Filistata insidiatrix ( Forsskål, 1775 )

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 : 155-161

Filistata insidiatrix ( Forsskål, 1775)

Figs 2 , 3C–D, I , 5 B, D, F , 6 , 8H , 10A, E, Q

Aranea insidiatrix Forsskål, 1775: 86 .

Filistata insidiatrix – Simon 1895: 1067.

Remarks

Please see WSC (2022) for a full synonymic list of this species, and Zonstein & Marusik (2019) for description, diagnosis, natural history, and additional distribution records.

Updated diagnosis

Males are similar to those of F. betarif sp. nov. and F. albens by having an elongate pedipalpal femur and tibia, and by a subconical copulatory bulb with a terminally hooked embolus ( Fig. 6 ; see also Zonstein & Marusik 2019: fig. 27). Males differ from those of F. albens by their longer copulatory bulb and from those of F. betarif sp. nov. by their inconspicuous embolic keel ( Fig. 8H (arrow), 5B, k) (conspicuous keel in F. betarif sp. nov., Fig. 5A ). Females have a pair of large spermathecae that may be partially divided or undivided ( Fig. 3C–D ) and their variation includes the observed morphology of F. betarif sp. nov.; thus females are not diagnosable through morphology alone, they can be diagnosed using DNA barcoding of the COI gene.

Material examined

ANGOLA • 1 ♀; Luanda, Luanda; [8.83833° S, 13.23444° E]; N. Chatelain leg.; USNM 1656 • 1 ♀; Namibe, Baia das Pipas , in house; [14.95° S, 12.18333° E]; Jan. 1960; Brühl leg.; ZMB .

EGYPT • 1 ♀; Cairo; [30.04442° N, 31.23571° E]; O. Manley leg.; ZMB 4375 .

FRANCE • 1 ♂, 1 ♀; Alpes-Côte d’Azur, Var, Brignoles , Gonfaron ; 43.34158° N, 6.29443° E; 2017; M. Stockmann leg.; MACN-Ar 39461 • 2 ♂♂, 4 ♀♀; Languedoc-Roussillon, Banyuls-sur-Mer ; [42.48343° N, 3.12887° E]; 21 Mar. 1968; H.W. Levi, F. Levi and L.R. Levi leg.; MCZ 39864 • 1 ♀; no further data; [42.44097° N, 3.16469° E]; Spatz leg.; ZMB 1053 • 2 ♀♀; southern France ; AMNH • 1 ♀; southern France ; ZMB 5021 .

GREECE • 1 imm.; Attica, Sounion ; [37.65915° N, 24.01476° E]; 30 Nov. 1978; B. Malkin leg.; AMNH • 2 ♀♀; Crete, Akrotiri ; [35.55°N, 24.13333° E]; Feb.–Mar. 1925; A. Schultz leg.; ZMB 6205 • 1 ♀; Crete, Agios Nikolaos ; 35.31006° N, 25.57781° E; Jul. 2019; M. Stockmann leg.; MACN-Ar 41844 • 3 ♀♀, 1 imm.; NW Crete; [35.55° N, 24.13333°E]; A. Schultz leg.; ZMB 6208 • 1 ♀; Crete; [36.39316° N, 25.46151° E]; Oertzen leg.; ZMB 6951 • 1 ♀; Cyclades Islands, Santorini ; 36.40922° N, 25.43221° E; May 1988; M.J. Ramírez leg.; MACN-Ar 20563 • 1 imm.; Kea ; [37.60758° N, 24.31037° E]; 30 Sep. 1925; A. Schultz leg.; ZMB 6203 • 2 ♀♀; Kythira ; [36.26327° N, 22.97737° E]; Aug. 1925; ZMB • 1 ♀; Rhodes, Kalavarda ; [36.33998° N, 27.94949° E]; 7 Dec. 1978; B. Malkin leg.; AMNH • 1 ♀; Thera ; [36.39316° N, 25.46151° E]; 1972; Hillas and Goerligen (?) leg.; ZMB .

