Volada; 20 Sep. 2000; B. Knoflach & K. Thaler leg.; NMBEAR4144. Paratype GREECE • 1 ♂; same collection data as for holotype; NMBE-AR4144. Other material examined TURKEY • 2 ♂♂; Isparta, Eğridir, surrounding of Eğridir Lake; 37.921° N, 30.918° E; 946 m a.s.l.; 10 Jun. 2007; O. Seyyar leg.; CMU. Description For a detailed description of male, refer to Bosmans (2009). Distribution This species was previously recorded in Greece (World Spider Catalog 2025). This is the first report of this species from Turkey, and represents, therefore, the easternmost occurrence of this species. Zodarion killini Bosmans, 2009 Fig. 63 Zodarion killini Bosmans, 2009: 289, figs 164–165. Diagnosis Refer to Bosmans (2009). Type material examined (holotype not examined) Paratypes GREECE • 2 ♀♀; Peloponnisos, Korinthia, Oros Killini, Pheneos Pass; 880 m a.s.l.; 19 Sep. 1993; B. Knoflach leg.; NMBE-AR4158. Paratype GREECE • 1 ♂ (subadult); same collection data as for holotype; NMBE-AR4158. Description For a detailed description of female, refer to Bosmans (2009). Distribution Greece. European Journal of Taxonomy 1016: 1–139 (2025) 82 Fig. 62. Zodarion karpathos Bosmans, 2009, holotype, ♂ (NMBE-AR4144). A. Habitus, dorsal view. B. Habitus, ventral view. C. Palp, prolateral view. D. Palp, ventral view. E. Palp, retrolateral view. Scale bars: A–B = 1 mm; C–E = 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 83 Zodarion konradi Bosmans, 2009 Figs 64–65 Zodarion graecum Brignoli, 1984: 311, figs 37–39. Zodarion konradi Bosmans, 2009: 268, figs 132–133, 160–161. Diagnosis Refer to Bosmans (2009). Fig. 63. Zodarion killini Bosmans, 2009, paratype, ♀ (NMBE-AR4158). A. Habitus, dorsal view. B. Habitus, ventral view. C. Intact epigyne, ventral view. D. Vulva, ventral view. Scale bars: A–B = 1 mm; C–D = 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 84 Type material examined Holotype GREECE • ♂; Peloponnisos, Lakonia, Oros Taigetos, Aghios Elias; 2400 m a.s.l.; 26 Sep. 1985; B. Knoflach and K. Thaler leg.; NMBE-AR3668. Fig. 64. Habitus of Zodarion konradi Bosmans, 2009. A–B. Holotype, ♂ (NMBE-AR3668). C–D. Paratype, ♀ (NMBE-AR3668). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 85 Paratype GREECE • ♀; Oros Taigetos, Moni Panaghia Giatrissa; 1100 m a.s.l.; 30 Sep. 1991; B. Knoflach and K. Thaler leg.; NMBE-AR3668. Other material examined GREECE • 1 ♀; Corfu, Agios Georgios; 8 Apr. 2004; F. Šťáhlavský leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Fig. 65. Copulatory organs of Zodarion konradi Bosmans, 2009. A–C. Holotype, male palp (NMBEAR3668). D. Paratype, female intact epigyne (NMBE-AR3668). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 86 Remarks As the prosoma of the specimen was missing, it could not be photographed. Distribution North Macedonia, Greece. Zodarion kossamos Bosmans, 2009 Fig. 66 Zodarion kossamos Bosmans, 2009: 253, figs 100–103, 146–147. Zodarion kossamos – Lecigne 2011: 75, figs P120–125, S4–5. Fig. 66. Zodarion kossomas Bosmans, 2009 (CMU), ♂. A. Habitus, dorsal view. B. Habitus, ventral view. C. Palp, prolateral view. D. Palp, ventral view. E. Palp, retrolateral view. Scale bars: A–B = 1 mm; C–E = 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 87 Diagnosis Refer to Bosmans (2009). Material examined GREECE • 1 ♂; Kos, Kefalos; 36.744° N, 26.970° E; 100 m a.s.l.; 18 Aug. 2010; A. Exnerová leg.; CMU. TURKEY • 2 ♂♂; Antalya, Akseki, surrounding of Altınbeşik Cave; 37.034° N, 31.635° E; 471 m a.s.l.; 17 Apr. 2008; O. Seyyar leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Distribution Greece, Turkey. Zodarion levyi Shafaie & Pekár sp. nov. urn:lsid:zoobank.org:act:F5D64ED9-3627-4905-A456-134F362A4231 Figs 67–69, 106 Diagnosis This new species is most similar to Z. beroni by the small body size and asymmetrically bi-concave embolar tip (Fig. 68G vs Fig. 37F) but can be distinguished by the following characters: (1) a more extended