Filistata betarif, Magalhaes & Aharon & Ganem & Gavish-Regev, 2022

Magalhaes, Ivan L. F., Aharon, Shlomi, Ganem, Zeana & Gavish-Regev, Efrat, 2022, A new semi-cryptic Filistata from caves in the Levant with comments on the limits of Filistata insidiatrix (Forsskål, 1775) (Arachnida: Araneae: Filistatidae), European Journal of Taxonomy 831, pp. 149-174 : 161-167

Filistata betarif sp. nov.

urn:lsid:zoobank.org:act:86B4AAD0-D69E-4C79-BF3F-DE5E7DC0EDB1

Figs 1 , 3E–F, H , 5A, C, E , 7–9

Filistata IFMsp142 – Magalhaes & Ramírez in press: 11, 13, 16.

Diagnosis

Males are similar to those of F. insidiatrix and F. albens by having a pedipalp with elongate femur and tibia, and by a subconical copulatory bulb with terminally hooked embolus ( Fig. 8A–B ; see also Zonstein & Marusik 2019: fig. 27). They differ from both species by the conspicuous embolic keel ( Fig. 8F (arrow), 5A, k) (inconspicuous keel in F. insidiatrix and F. albens ; Figs 5B , 10O ). Females have a pair of large spermathecae that appear undivided ( Fig. 9 ); they fall within the morphological variation observed for F. insidiatrix ( Fig. 3C–D ; Zonstein & Marusik 2019: fig. 17d–h) and thus females are not diagnosable through morphology only; they can be diagnosed using DNA barcoding of the COI gene.

Etymology

The specific epithet is a noun in apposition and refers to Bet ‘Arif, the name of the ‘ wadi ’ (dry riverbed) in which the two caves where the species was collected (Tinshemet and Oah) are located.

Type material examined

Holotype ISRAEL • 1 ♂; Judean hills , near Shoham, Hadom Shomeron Nature Reserve ; Bet ‘Arif wadi , Tinshemet cave , inside cave ; 31.99938° N, 34.96813° E; 122 m. a.s.l.; 20 Feb. 2020; I.L.F. Magalhaes, E. Gavish- Regev and S. Aharon leg.; HUJ-INVAr 21058 .

Paratypes ISRAEL • 1 ♀; Judean hills , near Shoham, Hadom Shomeron Nature Reserve ; Bet ‘Arif wadi , Oah cave , inside cave ; 32.0053° N, 34.9722° E; 25 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21055 • 2 ♀♀; same locality as for preceding; 9 Mar. 2014; S. Aharon and E. Gavish-Regev leg.; HUJ- INVAr 21059 • 1 ♂, 6 ♀♀; Judean hills , near Shoham, Hadom Shomeron Nature Reserve , Bet ‘Arif wadi , Tinshemet cave , inside cave ; 31.99938° N, 34.96813° E, 122 m. a.s.l.; 20 Feb. 2020; I.L.F. Magalhaes, E. Gavish-Regev and S. Aharon leg.; MACN-Ar 41222 • 2 ♂♂, 3 ♀♀, 2 imm.; same collection data as for preceding; HUJ-INVAr 20275 • 4 ♀♀, 1 imm.; same collection data as for preceding; MACN-Ar 41791 • 1 ♂; same collection data as for preceding; MACN-Ar 41793 • 2 imm.; same locality as for preceding, Mediterranean scrubland , outside the cave, under stones; 20 Feb. 2020; I.L.F. Magalhaes, E. Gavish-Regev and S. Aharon leg.; MACN-Ar 41829 • 1 ♀; same locality as for preceding; Mediterranean scrubland , outside the cave; 31.9994° N, 34.9681° E; 25 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21056 • 7 ♀♀; same locality as for preceding; inside cave; 31.99938° N, 34.96813° E; 25 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 20301 • 1 ♀; same collection data as for preceding; HUJ-INVAr 21057 • 3 imm; same locality as for preceding; 9 Mar. 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21060 .