ISRAEL • 1 ♂; upper Galilee, Ba’al-Shem-Tov Forest, Meron Mountains ; 32.9821° N, 35.47381° E; 13–20 Nov. 2007; T. Levanony leg.; SMNH • 1 ♂, 1 ♀, 1 imm.; upper Galilee, Dan ; 33.24° N, 35.655° E; 14 May 2012; S. Zonstein leg.; SMNH • 1 ♂; upper Galilee, En Ya’aqov ; 33.00764° N, 35.23889° E; 4–9 Feb. 2007; I. Shtirberg leg.; pitfall; SMNH • 1 ♂; upper Galilee, Nahal Keziv ; [33.04234° N, 35.17882° E]; 31 Jul. 2010; C. Drees and L. Friedman leg.; SMNH • 1 ♂; same locality as for preceding; 13 Nov. 2010; L. Friedman and C. Drees leg.; SMNH • 1 ♂, 2 ♀♀; upper Galilee, near Elon, Nahal Bezet Nature Reserve , Sharakh cave , entrance of cave; 33.074° N, 35.2379° E; 233 m. a.s.l.; 17 Feb. 2020; I.L.F. Magalhaes, E. Gavish-Regev, Z. Ganem, S. Aharon, N. Givon and M. Arnedo leg.; MACN- Ar 41218 • 1 ♂, 2 ♀♀; same collection data as for preceding; MACN-Ar 41226 • 1 ♂; same locality as for preceding; 17 Feb. 2020; I.L.F. Magalhaes, E. Gavish-Regev, Z. Ganem, S. Aharon, N. Givon and M. Arnedo leg.; twilight zone; MACN-Ar 41261 • 1 ♀; same collection data as for preceding; MACN-Ar 41825 • 1 ♂; upper Galilee, Senir ; 33.23333° N, 35.66667° E; 250 m. a.s.l.; 2 Nov. 2013; S. Zonstein leg.; SMNH • 1 ♀; upper Galilee, Yehiam , in cave; [32.99611° N, 35.22109° E]; 15 Nov. 1949; A. Shulov leg.; HUJ-INVAr 20290 • 1 ♀; upper Galilee, Yir’on , small cave; 33.0672° N, 35.4672° E; 28 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21049 • 1 ♀; upper Galilee, Yonim cave , in depth; 32.9236° N, 35.2168° E; 25 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ -INVAr 21050 • 1 ♂; upper Galilee, north slope of Mount Meron, Meron Field School ; 33.01667° N, 35.39167° E; 1 Nov. 2013; S. Zonstein leg.; SMNH • 2 imm.; lower Galilee, near Teverya ( Tiberias ), Berniki , large cave; 32.7775° N, 35.5401° E; 27 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21051 • 1 ♀; lower Galilee, near Teverya ( Tiberias ), Berniki , medium cave; 32.7768° N, 35.5413° E; 27 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21052 • 1 ♂; Mount Karmel, HaKarmel Nature Reserve, Karmel Ridge , Nahal Me’arot ; Gamal cave ; [32.6705° N, 34.96598° E]; 4 Nov. 2013; D. Rotem leg.; SMNH • 1 ♀; Mount Karmel, near Hefa (Haifa), HaKarmel Nature Reserve , Karmel Ridge ; Ornit cave , inside cave; 32.7567° N, 34.98975° E; 209 m. a.s.l.; 20 Feb. 2020; I.L.F. Magalhaes, E. Gavish-Regev, Z. Ganem, S. Aharon and M. Arnedo leg.; MACN-Ar 41215 • 1 ♂, 1 ♀; same collection data as for preceding; MACN-Ar 41813 • 1 ♂; Mount Karmel, Yagur ; [32.74153° N, 35.07683° E]; 4 Dec. 1939; HUJ-INVAr 20283 • 1 ♀; coast of Karmel, ‘ Atlit ; [32.69088° N, 34.94291° E]; 8 Sep. 1944; HUJ-INVAr 20287 • 1 ♂; coast of Karmel, Ma'agan Mikha'el , malaise trap; [32.55871° N, 34.91782° E]; 2–28 Dec. 2010; W. Kuslitzky leg.; SMNH • 1 ♂; same locality as for preceding; 12 Oct. 1992; Y. Lubin leg.; HUJ • 1 ♀; coast of Karmel , route to Zikhron-Y’aqov [32.57391° N, 34.95198° E]; 24 Mar. 1940; HUJ-INVAr 20286 • 1 ♀; Judean Hills, ‘ Adullam National Park, Midras ; 31.655° N, 34.94167° E; 15 Nov. 2012; S. Zonstein leg.; SMNH IFM-0272 • 1 ♂, 1 ♀; Judean Hills, ‘ Adullam National Park ; [31.64474° N, 34.96089° E]; Apr. 2003; U. Columbus leg.; SMNH • 2 ♂♂; same locality as for preceding; 20 May 2002; Y. Mandelik and A. Landsman leg.; SMNH • 1 ♂; Judean Hills, Ramat Avishur ; [31.65° N, 34.92° E]; 14 May 2002; Y. Mandelik leg.; pitfall trap; HUJ • 1 ♀; Judean Mountains , Jerusalem; [31.78333° N, 35.2° E]; 29 May 1944; A. Shulov leg.; HUJ-INVAr 20288 • 1 ♀, 1 imm.; same locality as for preceding; Oct. 1936; A. Shulov leg.; HUJ-INVAr 20284 • 1 ♀; Judean Mountains, Newe Ya’akov (= Kefar Ivri ); [31.84177° N, 35.24276°]; 30 Dec. 1939; A. Shulov leg.; HUJ-INVAr 20289 • 1 ♀; Judean Mountains, Qiryat'Anavim , under bark; [31.80991° N, 35.12084° E]; 20 Jan. 1940; HUJ-INVAr 20285 • 1 ♂; Samaria, Gilboa' ; [32.48248° N, 35.42137° E]; 30 Jan. 2011; L. Friedman and C. Drees leg.; SMNH • 1 ♀; Samaria, Sal’it cave ; 32.2454° N, 35.0456° E; 25 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21053 • 1 ♂; Sharon plain, Ramot Hashavim , in house; 32.16667° N, 34.88333° E; 9 Mar. 2013; T. Eshcoly leg.; SMNH • 1 ♂; Jordan Valley, Nahal HaGal ; [32.6341° N, 35.534° E]; 16 Jul. 2010; C. Drees and L. Friedman leg.; SMNH • 2 imm.; Jordan Valley, Nir Dawid (= Tel’ Amal ); [32.50406° N, 35.45762° E]; 29 Sep. 1939; A. Shulov leg.; HUJ-INVAr 21061 .