prolateral process of median apophysis (vs elongated, cf. Fig. 68B, E and Fig. 37B); and (2) slightly convergent embolar arms (vs divergent arms, cf. Fig. 68G and Fig. 37F). Females of Z. levyi sp. nov. can be recognized from the other members of the thoni-group by an extended median septum (Fig. 69A–B), whereas it is less extended in Z. beroni (Fig. 37D), Z. granulatum (Fig. 53D), Z. namrun (Fig. 82A–B), Z. reticulatum (Fig. 90D), Z. rhodiense (Fig. 92D), and Z. thoni (Fig. 96D). Etymology This species is named after the late Gershom Levy, an Israeli arachnologist. Type material Holotype ISRAEL • ♂; Odem Forest, Golan Heights; 1 May 1996; J. Levy leg.; HUJINV-Ar 21303. Paratype ISRAEL • 1 ♀; same collection data as for holotype; HUJINV-Ar 21304. Other material examined ISRAEL • 1 ♀; same collection data as for holotype; CMU • 3 ♀♀; Odem Forest; 33.206° N, 35.736° E; 29 Mar. 2023; S. Pekár and V. Opatová leg.; CMU • 1 ♀; same collection data as for preceding; 4 May 2023; M. Řezáč leg.; CMU • 1 ♂, 1 ♀; Har Hermon; 33.288° N, 35.764° E; 5 May 2023; M. Řezáč leg.; CMU. SYRIA • 1 ♂; W Homs, Al-Zarah; 28 Mar. 2001; O. Řezáč leg.; CMU. European Journal of Taxonomy 1016: 1–139 (2025) 88 Description Male (holotype HUJINV-Ar 21303) Habitus (Fig. 67A–B). Carapace and chelicerae light brown; mouthparts basally light brown and apically pale; sternum yellow. All legs yellow. Palpal segments yellow. Abdomen dorsally black, with a shiny, black scutum, venter of abdomen and spinnerets slightly dirty yellow; PVS present. Fig. 67. Habitus of Zodarion levyi Shafaie & Pekár sp. nov. A–B. Holotype, ♂ (HUJINV-Ar 21303). C–D. Paratype, ♀ (HUJINV-Ar 21304). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 89 MeasureMents. Total length 2.44; carapace 1.25 long, 0.89 wide. Clypeus 0.14 high. Eye sizes and interdistances: AME 0.13, ALE = PME 0.09, PLE 0.08, AME–AME 0.05, AME–ALE 0.03, ALE–ALE 0.29, AME–PME 0.05, PME–PME 0.33, PME–PLE 0.05, PLE–PLE 0.26, ALE–PLE 0.1. Legs: I 3.55 (0.99, 0.39, 0.72, 0.89, 0.56), II 2.97 (0.82, 0.36, 0.52, 0.75, 0.52), III 2.89 (0.79, 0.33, 0.56, 0.75, 0.46), IV 4.31 (1.15, 0.36, 0.92, 1.22, 0.66). Palp: 0.89 (0.29, 0.13, 0.05, 0.42). P alP (Fig. 68A–G). RTA broad, with a claw-shaped extension. Median apophysis broad, rectangular, with an oval leaf-shaped PMA. Embolus short, with a bifurcated tip including two convergent arms. Fig. 68. Male palp of Zodarion levyi Shafaie & Pekár sp. nov., holotype (HUJINV-Ar 21303). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Prolateral view. E. Ventral view. F. Retrolateral view. G. Embolar tip, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 96 Fig. 74. Habitus of Zodarion lutipes (O. Pickard-Cambridge, 1872) (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 97 Fig. 75. Copulatory organs of Zodarion lutipes (O. Pickard-Cambridge, 1872) (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 98 Zodarion milani Shafaie & Pekár sp. nov. urn:lsid:zoobank.org:act:6E2ADB03-2768-49EC-8744-4B9488A6DCB7 Figs 76–77, 106 Diagnosis Males of this new species are similar to those of Z. rhodiense in body size and the coloration of the prosoma and legs, but they can be distinguished by their flattened embolar tip and the spine-like appendix (Fig. 77G vs Fig. 92F). Etymology The species is named after our colleague, Milan Řezáč, who collected the specimens. Type material Holotype SYRIA • ♂; Mashta Al-Helu; 3 Apr. 2001; M. Řezáč leg.; NMP P6A 7509. Other material examined SYRIA • 1 ♂; Al-Qadmus, Al-Kahf Castle; 5 Apr. 2001; M. Řezáč leg.; CMU. Description Male (holotype NMP P6A 7509) Habitus (Fig. 76A–B). Carapace and chelicerae brown; chelicerae with a longitudinal black stripe. Sternum and mouthparts brownish. Cx greyish, black distally; Fe black, with two pairs of longitudinal yellow stripes; Pa I, Ti I–III, Mt and Ta I–IV pale. Pa II–IV and Ti IV pale ventrally and dark laterally. Palpal femur black, with two pairs of