Zaitunia Lehtinen, 1967

Sergei Zonstein & Yuri M. Marusik, 2016, A revision of the spider genus Zaitunia (Araneae, Filistatidae), European Journal of Taxonomy 214, pp. 1-97 : 6-11

Genus Zaitunia Lehtinen, 1967

Zaitunia Lehtinen, 1967: 275 .

Zaitunia – Zonstein 2009b: 126. — Fomichev & Marusik 2013: 85. — Zonstein et al. 2013: 69 . — Marusik & Zamani 2015: 129 .

Type species

Filistata schmitzi Kulczyński, 1911 , by monotypy.

Diagnosis

Zaitunia differs from other filistatine genera by a distinctly domed carapace lacking thoracic fovea, with a broadly rounded subvertical clypeus and by a shortened labium that is wider than long (in Filistata and Kukulcania Lehtinen, 1967 the clypeus is subhorizontal with a lateral constriction, without thickened setae, and the labium is distinctly longer than wide). It differs from Sahastata Benoit, 1968 by a uniseriate calamistrum (consisting in the latter genus of three setae rows) and from Microfilistata Zonstein, 1990 by spinose tarsi in females and pseudosegmented tarsi in males (vs aspinose and integral, respectively). Zaitunia can be recognized due to a gap-setose calamistrum (as shown in Figs 2 C–E), a short and wide male palpal tibia (which is longer and more slender in other members of the subfamily; only in Zaitunia is it noticeably wider than the palpal femur) and a characteristic dense comb on the clypeus, composed of stout reclined bristles and much more developed (especially in males) than in other genera of the Filistatinae.

Description

SIZE. Small to medium-sized filistatid spiders with body length 2.5–8.0 mm.

COLOUR AND PATTERN. Colour varying from pale yellowish-white to dark brown; carapace with more or less darkened eye tubercle, lateral margins, and foveal area. Pattern of carapace lacking in pale specimens. Legs unicolor or with dark, broad annulations. Abdomen with or without dorsal pattern composed by longitudinal median dark stripe or band and in some species with transverse stripes. Venter of abdomen in most species without pattern, book lungs usually paler than other parts, genital area and area in front of spinnerets darkened in some species.

PROSOMA. Carapace distinctly domed, broad oval, widely rounded anteriorly, covered with sparse thick bristles ( Figs 1 A–B, 42). Some bristles, usually one pair very strong and long (about ½ of carapace width). Thoracic fovea opened posteriorly and appears as shallow pit or indistinct. Cephalic area flattened, usually with median row of strong proclined bristles. Clypeus relatively short and subvertical, with ± developed comb of stout reclined bristles ( Fig. 1 C). Eye tubercle low. ALE> PLE ≈ PME> AME. Median ocular quadrangle wide trapezoidal. Chelicerae small, subvertical; cheliceral furrow and fang very short; cheliceral lamina well-developed. Sternum subcircular, sigillae indistinct. Labium slightly wider than long. Maxilla trapezoidal, with lateral corner about 90°. Palps in females short and thick, with short dentate claw ( Fig. 1 E).

LEGS. Leg formula 1423 (most species) or 4123. Leg spination very uniform in all species, but as a rule, males have more spines than conspecific females; additionally, these spines in males are usually longer.

All femora with one basodorsal spine and 1–2 smaller spines in pro- and retrodistal position. Patellae unarmed. Tibiae with 2–6 ventral spines (can be absent on tibiae I–II in females). Metatarsi usually long and slender, with 6–9 ventral spines, but in males of Z. psammodroma sp. nov. metatarsi shortened, dilated and covered with numerous ventrodistal spines. Tibiae and metatarsi with few (1–3) inclined trichobothria, which are 1.2–2 times shorter than diameter of the segment. Trichobotrial bases low and inclined, smooth outside and densely rifled inside ( Fig. 2 H–I). Tarsi ventrally with few small spines, and with short and dense bristles. Male tarsi curved and pseudosegmented. Tarsal organ with wide inclined opening ( Fig. 2 F–G, J). Calamistrum short, present in females only, formed by one composite row of a few thick, curved and flattened setae on a raised keel (crest); median part of calamistrum without setae ( Fig. 2 A–E). Paired tarsal claw narrow and slightly curved, with row of 8–15 long, dense teeth ( Fig. 1 H). Unpaired claw curved, with 5–8 dense teeth ( Fig. 1 I).