Additional material examined

None.

Description

Male (holotype, HUJ-INVAr 21058)

Coloration yellowish cream except where noted. Carapace lined with black pigment, with marbled brown clypeal markings and brown, V-shaped median pattern. Legs I–III with three brown ventral spots. Abdomen brownish gray. Anterior margin of the carapace unmodified, eye apodemes absent. Sternum subrounded, sigilla not visible. Total length 5.25. Carapace length 2.38, width 1.88. Clypeus length 0.28. Eye diameters and interdistances: AME 0.14, PME 0.15, ALE 0.20, PLE 0.18, AME–AME 0.06, PME– PME 0.14. Sternum length 1.26, width 1.05. Pedipalp: femur length 2.34, height 0.30, tibia length 2.10, height 0.34. Leg I: 14.88 (3.82, 1.09, 3.94, 3.91, 2.12). II: 11.29 (3.05, 0.94, 2.78, 3.17, 1.35). III: 10.23 (2.70, 0.84, 2.52, 2.91, 1.26). IV: 12.52 (3.57, 0.90, 3.28, 3.44, 1.33). Abdomen: length 2.97, width 1.72. Pedipalp macrosetae on ventral surface of femur. Leg macrosetae: fe I d1-0-0, dp0-0-2, ti I v0-2-2-2, mt I v2-2-2-4, fe II d1-1-0, dp0-0-2, ti II v0-2-2, mt II v2-2-4, fe III d1-0-0, dp0-0-1, dr0-0-2, ti III d1-2-2, v1-1, mt III dr0-0-1, v1-2-2-4, fe IV d1-1-0, dr0-0-2, ti IV v1-1, mt IV v1-1-2-4, all tarsi with ventral macrosetae. Pedipalp as in diagnosis ( Figs 5A, C, E , 8A–G ). State of the specimen: good, right leg I removed for molecular analysis, left pedipalp dissected.

Female (paratype, MACN-Ar 41222)

Coloration as in male. Eye apodemes absent. Sternum subrounded, with one pair of posterior sigilla. Total length 7.76. Carapace length 3.17, width 2.51. Clypeus length 0.40. Eye diameters and interdistances: AME 0.14, PME 0.19, ALE 0.27, PLE 0.23, AME–AME 0.04, PME–PME 0.23. Sternum length 1.64, width 1.50. Pedipalp: femur length 1.70, height 0.67, tibia length 1.06, height 0.65. Leg I: 13.60 (3.66, 1.37, 3.49, 3.23, 1.85). II: 9.66 (2.84, 1.13, 2.26, 2.16, 1.27). III: 7.82 (2.34, 1.03, 1.66, 1.64, 1.15). IV: 10.54 (3.30, 1.25, 2.64, 2.27, 1.08). Abdomen: length 4.67, width 3.08. Palpal macrosetae on ventral surface of tibia and tarsus. Leg macrosetae: fe I d1-0-0, ti I v0-2-0, mt I v2-2-2-4, fe II d1-0-0, ti II v0- 2-0, mt II v2-2-2-4, fe III d1-0-0, ti III d0-2-0, v0-1-2, mt III dr1-1-1, v1-2-2-4, fe IV d1-0-0, ti IV v1-1, mt IV v1-1-1-4, all tarsi with ventral macrosetae. Calamistrum with three staggered rows with <6 setae; calamistrum gap present. Epigastric furrow unmodified. Endogyne of females from the same locality as in the diagnosis ( Fig. 9 ). State of the specimen: good.

Variation

Males (N = 3): total length 4.89–5.67 (5.27), carapace length 2.25–2.78 (2.47), femur I length 3.62–4.04 (3.83), femur/carapace ratio 1.45–1.61 (1.56). Females (N = 5): total length 6.47–8.95 (7.79), carapace length 3.17–3.98 (3.57), femur I length 3.5–4.18 (3.86), femur/carapace ratio 1.01–1.15 (1.08). The shape of the spermathecae varies only slightly among the dissected females.

