Systematics and phylogeography of western Mediterranean tarantulas (Araneae: Theraphosidae) Author Korba, Jan Author Opatova, Vera Author Calatayud-Mascarell, Arnau Author Enguídanos, Alba Author Bellvert, Adrià Author Adrián, Silvia Author Sánchez-Vialas, Alberto Author Arnedo, Miquel A text Zoological Journal of the Linnean Society 2022 2022-10-01 196 2 845 884 https://academic.oup.com/zoolinnean/article/196/2/845/6619232 journal article 164207 10.1093/zoolinnean/zlac042 44bacddc-4f3a-42e3-8a6e-9f5d9180968d 0024-4082 7184518 ISCHNOCOLUS ELONGATUS ( SIMON, 1873 ) ( FIGS 6 A-L, 7, 8, 9C, D, 10C–F, 13C–F, 21A–N, 22A– G, 23A–F, 24A–H) Cyrtauchenius elongatus Simon, 1873: 32 (description of female). Moggridge (1874: 182 , 189, 248, pl. XIII, fig. B; burrow entrance); Savory (1928: 290) . Presumably deposited in MNHN, not found by Zonstein (2018) , not examined. Topotypes (Ksar el Kebir) were included in this study. Leptopelma africana Ausserer, 1875: 167 (description of female). Synonymized with Cyrtauchenius elongatus Simon, 1873 by Simon (1889: 396) . Leptopelma elongata Simon 1889: 395 , pl. XIII, fig. 2 (female, burrow entrance), Simon (1909: 9) ; Reimoser (1919: 7) ; Berland (1932: 110 , fig. 219; burrow entrance). Luphocemus insidiosus Denis, 1960: 186–189 (description of female), illustration of burrow (fig. 2, p.187). Deposition place unknown. Type not examined. Topotypes (Benslimane) were included in this study. New synonymy . Harpactirella insidiosa Benoit, 1965: 297 ; CalatayudMascarell & Sánchez-Vialas (2020: figs 2–4, adult female and burrow). Ischnocolus hancocki Smith, 1990: 127 , figs 803–818 (female); Guadanucci & Wendt (2014: 394 , fig. 4A; female); Zonstein (2018: 107 , figs1–8; male). Deposited in BMNH, type not examined. Topotypes (Larache) were morphologically examined. New synonymy . Ischnocolus elongatus: Zonstein (2018: 106) . Type material: Type locality Ksar el Kebir ( Morocco ), female holotype presumably in MNHN, not found by Zonstein (2018) , not examined. Topotypes were included in this study. Material examined: Morocco : 9♀♀ ( CRBAMM000430 , CRBAMM000432,CRBAMM000424,CRBAMM000425, CRBAMM000426,CRBAMM000427,CRBAMM000428, CRBAMM000429, CRBAMM000431), province of Fés, Imouzer, 33°38 ′ 39 ″ N , 5°04 ′ 09 ″ W , 12.iv.2010 ( V . Opatova, M, Arnedo leg.) . 1♀ , 2 juv. ( CRBAMM000351 , CRBAMM000350), province of Taroudant , Cova Maravilles , 30°10 ′ 40 ″ N , 8°17 ′ 38 ″ W , 8.iv.2010 ( V . Opatova & M.Arnedo leg.) . 3♀♀ , 3 juv. ( CRBAMM000372 , CRBAMM000363,CRBAMM000371,CRBAMM000364, CRBAMM000365, CRBAMM000367), province Al Haouz , Asni , 31°11 ′ 22 ″ N , 8°03 ′ 27 ″ W , 9.iv. 2010 ( V . Opatova , M. Arnedo leg.) . 1♀ ( CRBAMM000534 ), province of Taounate , Rhafsai , 34°37 ′ 55 ″ N , 4°55 ′ 52 ″ W , 16.iv.2010 ( V . Opatova , M. Arnedo leg.) . 1♂ , 1 juv. ( CRBAMM000397 , CRBAMM000403), province of Azilal , Tilouguite , 32°05 ′ 04 ″ N , 6°20 ′ 00 ″ W , 10.IV.2010 ( V . Opatova & M. Arnedo leg.) . 