ITALY • 1 ♂, 1 ♀; Sardinia, Baunei ; [40.03175° N, 9.66327° E]; J. Wunderlich leg.; SMF 59579 • 1 ♂; Sardinia, Porto Torres ; [40.83134° N, 8.40587° E]; 7 Apr. 1952; Esk. Zool. Just. Ffn. Leg.; SMF • 1 ♀; Sorgono ; [40.0272° N, 9.10196° E]; 1917; Häuser and Krausse leg.; ZMB • 7 ♀♀, 6 imm.; Toscania, Siena, 4 km S of San Giminiano, Fattoria Voltrona ; [43.46728° N, 11.04348° E]; 12 Dec. 2001; M.J. Ramírez leg.; MACN-Ar 10378 • 1 ♀; same collection data as for preceding; MACN-Ar 10379 .

JORDAN • 1 ♂; Amman, Fuhais ; [32.00408° N, 35.78451° E]; Dec. 1980; R. Kinzelbach leg.; SMF 57068 .

MOROCCO • 1 ♀; Saffi ; [32.30082° N, 9.2272° W]; Quedenfeldt leg.; ZMB • 1 ♂, 3 ♀♀; Sefrou , Travertine Falls; 33.83333° N, 4.83333° W; 25 Dec. 1986; V. and B. Roth leg.; CAS 9060650 • 1 imm.; same locality as for preceding; 25 Dec. 1996; V. Roth leg.; CAS 9057617 • 1 ♀, 1 imm.; Souss-Massa- Draâ , Ouarzazate , Amerzgane, Telouet; 31.06659° N, 7.14178° W; Oct. 2016; M. Stockmann leg.; MACN-Ar 41845 • 1 ♂, 1 ♀; Tafilalt , Errachidia , Aoufous, Ziz Valley; 31.75176° N, 4.1968° W; Oct. 2016; M. Stockmann leg.; MACN-Ar 39463 .

PALESTINE • 1 ♀; Judean desert, 15 km E of Jerusalem, Fawar Spring, Wadi Qelt; [31.8401° N, 35.3489° E]; 7 Mar. 1975; H.W. Levi leg.; MCZ 39890 • 1 ♀, 1 imm.; Judean desert, Nahal Perat Nature Reserve, Inbal cave; 31.8332° N, 35.3019° E; 26 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 21054 • 1 ♂, 1 ♀; Judean desert, Nahal Perat Nature Reserve, I’qul cave; [31.8318° N, 35.30842° E]; 26 May 2014; S. Aharon and E. Gavish-Regev leg.; HUJ-INVAr 20276 .