orange stripes, patella and tibia pale ventrally and dark laterally, and cymbium brown, with yellow tip. Abdomen dorsum black, with an oval scutum, venter black except for brownish epiandrous; PVS present. Spinnerets black basally and pale apically. MeasureMents. Total length 4.55; carapace 2.34 long, 1.58 wide. Clypeus 0.26 high. Eye sizes and interdistances: AME 0.15, ALE = PME 0.1, PLE 0.08, AME–AME 0.06, AME–ALE 0.01, ALE–ALE 0.4, AME–PME 0.07, PME–PME 0.45, PME–PLE 0.1, PLE–PLE 0.27, ALE–PLE 0.2. Legs: I 7.77 (1.98, 0.72, 1.65, 2.14, 1.28), II 7 (1.91, 0.72, 1.38, 1.94, 1.05), III 7.01 (1.91, 0.69, 1.28, 2.14, 0.99), IV 9.64 (2.70, 0.75, 2.24, 2.8, 1.15). Palp: 2.37 (0.85, 0.39, 0.16, 0.95). Fig. 76. Habitus of Zodarion milani Shafaie & Pekár sp. nov., holotype, ♂ (NMP P6A 7509). A. Dorsal view. B. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 99 PalP (Fig. 77A–G). RTA broad, with a claw-shaped extension. Median apophysis rectangular, with a beak-shaped PMA, companied by a blunt prolateral extension. Embolus with a flattened embolar tip accompanied by a lateral spine-like appendix. Fig. 77. Male palp of Zodarion milani Shafaie & Pekár sp. nov., holotype (NMP P6A 7509). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Prolateral view. E. Ventral view. F. Retrolateral view. G. Embolar tip, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 100 Female Unknown. Size variation Total body length 4.55–4.57, carapace length/width 2.34–2.35/1.58–1 in males (N = 2). Distribution At present, known only from the type locality in Syria. Zodarion morosum Denis, 1935 Figs 78–79, 105 Zodarion morosum Denis, 1935b: 78, figs 22–24. Zodarion aegaeum Denis, 1935b: 81, fig. 25. Zodarion morosum – Karol 1969: 201, figs 1–11. — Deltshev 1987: 19, fig. 1.1–2. — Kovblyuk 2003: 177, figs 1–8, 14–16. — Bosmans 2009: 236, figs 20–22, 54–55. — Lecigne 2011: 75, figs P125– 128, S6–7. — Seropian et al. 2023: 293, figs 136–137. — Nae 2023: 56, figs 1–6. Diagnosis Refer to Bosmans (2009). Material examined GREECE • 2 ♀♀; E Olympos, Leptokarya, Olympic River; 9 Jun. 1996; J. Dolanský leg.; CMU • 1 ♀; same collection data as for preceding; 22 Jun. 2008; M. Schröder leg.; CMU • 1 ♂; Aladjagiola Wetland (N of Chrisoupoli); 41.01° N, 24.703° E; 18 Jun. 2004; S. Buchholz leg.; CMU • 1 ♀; same collection data as for preceding; 22 Jun. 2008; M. Schröder leg.; CMU • 4 ♂♂, 1 juv.; Lesvos; 18 May 2006; W. Nentwig leg.; CMU • 14 ♂♂, 2 ♀♀, 1 juv.; same collection data as for preceding; 15 Jun. 2006; W. Nentwig leg.; CMU. SYRIA • 1 ♀; Latakia; MNHN-AR2857. Description For a detailed description, refer to Bosmans (2009). Remarks Examination of the material from the Simon collection (MNHN-AR2857) revealed a female specimen of Z. morosum. Distribution This species was formerly recorded from North Macedonia, Bulgaria, Albania, Greece, Turkey, Ukraine, and Russia (World Spider Catalog 2025); this is the first report of this species from Syria. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 101 Fig. 78. Habitus of Zodarion morosum Denis, 1935 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 102 Fig. 79. Copulatory organs of Zodarion morosum Denis, 1935 (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 103 Zodarion namrun Dimitrov, 2024 Figs 80–82, 106 Zodarion namrun Dimitrov, 2024: 116, figs 15–17, 19–20. Fig. 80. Habitus of Zodarion namrun Dimitrov, 2024. A–B. ♂ (NMP P6A 7503). C–D. ♀ (NMP P6A 7505). A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 104 Diagnosis For male diagnosis, refer to Dimitrov (2024). The general epigynal shape of Z. namrun is similar to that of Z. thoni, except for the spermathecae strongly converging posteriorly (vs slightly converging posteriorly, cf. Fig. 82A–B and Fig. 96D) and the size of distal spermathecal ducts (smallest vs with the same size, cf. Fig. 82C and Fig. 96E). Material examined TURKEY • 2 ♂♂; Kahramanmaraş, Göksun, Püren Moutain Pass; 