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 .

PALESTINE • 3 ♀♀; Jericho or Bethlehem; MNHN-AR2849 .

Redescription

Male (HUJINV-Ar 21301)

HABITUS (Fig. 29A–B). Carapace dark brown; chelicerae and mouthparts brown; sternum light brown, with dark spots radiating towards the center. Legs entirely yellow except for the first pair of Fe, Pa and Ti laterally greyish. Palp segments grey to yellow. Abdomen dorsum black, venter apically pale and turned to dark brown distally; PVS present. Spinnerets black basally and pale apically.

MEASUREMENTS. Total length 4; carapace 2 long, 1.5 wide. Clypeus 0.1 high. Eye sizes and interdistances: AME 0.1, ALE= PME 0.06, PLE 0.04, AME–AME 0.06, AME–ALE 0.02, ALE–ALE 0.27, AME–PME 0.06, PME–PME 0.41, PME–PLE 0.06, PLE–PLE 0.23, ALE–PLE 0.1. Legs: I 7.52 (1.69, 0.64, 1.63, 2.03, 1.52), II 6.64 (1.59, 0.56, 1.53, 1.86, 1.1), III 6.8 (1.63, 0.87, 1.39, 1.85, 1.06), IV 8.14 (2.23, 0.64, 1.66, 2.37, 1.24). Palp: 1.28 (0.52, 0.25, 0.16, 0.35).

PALP (Fig. 30A–F). For details, refer to Levy (1992).

Female (HUJINV-Ar 21302)

HABITUS (Fig. 29C–D). Carapace, chelicerae and abdomen similar to those in male except abdomen venter slightly lighter. Mouthparts lighter than those in male. Fe I and II dark brown, with longitudinal, parallel lighter stripes; Fe III and IV darkened on the second half. Ti of all legs light, with dark lateral sides. Pa, Mt and Ta of all legs brown. Spinnerets brown basally and pale apically; PVS present.

MEASUREMENTS. Total length 6.8; carapace 2.59 long, 1.85 wide. Clypeus 0.18 high. Eye sizes and interdistances: AME 0.1, ALE= PME =PLE 0.06, AME–AME 0.08, AME–ALE 0.04, ALE–ALE 0.4, AME–PME 0.08, PME–PME 0.42, PME–PLE 0.06, PLE–PLE 0.24, ALE–PLE 0.2. Legs: I 5.85 (1.41, 0.54, 1.61, 1.29, 1), II 5.37 (1.31, 1.15, 0.54, 1.47, 0.9), III 5.03 (1.23, 0.52, 1.1, 1.47, 0.71), IV 6.59 (1.69, 0.77, 1.4, 1.86, 0.87). Palp: 1.52 (0.54, 0.33, 0.19, 0.46).

EPIGYNE (Fig. 31A–F). For details, refer to Levy (1992).

Size variation

Total body length 2.5–4.4, carapace length/width 1.8–2/ 1.48–1.6 in males (N= 6). Total body length 4.6–6.8, carapace length/width 2.1–2.6/ 1.6–1.7 in females (N=6). For the carapace length/width ratio, refer to Fig. 87.

Remarks

Although taxonomic studies on the genus Zodarion have a long history, little is known about its type species, Z. nitidum . In fact, Z. nitidum was not properly illustrated until Denis prepared two distinct epigyne drawings in separate manuscripts, each claiming to be the first illustration of Z. nitidum ’s epigyne (Denis 1937, 1962). The reason for Denis’ failure to address this inconsistency in his publications from 1937 to 1962 remains unclear (World Spider Catalog 2025).

Thirty years later, Levy (1992) acknowledged that the types of Z. nitidum – Enyo nitida (♀) and Enyo longipes (♂) – were presumably lost. However, he examined the holotypes of Zodarium nicki Strand, 1914 and Zodarion cambridgei Denis, 1938, along with a large number of newly collected specimens from various localities across Israel, Jordan, and Sinai. Based on the observed similarities in copulatory structures, Levy assigned all of these specimens to Z. nitidum . Despite Levy being the first to provide detailed descriptions and illustrations for both sexes within the Z. nitidum complex, his conclusions were based solely on the newly collected material. Consequently, he neither compared these specimens with the original (now lost) types nor provided illustrations that might serve as proxies for future taxonomic assessments.

Ultimately, all morphologically variable specimens were subsumed under the name Z. nitidum . However, recent comprehensive studies on the biology and ecology of this species complex (Pekár et al. 2022) combined with molecular phylogenomic analyses (Ortiz et al. 2024) have revealed that the complex consists of two distinct, diagnosable species, now recognized as Z. nitidum and Z. nicki .

Examination of the material from the Simon collection, who frequently combined specimens of different species or from different localities in a single tube, demonstrated that six female individuals, preserved in three different tubes (MNHN-AR2847: 2 ♀♀; MNHN-AR2849: 3 ♀♀; MNHN-AR2854: 1 ♀), had been misidentified and are, in fact, assignable to Z. nitidum . It should be noted that the MNHN-AR2854 tube contains material from multiple species, including the neotypes of Z. tunetiacum and Z. nitidum . The epigyne of the single Z. nitidum specimen from this tube closely corresponds with the illustrations provided by Denis (1937) (Fig. 31E cf. Fig. 31F). Levy (1992) erroneously lumped specimens of Z. nitidum and Z. nicki under a single taxon, as he found no discernible differences in their copulatory organ structures.

Levy (1992) also pointed out that earlier arachnologists had merged E. nitida (♀) and E. longipes (♂) – both described by Audouin (1826) – and subsequently treated them as Z. nitidum . However, whether these original associations were taxonomically accurate remains uncertain.

Distribution

North Africa, Middle East.