Natural history

Filistata betarif sp. nov. is a troglophilic species, inhabiting both epigean and hypogean (caves) habitats. It is known from the mesic Mediterranean region of Israel, in scrubland habitats, and usually hides in a refugium made of cribellate silk in crevices, holes or under stones and on rocky walls ( Fig. 1A–B ). All specimens were collected from Bet ‘Arif wadi near Shoham, Hadom Shomeron Nature Reserve, central Israel. The species was found in low abundances outside of caves ( Fig. 1D ) and in very high abundances, year-around, in Tinshemet cave that is characterized by a high guano amount of the Egyptian fruit bat [ Rousettus aegyptiacus (Geoffroy, 1810) ]. Tinshemet (Arabic name: Mugharet al-Watwat) is a small to medium-sized Paleolithic cave, formed in a limestone rock of the Turonian Bina formation. It is located on the east bank of Bet ‘Arif wadi (Arabic name: Wadi Adasiyeh), 95 meters a.s.l. and 15 meters above the wadi ( Frumkin et al. 2016). It has a main opening on the side of the cliff, and an additional chimney opening, which the frugivorous bats frequently use (personal observation). Filistata betarif sp. nov. was also found in lower abundances in the nearby Oah cave that houses no bats ( Cuff et al. 2021; Gavish- Regev et al. 2021). Oah is a small cave, situated on the same east bank of Bet ‘Arif wadi, north to Tinshemet cave, and is formed in a similar rock and age (Boaz Langford, pers. comm.).

In both caves, Filistata betarif sp. nov. was found mainly at the entrance zones, on the cave walls ( Fig. 1A–B ). Other caves surveyed by us, at the mesic Mediterranean, semi-arid, and arid regions of Israel and Palestine, were not found to be occupied by F. betarif sp. nov., but instead by its sibling troglophilic species F. insidiatrix [Berniki, Yir’on, Yonim (Galilee); Ezba’, Oren (Karmel); Sal’it (northern Samaria); Andartat HaBiqa’ (central Jordan Valley); Perat (northern Judean desert); Qumeran (northern Dead-Sea area)] ( Cuff et al. 2021; Gavish-Regev et al. 2021).

Distribution

Known only from two caves and under stones in their surroundings in Bet ‘Arif wadi, Hadom Shomeron Nature Reserve, central Israel ( Fig. 4 ).

Figures

Fig. 1. A–B. Habitat and webs of Filistata betarif sp. nov. in Tinshemet cave. C. Paratype male in life (MACN-Ar 41793). D. Surroundings of Tinshemet cave. Note the entrance to the cave at the right side of the picture. Photos A–B by Shlomi Aharon and C–D by Ivan L.F. Magalhaes.

Fig. 3. A–G. Endogynes Filistata Latreille, 1810, ventral view. A. Filistata wunderlichi Zonstein & Marusik, 2019 from Spain, Boca de los Frailes (CRBA IFM-2197). B. Filistata IFMsp191 from Israel, Odem (MACN-Ar 41262). C. Filistata insidiatrix (Forsskål, 1775) from France, Gonfaron (MACN-Ar 39461). D. Filistata insidiatrix from Israel, Beriniki (HUJ-INVAr 21052). E. Filistata betarif sp. nov. from Israel, Oah cave (HUJ-INVAr 21055). F. Filistata betarif sp. nov. from Israel, Tinshemet cave (MACN-Ar 41222). G. Filistata IFMsp99 (subadult ♀?) from Palestine, En Fashkha (HUJ-INVAr 20277). — H–I. ♂♂, left pedipalp, prolateral. H. Filistata betarif sp. nov., holotype from Israel, Tinshemet cave (HUJ -INVAr 21058). I. Filistata insidiatrix from Spain, Mallorca (MACN-Ar 42346). Scale bar = 0.1 mm, all figures to scale. Colours correspond to clades in Fig. 2.