ABDOMEN. Abdomen elongate oval, overhangs posterior part of carapace. Spinnerets shifted anteriorly. Spinneret group set relatively far from posterior edge of abdomen. Cribellum small, bipartite trapezoidal, present in both sexes ( Figs 3 A, D, 4D) but reduced in size and lacking functional spigots in males. ALS and PLS subequal in size ( Figs 3 A, C, 4C); PMS much smaller, with two probably paracribellar spigots ( Fig. 4 F, I). Tracheal spiracle wide and located in posterior ⅓ of abdomen between epigastral furrow and spinnerets ( Fig. 3 A–C). Tracheae thin and short (about as long as spiracle width). Two pairs of tracheal stems: median branches longer and inclined sideward, lateral branches much shorter ( Figs 3 B, 33E–F). As in the prosoma and legs, abdomen covered with ciliate hairs only, plumose hairs absent.

MALE PALP. Relatively short compared to other filistatines, although 1.3‒1.8 longer than carapace. The shortest palp in comparison to the carapace is in the type species (palp/carapace length = 1.3). Femur unmodified, cylindrical; equal in length to or longer than tibia. Patella unmodified, short, about one third of femur length. Tibia long and swollen, 1.5–2 times thicker than femur; ventromedially with patch of dense, suberect setae.

CYMBIUM. Conical at base, and the rest is cylindrical. Only Z. logunovi sp. nov. has a conical dorsal outgrowth in the terminal one-third ( Fig. 1 K–L). Tip of cymbium without dorsoapical excavation, its margin varies from strongly slanting (some specimens of Z. schmitzi , Fig. 7 A–D) to almost straight (perpendicular to axis of cymbium). Dorsum of cymbium with brush of dense, long suberect hairs; some hairs as long as cymbium. One species, Z. logunovi sp. nov., with set of thick hairs along ventral and lateral edges of cymbium ( Fig. 1 J). Terminal part of dorsum hairless, semitransparent.

BULB. Ranges from conical (most of the species) to globular ( Z. logunovi sp. nov., Z. minoica sp. nov.) and subcylindrical ( Z. schmitzi (Kulczyński, 1911)) , basal part embedded in cymbium. Length of bulb + embolic part varies from long (longer than cymbium) to short (shorter than cymbium). Spermophore forms 3‒4 coils, coils not spaced; spermophore ranges from very thick and occupying the whole bulb ( Z. schmitzi ) to moderately thin.

EMBOLUS (embolic part of the bulb). Without distinct separation from the tegulum. We recognize the neck of the embolus (Ne = part of embolic base wider than spermophore) and the embolus “proper” (Em = diameter equal to diameter of spermophore). Embolus can be distinguished from the neck by its dark colour, much darker then rest of the bulb. Embolic part of bulb variable in size and shape, from very short, lacking neck ( Z. logunovi sp. nov., Z. schmitzi ) to long, longer than bulb and as long as cymbium ( Z. minoica sp. nov.). Embolic part from straight to screw-shaped ( Z. logunovi sp. nov., Z. schmitzi ). Embolic neck in most species with distinct ventral keel or without ( Z. minoica sp. nov.). Shape of keel in some cases is species-specific. Tip of embolic part bent ventrally or screw-shaped.

ENDOGYNE (VULVA). Adult females can be hardly recognized. Compared to subadult females, they have a slightly swollen edge of the genital area. Receptacles are enveloped in a trapezoidal or semicircular fold formed by the epigastric integument (making it difficult to observe and figure correctly). Number of receptacles varies from one to two pairs. In species with four receptacles, all receptacles are usually isolated from each other, but in some cases median and lateral receptacles are connected with each other and can appear as one receptacle with two heads. Shape of receptacles varies and can be cylindrical ( Figs 9 D–F, 21D–G, 39A–D), globular ( Figs 7 E–K, 10F–G, 11F–G, 27H, 30C–E), subconical ( Fig. 25 ), or dumbbell- or boomerang-shaped ( Z. afghana ( Roewer, 1962) , Z. brignoliana sp. nov.). Cylindrical receptacles can be bent or straight. Straight cylindrical and subconical receptacles have a corrugated transparent stem. Median (or mesal) receptacles are usually longer and wider than lateral, although in some species the opposite is true. Receptacles bear glandular pores (not recognized in Z. persica Brignoli, 1982 ). Each pore equipped with a kind of cilia. Pores can be evenly spread along receptacles (in species lacking a corrugated stem) or can be concentrated on the heads. In some species, females are almost indistinguishable from each other by the structure of the endogyne (e.g., Z. ferghanensis sp. nov., Z. maracandica ( Charitonov, 1946) , Z. wunderlichi sp. nov.), whereas males belonging to the same species are easily differentiated (cf. Figs 33 A–B, D, 34E–J, 38E–I and 32C–G, 35A–D, 37A–C, respectively).