SPAIN • 4 ♀♀; Alicante, Benidorm ; [38.54106° N, 0.12249° W]; 1961; K.W. Haller leg.; AMNH • 1 ♂, 1 ♀, 3 imm.; same collection data as for preceding; AMNH • 1 ♀; Almeria, 25 km from Almeria, Salinas de Cerillos ; [36.70498° N, 2.66982° W]; 25 Jun. 1967; B. Malkin leg.; AMNH • 1 imm.; Balearic Islands, Mallorca, Colonia de Sant Jordi , 50 km SE of Palma, on house walls; 39.31727° N, 2.99381° E; 15 Sep. 2007; M.J. Ramírez leg.; MACN-Ar 20558 • 1 ♀; same collection data as for preceding; MACN-Ar 20560 • 1 ♀; same collection data as for preceding; MACN-Ar 20561 • 1 imm.; same collection data as for preceding; MACN-Ar 20564 • 1 ♀; Balearic Islands, Mallorca, Colonia Saint Jordi , 50 km SE of Palma, on walls of houses; [39.31727° N, 2.99381° E]; 15 Sep. 2007; M.J. Ramírez leg.; MACN- Ar 42348 • 1 ♂; same collection data as for preceding; MACN-Ar 42347 • 1 ♂; same collection data as for preceding; MACN-Ar 42346 • 1 ♀; Barcelona ; [41.38504° N, 2.17347° E]; 13 Sep. 1986; J.A. Coddington leg.; caught in the middle of an imaginal molt, has two sets of spermathecae; USNM 1656 • 1 ♀; same collection data as for preceding; USNM 1656 • 2 ♂♂; Castellón , N of L’Alcora, stone walls; 40.08974° N, 0.19945° W; 300 m. a.s.l.; 30 May 2010; S. Huber and A. Schönhofer leg.; at night; SMF • 1 ♀; Cataluña, Barcelona, Barcelona ; 41.40423° N, 21.54461° E; Jan. 1988; M.J. Ramírez leg.; MACN- Ar 20562 • 1 ♀, 1 imm.; Madrid; [40.41678° N, 3.70379° W]; 1961; K.W. Haller leg.; AMNH • 3 ♀♀, 1 imm.; Madrid, Navas del Rey ; [40.38776° N, 4.25° W]; 1961; K.W. Haller leg.; AMNH • 9 ♀♀, 11 imm.; Madrid, Vaciamadrid ; [40.35191° N, 3.53573° W]; 1961; K.W. Haller leg.; AMNH • 22 ♀♀, 6 imm.; Madrid, Vaciamadrid ; 1961; K.W. Haller leg.; AMNH • 1 ♀; Murcia, Fortuna, Espacio Natural de Ajanque ; [38.14949° N, 1.10128° W]; CRBA 861 • 1 ♂; same locality as for preceding; 19 Jun. 2004; CRBA 860 • 4 ♀♀, 1 imm.; Tarragona, Monte Poulet ; 41.11903° N, 1.24451° E; 23 Dec. 1986; M.J. Ramírez leg.; MACN-Ar 20559 • 1 ♂; Teruel, Molinos ; 40.81945° N, 0.45083° W; Apr. 1985; J. Moles leg.; MACN-Ar 20565 • 1 ♀, 1 imm.; no further data, on a slope; [39.66422° N, 0.22674° W]; 17 Jun. 1993; S. Lideese leg.; ZMB 30578 .

TUNISIA • 1 ♀; Djerba; [33.8076° N, 10.84515° E]; V. Bilgner leg.; ZMB K.786 .

VENEZUELA • 1 ♀; Miranda, Caracas suburbs, caves near Cementerio del Este ; [10.45038° N, 66.81118° W]; Jul. 1986; J.A. Coddington leg.; USNM 3222 .

NO DATA • 1 ♂, 2 ♀♀, 2 imm.; Mediterranean region ; ZMB • 1 ♀; ZMB 533 • 1 ♀; ZMB 580 • 1 ♀; MNRJ 1434 .

Distribution

Widely distributed in the Mediterranean region and the Middle East ( Fig. 4A ), with records (probably introduced) in the Azores, Socotra Island, Cabo Verde, Angola and Venezuela. In Israel, it is distributed in Mediterranean areas in the north ( Fig. 4B ); in some localities only females are known, and thus it is possible these records could belong to F. betarif sp. nov., although it is more likely these records are of F. insidiatrix since they broadly overlap with records of males.

Figures

Fig. 2. Phylogenetic tree of cytochrome c oxidase subunit I haplotypes of different species of Filistata Latreille, 1810 inferred under maximum likelihood. Numbers below branches are bootstrap supports. The histogram shows the distribution of pairwise Kimura two-parameter genetic distances within and between F. insidiatrix (Forsskål, 1775) and F. betarif sp. nov. (for pairwise distances of other species, see Table 2). Note that some intraspecific genetic distances within F. insidiatrix are larger than interspecific distances between F. insidiatrix and F. betarif sp. nov., denoting a lack of a barcode gap in this group, or cryptic species within F. insidiatrix.

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. 6. Filistata insidiatrix (Forsskål, 1775), left bulb, prolateral (A, C, E, G) and dorsal (B, D, F, H) views. A–B. France (MACN-Ar 39461, ♂ collected with ♀ IFM-1702). C–D. Spain (CRBA IFM- 2196). E–F. Israel, Ornit cave (MACN-Ar 41813, IFM-2175). G–H. Israel, Sharakh cave (MACN-Ar 41218, IFM-2144). Scale bar = 0.1 mm, all figures to scale.

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. 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).