Fig. 5. A, C, E. Filistata betarif sp. nov., paratype, ♂, from Israel, Tinshemet cave, (MACN-Ar 41793), left pedipalp. B, D, F. Filistata insidiatrix (Forsskål, 1775), ♂, from Spain, Mallorca (MACN-Ar 42347), right pedipalp, mirrored. A–B. Prolateral view. C–D. Dorsal view. E–F. Retrolateral view. Abbreviations: k = embolic keel; ps = prolateral setae. Scale bars = 0.1 mm.

Fig. 7. Filistata betarif sp. nov., habitus. A–C. Holotype, ♂, from Tinshemet cave (HUJ -INVAr 21058). A. Dorsal view. B. Lateral view. C. Ventral view. — D–F. Paratype, ♀, from the same locality (MACN- Ar 41222). D. Lateral view. E. Dorsal view. F. Ventral view. Scale bars = 1 mm.

Fig. 8. A–G. Filistata betarif sp. nov., holotype, ♂ (HUJ -INVAr 21058), left pedipalp. A–B, E. Prolateral view. Arrow points to prolateral strong setae in cymbium. C, F–G. Retrolateral view. Arrow points to embolic keel. D. Dorsal view. — H. Filistata insidiatrix (Forsskål, 1775) (MACN-Ar 39461), bulb, retrolateral view. Arrow points to embolic keel. Scale bars: A–F = 0.2 mm; G–H = 0.05 mm.

Fig. 9. Filistata betarif sp. nov., female spermathecae, ventral. A. Oah cave (HUJ-INVAr 21055). B. Tinshemet cave (MACN-Ar 41222). C–D. Tinshemet cave (MACN-Ar 41791). Scale bars: A–C = 0.2 mm; D = 0.05 mm.

Fig. 10. Filistatid morphology. A. Filistata insidiatrix (Forsskål, 1775), ♀, leg tarsus, arrow points to macrosetae. B. Pritha nana (Simon, 1968), arrow points to right calamistrum. C. Sahastata aravaensis Ganem et al., 2022, left calamistrum. D. Zaitunia schmitzi Kulczyński, 1911, left calamistrum, arrow points to gap between setae. E. Filistata insidiatrix, left calamistrum. F. Filistata lubinae Zonstein & Marusik, 2019, ♂, tarsus I. G. Sahastata aravaensis, ♀. H. Pritha IFMsp185, ♀. I. Pritha sp., male abdomen, dorsal view. J. Zaitunia schmitzi, ♂. K. Zaitunia schmitzi, spermathecae ventral view. L. Pritha tenuispina (Strand, 1914), ♂, left pedipalp, prolateral view, arrow points to cymbium. M. Filistata lubinae, left copulatory bulb, prolateral view. N. Zaitunia schmitzi, ♂, left pedipalp, prolateral view, arrow points to cymbium. O. Filistata albens Zonstein & Marusik, 2019, left copulatory bulb, prolateral view. P. Zaitunia schmitzi, male clypeus, lateral view, arrow points to stout setae. Q. Filistata insidiatrix, female clypeus, lateral view. Scale bars = 0.2 mm.

Fig. 4. Distribution maps. A. Filistata insidiatrix (Forsskål, 1775) in the Mediterranean and Middle East (note the species has been recorded from Angola, Venezuela and the islands of Socotra, Azores and Cabo Verde, but these records seem to be introduced); literature records have been taken from Benoit (1968), Brignoli (1982), Marusik & Zonstein (2014), Marusik & Zamani (2015a) and Zonstein & Marusik (2019). B. Filistata insidiatrix (circles) and F. betarif sp. nov. (star) in Israel and Palestine; only specimens examined by us are included in the map. Darker shades of orange are records that include at least one male specimen. C. Other species of Filistata Latreille, 1810 in Israel (except F. insidiatrix).