Composition

24 species: Z. afghana ( Roewer, 1962) (Afghanistan) , Z. akhanii Marusik & Zamani, 2015 (Iran) , Z. alexandri Brignoli, 1982 (Iran) , Z. annulipes ( Kulczyński, 1908) (Cyprus) , Z. beshkentica ( Andreeva & Tyshchenko, 1969) (Uzbekistan, Tajikistan), Z. brignoliana sp. nov. (Iran), Z. ferghanensis sp. nov. (Kyrgyzstan, Uzbekistan), Z. feti sp. nov. (Turkmenistan), Z. halepensis sp. nov. (Syria), Z. huberi sp. nov. (Afghanistan), Z. inderensis Ponomarev, 2005 (Kazakhstan) , Z. kunti sp. nov. (Turkey, Cyprus), Z. logunovi sp. nov. (Kazakhstan, Kyrgyzstan), Z. maracandica ( Charitonov, 1946) (Uzbekistan, Kazakhstan), Z. martynovae ( Andreeva & Tyshchenko, 1969) (Turkmenistan, Tajikistan), Z. medica Brignoli, 1982 (Iran) , Z. minoica sp. nov. (Greece), Z. minuta sp. nov. (Uzbekistan), Z. persica Brignoli, 1982 (Iran) , Z. psammodroma sp. nov. (Turkmenistan), Z. schmitzi ( Kulczyński, 1911) (Egypt, Israel), Z. spinimana sp. nov. (Kazakhstan, Turkmenistan), Z. wunderlichi sp. nov. (Kyrgyzstan) and Z. zonsteini Fomichev & Marusik, 1969 (Kazakhstan) .

Species grouping

To assist with identifications, the species treated here are assigned to seven informal species groups. These assignments are preliminary, considering that males in many species are unknown, and not based on a monophyletic grouping, though some of the groups may actually reflect phylogenetic relationships.

Distribution

The genus is known from Crete to Eastern Kazakhstan, south to the Sinai Peninsula in Egypt, Fars and Kerman Provinces of Iran and Southeastern Afghanistan, and north to Northwestern and Northeastern Kazakhstan ( Figs 46–49 ).

Ecology

Most species inhabit more or less arid habitats: deserts, semi-deserts, steppes, maquis or deciduous shrubland, where spiders may occur under rocks and in crevices of clay or rocky escarps where they build small tubular webs (see Fig. 50 ). Adult males were collected manually and by pitfall traps, generally during late spring or summer. Only one species, Z. wunderlichi sp. nov., was found in a humid dense broad-leaved and mixed mountain forest dominated by walnut ( Juglans regia ). Concluding from the known data, Zaitunia avoid the true sandy deserts, with the only exception being Z. psammodroma sp. nov., known currently only from males. Contrary to all other congeners, males of Z. psammodroma sp. nov. possess short and distally dilated metatarsi IV with numerous ventrodistal spines, a clear adaptation to moving on loose sandy substrate. Zaitunia species are found from the seashore to altitudes over 2000 m ( Z. beshkentica ).

Figures

Fig. 1. Morphology of Zaitunia. A – C. Z. martynovae (Andreeva & Tyshchenko, 1969), subadult ♀ from Gandzhina. D – I. Same, adult ♀. J – L. Z. logunovi sp. nov., paratype, ♂. — A, C. Anterior part of prosoma, dorsal and lateral. B. Prosoma, dorsal. D. Leg IV, prolateral. E. Palp, prolateral. F. Cuticle and setae covering venter of abdomen. G. Leg surface showing large and fine (possibly chemosensory) setae. H – I. Tarsal claws of leg IV, prolateral. J – L. Terminal part of palp, prolateral, dorsal and ventroretrolateral. Abbreviations: see Material and methods.

Fig. 2. Morphology of Zaitunia leg. A – K. Z. martynovae (Andreeva & Tyshchenko, 1969), ♀ from Gandzhina. L. Z. logunovi sp. nov., paratype, ♂. — A. Calamistrum, dorsal. B – C. Same, lateral. D. Calamistral area with removed setae, dorsal. E. Setae of calamistrum, lateral. F – G. Tarsal organ. H – I. Base of trichobothria. J. Part of dorsal leg surface showing two slit organs. K. Long slit organ. L. Part of male tarsus IV, prolateral, showing pseudosegmentation. Abbreviations: see Material and methods.