37.945° N, 36.511° E; 1518 m a.s.l.; 17 Jun. 2008; O. Seyyar leg.; NMP P6A 7503, NMP P6A 7504 • 1 ♀; same collection data as for preceding; NMP P6A 7505 • 1 ♂; same collection data as for preceding; CMU. Fig. 81. Male palp of Zodarion namrun Dimitrov, 2024 (NMP P6A 7503). A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Prolateral view. E. Ventral view. F. Retrolateral view. G. Embolar tip, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 105 Description For a detailed description of male, refer to Dimitrov (2024). Female (NMP P6A 7505) H abitus (Fig. 80C–D). Carapace distally yellowish-brown, anteriorly brown; clypeus and chelicerae brown; mouthparts yellow basally and pale apically; sternum greyish. All legs yellow except for entirely dark Fe I and II. Palpal segments yellow. Abdomen dorsum black, with no pattern; venter mostly dark except for a pair of pale, converging patches and a light area around the copulatory organ; PVS present. Spinnerets light grey basally, whitish apically. MeasureMents. Total length 5.01; carapace 2.21 long, 1.48 wide. Clypeus 0.33 high. Eye sizes and interdistances: AME 0.12, ALE 0.1, PME = PLE 0.08, AME–AME 0.07, AME–ALE 0.02, ALE–ALE 0.41, AME–PME 0.07, PME–PME 0.45, PME–PLE 0.1, PLE–PLE 0.27, ALE–PLE 0.21. Legs: I 6.68 (1.74, 0.66, 1.45, 1.81, 1.02), II 5.96 (1.61, 0.66, 1.12, 1.65, 0.92), III 6.21 (1.81, 0.69, 1.08, 1.81, 0.82), IV 8.25 (2.21, 0.77, 1.94, 2.44, 0.89). Palp: 1.87 (0.69, 0.33, 0.33, 0.52). ePigyne (Fig. 82A–D). Epigyne with strongly converging spermathecae; vulva comprising three globular, voluminous and convergent bodies, arranged by size from largest to smallest. Size variation Total body length 3.56–4.12, carapace length/width 1.8–1.84/1.3–1.42 in males (N = 3). Distribution At present, known only from Turkey. Fig. 82. Female copulatory organ of Zodarion namrun Dimitrov, 2024 (NMP P6A 7505). A. Intact epigyne, ventral view. B. Epigyne, ventral view. C. Vulva, ventral view. D. Vulva, ventral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 112 Diagnosis Refer to Bosmans (2009). Material examined GREECE • 1 ♂; Aladjagiola Wetland (N of Chrisoupoli); 41.007° N, 24.703° E; 8 May 2004; S. Buchholz leg.; CMU • 1 ♂; same collection data as for preceding; 8 July 2008; M. Schröder leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Distribution Bulgaria, Greece. Zodarion reticulatum Kulczyński, 1908 Figs 89–90, 105 Zodarion reticulatum Kulczyński, 1908: 61, pl. 2 fig. 16. Zodarion reticulatum – Levy 1992: 93. — Bosmans 2009: 280, figs 174–175, 182–183. — Lecigne & Henrard 2022: 37, fig. 5d, g. Diagnosis Refer to Bosmans (2009) and Lecigne & Henrard (2022). Material examined CYPRUS • 1 ♂, 1 ♀; Olympus Mount, Apliki; 34.941° N, 33.123° E; 720 m a.s.l.; 27 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU • 1 ♂ (subadult); Olympus Mount, Moutoulas Mountain; 34.977° N, 32.828° E; 800 m a.s.l.; 25 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU • 3 juv.; Lefka Dam; 35.063° N, 32.837° E; 288 m a.s.l.; 25 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU. GREECE • 1 ♀; Grevena, Kalamitsi; 15 Aug. 1999; CMU. TURKEY • 2 ♂♂, 1 ♀; Konakli, Macchie Hill, 10 km W of Alanya; 20 May 2005; J. Dolanský leg.; CMU • 2 ♂♂, 1 ♀; same collection data as for preceding; 19 May 2005; J. Dolanský leg.; CMU • 1 ♀; s. loc.; 21 Jun. 2003; CMU • 1 ♀; Erzincan, Çağlayan; 39.589° N, 39.705° E; 1541 m a.s.l.; 24 Jun. 2002; M. Řezáč leg.; CMU. Description For a detailed description, refer to Lecigne & Henrard (2022). Remarks As in Z. granulatum, males of Z. reticulatum possess longer PVS, arranged in a somewhat radiating pattern (Fig. 89B), whereas females exhibit shorter, more parallel PVS structures (Fig. 89D). Distribution Previous studies recorded Z. reticulatum in Cyprus and Greece (World Spider Catalog 2025). This is the first report of this species from Turkey and represents, therefore, the northwesternmost occurrence of this species across its distribution range. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 113 Fig. 89. Habitus of Zodarion reticulatum Kulczyński, 1908 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 114 Zodarion rhodiense Caporiacco, 1948 Figs 91–92, 105 Zodarion rhodiense Caporiacco, 1948: 47, fig. 5. Zodarion rhodiense – Brignoli 1984: 317, figs 50–53. — Dimitrov 2024: 113, figs 1–9. Zodarion rhodiense nigrifemur – Caporiacco 1948: 49. Fig. 90. Copulatory organs of Zodarion reticulatum Kulczyński, 1908 (CMU). A–C. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. F. Detail of embolus, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 115 Zodarion nigrifemur – Bosmans 2009: 279, figs 172–173, 180–181. — Lecigne & Henrard 2022: 33, figs 1a–d, 2a–g, 3a–e, 4a–h, 5a. — Seropian et al. 2023: 293, figs 138–140. — Dimitrov 2024: 113, figs 1–9. Zodarion thoni – Lecigne 2021: 34, fig. 55a–b. Diagnosis Refer to Lecigne & Henrard (2022) and Dimitrov (2024). Fig. 91. Habitus of Zodarion rhodiense Caporiacco, 1948 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 116 Material examined CYPRUS • 4 ♀♀, 5 juv.; Kyrenia Mountain range, Saint Hilarion Castle, Sysklipos (Akcicek); 35.309° N, 33.202° E; 329 m a.s.l.; 27 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU. GREECE • 2 ♀♀, 1 juv.; Lesvos; 39.220° N, 26.291° E; 967 m a.s.l.; 15 Jun. 2006; W. Nentwig leg.; CMU. SYRIA • 2 ♀♀; Hanna Area; 11 Apr. 2001; M. Řezáč leg.; CMU. TURKEY • 1 ♀; Isparta, Aksu, Dedegöl Mountain; 37.656° N, 31.291° E; 8 Jul. 2003; M. Řezáč leg.; CMU. Description For a detailed description, refer to Lecigne & Henrard (2022). Fig. 92. Copulatory organs of Zodarion rhodiense Caporiacco, 1948 (CMU). A–C, F. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. F. Detail of embolus, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 117 Distribution This species was previously reported from Georgia, Greece and Turkey; these are the first reports of this species from Cyprus and Syria, and represent, therefore, the southernmost occurrences of this species across its distribution range. Zodarion spinibarbis Wunderlich, 1973 Figs 93–94 Zodarion spinibarbis Wunderlich, 1973: 173, figs 4–10. Zodarion spinibarbis – Brignoli 1984: 319, fig. 45. Zodarion spinibarbe – Bosmans 2009: 244, figs 72–75, 136–137. — Coşar et al. 2022: 232, figs 22–32. — Coşar & Danışman 2024: 268, figs 23–24. Fig. 93. Habitus of Zodarion spinibarbis Wunderlich, 1973 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 118 Diagnosis Refer to Bosmans (2009). Material examined GREECE • 1 ♀; Kythira, Agia Pelagia; 36.325° N, 22.982° E; 383 m a.s.l.; 28 May 2003; J. Dolanský leg.; CMU. TURKEY • 1 ♂, 1 ♀; Kahramanmaraş, Andiran, Geben, Değirmendere; 37.889° N, 36.448° E; 1518 m a.s.l.; 15 May 2008; O. Seyyar leg.; CMU. Description For a detailed description, refer to Bosmans (2009). Distribution Greece and Turkey. Zodarion thoni Nosek, 1905 Figs 95–96, 105 Zodarion thoni Nosek, 1905: 128, fig. 10. Refer to World Spider Catalog 2025. Fig. 94. Copulatory organs of Zodarion spinibarbis Wunderlich, 1973 (CMU). A–C. Male palp, with rotated bulb. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 119 Diagnosis Refer to Bosmans (2009) and Lecigne & Henrard (2022). Material examined CYPRUS • 2 ♂♂, 1 ♀; Latchi, The Olympians Latchi Beach Villas; 35.041° N, 32.413° E; 7 m a.s.l.; 7–27 Aug. 2021; K.G. Mikhailov leg.; ZMMU. Fig. 95. Habitus of Zodarion thoni Nosek, 1905 (CMU). A–B. ♂. C–D. ♀. A. Dorsal view. B. Ventral view. C. Dorsal view. D. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 120 GREECE • 2 ♂♂; Kos, Kefalos; 36.744° N, 26.970° E; 100 m a.s.l.; 18 Aug. 2010; A. Exnerová leg.; CMU. IRAN • 1 ♂, 1 ♀; Urmia, Band; 37.461° N, 44.932° E; 1400 m a.s.l.; 29 Jun. 2022; S. Shafaie leg.; CMU. TURKEY • 2 ♀♀; Agri, Dogubeyazit; 39.547° N, 44.083° E; 1625 m a.s.l.; 16 May 1997; V. Bryja leg.; CMU • 1 ♂, 1 ♀; Tunceli, Munzur River; 38.957° N, 39.561° E; 22 Jun. 2002; J. Dolanský leg.; CMU • 1 ♂; Sivas, Demiryurt; 39.883° N, 37.6° E; 1542 m a.s.l.; 28 Jun. 2002; J. Dolanský leg.; CMU • 2 ♂♂, 1 ♀; Tokat, Pazar, Ballica Cave; 40.227° N, 36.301° E; 1085 m a.s.l.; 1 Jul. 2002; J. Dolanský leg.; CMU • 1 ♀; Konakli, Macchie Hill, 10 km W of Alanya; 36.589° N, 31.875° E; 800 m a.s.l.; 20 May 2005; J. Dolanský leg.; CMU • 5 ♂♂; Kayseri, Yahyali, near the Kapuzbaşı Waterfall; 26 May 2007; O. Seyyar leg.; CMU • 1 ♂, 1 ♀, 31 juv.; Cankiri, Ankara-Cankiri Highway; 40.389° N, 33.570° E; 689 m a.s.l.; 15 Sep. 2010; Y.M. Marusik leg.; ZMUT • 4 juv.; Bilecik, Mezit; 39.950° N, 29.701° E; 500 m a.s.l.; 26 Sep. 2010; Y.M. Marusik leg.; ZMUT. Fig. 96. Copulatory organs of Zodarion thoni Nosek, 1905 (CMU). A–C, F. Male palp. D–E. Female copulatory organ. A. Prolateral view. B. Ventral view. C. Retrolateral view. D. Intact epigyne, ventral view. E. Vulva, ventral view. F. Detail of embolus, ventral view. Scale bars: 0.2 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 121 Description For a detailed description, refer to Lecigne & Henrard (2022). Distribution Previous studies recorded this species in Eastern Europe, Cyprus, Turkey, the Caucasus, and Lebanon. The current study reported this species from Greece and Iran for the first time. Zodarion tunetiacum Strand, 1906 Figs 97–98, 106 Zodarium tunetiacum Strand, 1906: 669. Zodarium tunetiacum – Strand 1908: 86. Zodarion tunetiacum – Denis 1937: 17, pl. 3 fig. 21. Diagnosis This species can be separated from Z. cyrenaicum by the presence of a horizontally curly brace-shaped protuberance in the middle of epigyne (vs absent in Z. cyrenaicum; cf. Fig. 98A–B and Fig. 46D). Type material examined Neotype (here designated) EGYPT • ♀; Alexandria; MNHN-AR2854. Remarks Examination of material from the Simon collection (MNHN-AR2854) led to the identification of a female specimen belonging to Z. tunetiacum. Although this species has not been listed as a nomen Fig. 97. Habitus of Zodarion tunetiacum Strand, 1906. A–B. Neotype, ♀ (MNHN-AR2854). A. Dorsal view. B. Ventral view. Scale bars: 1 mm. European Journal of Taxonomy 1016: 1–139 (2025) 128 Other material examined CYPRUS • 1 ♂ (subadult), 2 juv.; Agios Georgios; 34.822° N, 32.897° E; 840 m a.s.l.; 29 Apr. 2022; S. Pekár, S. Shafaie, V. Šoltýsová and O. Michálek leg.; CMU. ISRAEL • 16 juv.; same collection data as for holotype; CMU • 7 juv.; Geshur; 32.816° N, 35.718° E; 30 Mar. 2023; S. Pekár leg.; CMU • 1 ♂, 8 juv.; Kadarim; 32.894° N, 35.478° E; 31 Mar. 2023; S. Pekár and V. Opatová leg.; CMU • 1 juv.; Har Heron; 32.996° N, 35.410° E; 3 May 2023; M. Řezáč leg.; CMU. Description Male (holotype NMP P6A 7510) Habitus (Fig. 103A–B). Carapace reddish-brown companied by a lighter, butterfly-like mark on behind of fovea; clypeus, chelicerae reddish-brown; labium and gnathocoxae brownish, with lighter tips. Sternum yellow at the middle and slightly darker at borders. Cx I and Fe I and II entirely dark, Cx II dark on lateral sides, Cx III and IV yellowish, Fe III and IV light brown proximally and darker distally, with a pair of longitudinal, light stripes. Pa and Ti of all legs yellow, with greyish laterals, other segments entirely yellow. Palpal segments yellow, with pale cymbium. Abdomen dorsum entirely dark, venter dark except for lighter parts around the epiandrous and PVS. Spinnerets pale, with dark basal part. M easureMents . Total length 4.49; carapace 2.16 long, 1.76 wide. Eye sizes and interdistances: AME 0.15, ALE 0.11, PME = PLE 0.08, AME–AME 0.1, AME–ALE 0.02, ALE–ALE 0.36, PME–PME 0.23, PLE–PLE 0.4, PME–PLE 0.03. Legs: I 9.59 (2.6, 0.8, 2.26, 2.53, 1.4), II 9.19 (2.53, 1, 2.13, 2.4, 1.13), III 8.38 (2.26, 0.73, 1.66, 2.6, 1.13), IV 10.06 (3, 1, 2.53, 3.33, 1.2). Palp: 3.26 (1, 0.46, 0.2, 1.6). PalP (Fig. 104A–C). Palp entirely similar to that of Z. lutipes but markedly larger (for a detailed description, refer to Levy 1992: cf. figs 67–69 and Bosmans 2009: cf. figs 184–185). Fig. 103. Habitus of Zodarion zonsteini Shafaie & Pekár sp. nov., holotype, ♂ (NMP P6A 7510). A. Dorsal view. B. Ventral view. Scale bars: 1 mm. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 129 Remarks Although no structural differences were detected in the copulatory organs of Z. zonsteini sp. nov. and Z. lutipes, the two representatives exhibit notable disparities in overall body and copulatory organ dimensions. Particularly, the carapace and cymbium of Z. zonsteini are 1.31 and 1.76 times longer, respectively, than those of Z. lutipes. Such divergence in absolute copulatory organ dimensions may lead to mechanical incompatibility between sexes, potentially impeding successful copulation (Ramos et al. 2005). For instance, the relatively large palpal organs of Z. zonsteini might not be effectively paired with the smaller, heavily sclerotized epigyne of Z. lutipes. Moreover, the two species differ in somatic morphology, with Z. lutipes exhibiting a distinct bicolored carapace and lighter legs compared to Z. zonsteini. Similar patterns of morphological divergence have previously been observed in Z. nitidum and Z. nicki (refer to Pekár et al. 2022 and corresponding comments in this study). Based on these consistent morphological differences, we treat Z. zonsteini and Z. lutipes as separate species. Nevertheless, further examination of additional material will be necessary to fully resolve issues of species delimitation and taxonomic consistency. Distribution At present, known only from Israel and Cyprus. Fig. 104. Male palp of Zodarion zonsteini Shafaie & Pekár sp. nov., holotype (NMP P6A 7510). A. Prolateral view. B. Ventral view. C. Retrolateral view. Scale bars: 0.2 mm. European Journal of Taxonomy 1016: 1–139 (2025) 130 Discussion Although several arachnologists have revised the zodariid species of the Middle East and Eastern Mediterranean (Levy 1992; Bosmans 1994, 1997, 2009; Zamani & Marusik 2021), our knowledge on the taxonomy of these spiders in this region remains fragmentary. The primary issue stems from the vast geographical area that hosts many species, and the relatively few arachnologists working in the region. Moreover, political instability in some countries further hampers progress in the research. Another significant issue lies in the overreliance on specific diagnostic characters. Early taxonomic studies in the 19 th and early 20 th centuries primarily concentrated on somatic characters of zodariids, whereas later research shifted its emphasis to copulatory traits. This transition initially led to a proliferation of recognized species, followed by a subsequent taxonomic reduction. Although the emphasis on copulatory organ morphology has proven reliable, an exclusive focus on these traits – at the expense of somatic traits and body dimensions – may hinder consensus on which characters should receive priority and what degree of variation is acceptable for defining species boundaries. Fig. 105. Distribution map showing new country records across the Middle East and Eastern Mediterranean, one sign may refer to several points, each number may refer to several localities. 1 = Acanthinozodium odem (Levy, 2007); 2 = A. tibesti Jocqué, 1991; 3 = Zodarion abantense Wunderlich, 1980; 4 = Z. granulatum Kulczyński, 1908; 5 = Z. karpathos Bosmans, 2009; 6 = Z. morosum Denis, 1935; 7 = Z. reticulatum Kulczyński, 1908; 8 = Z. rhodiense Caporiacco, 1948; 9 = Z. thoni Nosek, 1905. SHAFAIE S. et al., New data on Middle East and Eastern Mediterranean zodariids (Araneae) 131 Recent studies that questioned this long-held traditional approach was conducted by Pekár et al. (2022) and Ortiz et al. (2024). Previously, Z. nicki has been considered a synonym of Z. nitidum, with their distinct body coloration – blackish in Z. nicki and yellowish in Z. nitidum – interpreted as