Fig. 3. Morphology of Zaitunia spp. A – B. Z. logunovi sp. nov., paratype, ♂. C. Z. schmitzi (Kulczyński, 1911), ♀ from Jerusalem area. D, H. Z. martynovae (Andreeva & Tyshchenko, 1969), ♀ from Gandzhina. E, G. Z. wunderlichi sp. nov., paratype, ♂. F. Z. beshkentica (Andreeva & Tyshchenko, 1969), ♂ from Khurshedi. — A – D. Abdomen, ventral. E – F. Terminal part of palp, retrolateral. G. Same, dorsal. H. Genital area of subadult female. Abbreviations: see Material and methods. Scale bars: A – B, D, G = 0.2 mm; C, H = 0.1 mm.

Fig. 4. Spinneret morphology of Zaitunia martynovae (Andreeva & Tyshchenko, 1969), adult ♀ from Gandzhina. A – B. Spinnerets and tracheal spiracle. C. Right posterior lateral spinneret. D. Cribellum. E. Left anterior lateral spinneret. F. Left posterior median and lateral spinnerets. G. Right posterior lateral spinneret, frontal. H. Right anterior lateral spinneret. I. Posterior median spinnerets. Abbreviations: see Material and methods. Scale bars: A – B = 100 μm; C – I = 10 μm.

Fig. 7. Copulatory organs of Zaitunia schmitzi (Kulczyński, 1911), ♂ (A – D) and ♀ (E – K) from Jerusalem area. — A – B. Palp, retrolateral and prolateral. C – D. Same, ventro-prolateral and ventroretrolateral. E, G, K. Endogyne, anterior. F. Same, posterior. H, J. Receptacles, dorsal and anterior. I. Endogyne, dorsal.

Fig. 9. Copulatory organs of Zaitunia annulipes (Kulczyński, 1908), ♂ from Paramali (A – C) and ♀ from Germasogenea (D – F). — A – C. Palp, prolateral, ventro-retrolateral and retrolateral. D. Endogyne, anterior. E – F. Same, dorsal. Scale bars: A – C = 0.2 mm; D – F = 0.1 mm.

Fig. 25. Palp and somatic characters of Zaitunia martynovae (Andreeva & Tyshchenko, 1969), ♂ from Gandzhina. A, D. Palp, retro- and prolateral. B – C. Terminal part of palp, retro-ventral and prolateral. E. Cephalic part of carapace, dorsal. F. Abdomen, ventral.

Fig. 33. Somatic characters and copulatory organs of Zaitunia wunderlichi sp. nov., paratype, ♀ (A – B, D), Z. beshkentica (Andreeva & Tyshchenko, 1969), ♀ from Babatagh Mts (C), Z. logunovi sp. nov., paratype, ♂ (E, G) and Z. schmitzi (Kulczyński, 1911), ♀ from Jerusalem area (F). — A – D. Endogyne, dorsal (A, C – D) and ventral (B). E – F. Posterior respiratory system, dorsal. G. Close-up view of male tarsus IV, prolateral, showing pseudosegmentation. Scale bars: A – B, D – G = 0.1 mm; C = 0.2 mm.

Fig. 46. Distribution of Zaitunia schmitzi and Zaitunia annulipes species groups.

Fig. 47. Distribution of Zaitunia persica species group.

Fig. 48. Distribution of Zaitunia logunovi and Zaitunia martynovae species groups.

Fig. 49. Distribution of Zaitunia beshkentica and Zaitunia maracandica species groups.

Fig. 50. Live specimens of Zaitunia and habitats of Z. martynovae (Andreeva & Tyshchenko, 1969) in Southern Tajikistan. A. ♀ of Z. schmitzi (Kulczyński, 1911) on the turned stone bottom (Israel, Shoham, image courtesy of Amir Weinstein). B. Surroundings of Gandzhina: the spiders occur under the stones. C. Pyandzh-Karatau Mts, one of the co-authors (YM) near the clay escarps inhabited by filistatids. D. Foothills of the salt dome Khodzha-Mumin: the spiders inhabit crevices and cavities in the rock outcrops. E. One of the figured escarps, a close-up photo showing the twin entrance leading to the inhabited cavity. F. Captured ♀ of Z. martynovae (Andreeva & Tyshchenko, 1969) weaving a web in the cover of a collecting tube.