intraspecific variation. However, Pekár et al. (2022) and Ortiz et al. (2024) provided compelling evidence for the importance of somatic characters in distinguishing these two species of Zodarion. Although copulatory organs did not provide a clear differentiation, molecular, ecological, and behavioral data effectively separated the species, corroborating the distinction based on somatic traits and affirming the validity of both species. Another issue is the quality of available illustrations. Many species have been described primarily using copulatory organ drawings. While these illustrations are taxonomically valuable, they can arguably introduce inaccuracies and inconsistencies in species identification, as their interpretation is often subjective and influenced by the taxonomist’s perspective. In this study, we documented 50 zodariid representatives, providing detailed illustrations and photographs of both somatic and copulatory organs. Following the revised taxonomy presented here, the zodariid fauna of the Middle East and Eastern Mediterranean now includes the following genera, with the number of species known from each indicated in parentheses: Acanthinozodium (14), Cydrela Thorell, 1873 (1), Dusmadiores Jocqué, 1987 (1), Indozodion (1), Lachesana (11), Palaestina (3), Parazodarion (1), Pax Fig. 106. Distribution records across the Middle East and Eastern Mediterranean. 1 = Indozodion lindbergi (Roewer, 1960); 2 = Lachesana gavishae Shafaie & Pekár sp. nov.; 3 = L. lubinae Shafaie & Pekár sp. nov.; 4 = Parazodarion raddei (Simon, 1889); 5 = Pax akilae Shafaie & Pekár sp. nov.; 6 = Zodarion crewsae Coşar, Danışman & Kunt, 2022; 7 = Z. jakubi Shafaie & Pekár sp. nov.; 8 = Z. levyi Shafaie & Pekár sp. nov.; 9 = Z. milani Shafaie & Pekár sp. nov.; 10 = Z. namrun Dimitrov, 2024; 11 = Z. nicki Strand, 1914; 12 = Z. nitidum Audouin, 1826; 13 = Z. tunetiacum Strand, 1906; 14 = Z. zonsteini Shafaie & Pekár sp. nov. European Journal of Taxonomy 1016: 1–139 (2025) 132 (8), Trygetus (6), Zodariellum Andreeva & Tystshenko, 1968 (1), and Zodarion (75). The highest species richness is observed in Greece (43), Turkey (40), Isreal (26), and Iran (22) (Fig. 107). In conclusion, this study represents a thorough survey of the taxonomy of zodariid spiders in the Middle Eastern and East Mediterranean. To achieve a more precise estimate of the actual number of species, it is crucial to integrate comprehensive morphological documentation with data derived from multiple complementary approaches. Acknowledgments We wish to express our warmest thanks to C. Kropf (Switzerland) for providing us with the type specimens of Z. barbarae, Z. karpathos, Z. killini and Z. konradi; to W. Wins and T. Szűts (Hungary) for the type of Z. aculeatum; to P. Jäger and J. Altmann (Germany) for the type of Z. nicki; to Z. Simmons and R. Douglas (UK) for the types of Z. expers, Z. luctuosum and Z. lutipes; to E. Gavish-Regev (Israel) for the types of P. eremica, P. engediensis, P. palmonii, Z. geshur, Z. judaeorum and Z. odem; to C. Rolland and J. Pétillon (France) for organizing the first author’s stay in Paris and loaning important material; to D. Harms and N. Dupérré (Museum of Nature Hamburg, Germany) for preparing the figures of Z. parashi; to J. Mitáček (Czech Republic), O. Mirshamsi (Iran), P. Pantini (Italy), Y.M. Marusik and K.G. Mikhailov (Russia), and M. Chatzaki (Greece) who kindly sent us the material deposited in their collections. We are particularly grateful to R. Jocqué (Belgium), A. Henrard and an anonymous reviewer for constructive comments on an earlier version of the manuscript. We are also grateful to S. Korenko, E. Líznarová, O. Michálek, D. Ortiz, L. Sentenská, and V. Šoltýsová for their kind help in the field; and to M. Řezáč, F. Šťáhlavský, J. Dolanský, E. Gavish, Y. Lubin, G. Levy, and A. Exnerová for providing their material. 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