3♀♀ , 4 juv. ( CRBAMM 000411 , CRBAMM 000406, CRBAMM000407,CRBAMM000408,CRBAMM000410, CRBAMM000412, CRBAMM000413), province of Béni- Mellal , Ab el Hamam, 32°31 ′ 19 ″ N , 4°55 ′ 52 ″ W , 11.iv.2010 ( V. Opatova & M. Arnedo leg.) . 3♀♀ ( CRBAMM000468 , CRBAMM000467, CRBAMM000470), province of Taza , Sidi Abdulah , 32°31 ′ 20 ″ N , 6°02 ′ 02 ″ W , 13.iv.2010 ( V . Opatova & M.Arnedo leg.) . 1♀ , 1 juv. ( CRBAMM000483 , CRBAMM000484), province of Oujda , Ain Sfa , 34°49 ′ 28 ″ N , 2°05 ′ 12 ″ W , 14.IV.2010 ( V . Opatova & M. Arnedo leg.) . 4♀♀ , 4 juv. ( CRBAMM000501 , CRBAMM000502,CRBAMM000503,CRBAMM000504, CRBAMM000506,CRBAMM000507,CRBAMM000508, CRBAMM000509), province of Al Hoceima , Bni Hadifa , 35°00 ′ 19 ″ N , 4°10 ′ 44 ″ W , 15.iv.2010 ( V . Opatova & M. Arnedo leg.) . 1♀ , 2 juv. ( CRBA004970 , CRBA004971, CRBA004972), province of Taroudant , Imgoune , 30°16 ′ 29 ″ N , 8°16 ′ 26 ″ W , 22.ii.2020 ( J. Korba leg.) . 2♀♀ , 3 juv. ( CRBA004973 , CRBA004974, CRBA004975, CRBA004976, CRBA004977), province of Tiznit , Tafraout , 29°40 ′ 54 ″ N , 9°01 ′ 54 ″ W , 23.ii.2020 ( J. Korba leg.) . 2 juv. ( CRBA004985 , CRBA004986), province of Guelmim , Mesti , 29°11 ′ 52 ″ N , 10°05 ′ 15 ″ W , 25.ii.2020 ( J. Korba leg.) . 5♀♀ , 2 juv. ( CRBA004992 , CRBA004993, CRBA 004994, CRBA 004995, CRBA 004996, CRBA004997, CRBA004998), province of Essaouira , Ounagha , 31°31 ′ 32 ″ N , 9°37 ′ 38 ″ W , 28.ii.2020 ( J. Korba leg.) . 2♀♀ ( CRBAMM000446 , CRBAMM000447), province of Fés , Pantano , 34°03 ′ 33 ″ N , 5°18 ′ 12 ″ W , 12.iv.2010 ( V . Opatova , M. Arnedo leg.) . 2 juv. ( CRBAMM000352 , CRBAMM000353), province of Taroudant , Azoura , 30°01 ′ 31 ″ N , 8°35 ′ 24 ″ W , 8.iv.2010 ( V . Opatova , M. Arnedo leg.) . 4 juv. ( CRBAMM000373 , C R B A M M 0 0 0 3 7 4, C R B A M M 0 0 0 3 7 5 CRBAMM000376), province of Azilal , near Ouzoud falls , 31°57 ′ 35 ″ N , 6°46 ′ 05 ″ W , 10.IV.2010 ( V . Opatova & M. Arnedo leg.) . 1♀ ( CRBAMM000419 ), province of Khénifra , Oumer Riba , 32°55 ′ 41 ″ N , 5°30 ′ 34 ″ W , 11.iv. 2010 ( V . Opatova , M. Arnedo leg.) . 2♀♀ ( CRBAMM000448 , CRBAMM000449), province of Fés , Djebel Zalach , 34°06 ′ 23 ″ N , 4°58 ′ 10 ″ W , 12.IV.2010 ( V . Opatova & M.Arnedo leg.) . 3♀♀ , 1 juv. ( CRBAMM000488 , C R B A M M 0 0 0 4 8 9, C R B A M M 0 0 0 4 9 0 CRBAMM000491), province of Berkane , Beni Snassen , 34°48 ′ 12 ″ N , 2°23 ′ 48 ″ W , 14.IV.2010 ( V . Opatova & M. Arnedo leg.) . 1♀ ( CRBA004980 ), Bou Tazlaft , 29°37 ′ 40 ″ N , 9°52 ′ 52 ″ W , 24.ii.2020 ( J. Korba leg.) . 1♀ ( CRBA004991 ), province of Essaouira , Ida Ou Guelloul , 30°54 ′ 00 ″ N , 9°43 ′ 04 ″ W , 27.ii.2020 ( J. Korba leg.) . 2♀♀ ( CRBA004978 , CRBA004979), province of Tiznit , Tighmi , 29°34 ′ 32 ″ N , 9°23 ′ 51 ″ W , 24.ii.2020 ( J. Korba leg.) . 1 juv. ( CRBAMM000357 ), province of Taroudant , Tafingoul , 30°44 ′ 32 ″ N , 8°24 ′ 06 ″ W , 9.iv.2010 ( V . Opatova & M. Arnedo leg.) . 1 juv. ( CRBAMM000356 ), province of Tiznit , Tizegzauine , 29°50 ′ 42 ″ N , 8°56 ′ 14 ″ W , 8.iv.2010 ( V . Opatova & M. Arnedo leg.) . 1♀ ( CRBAMM000415 ), province of Beni Mellal , Cheikh , 32°37 ′ 23 ″ N , 5°58 ′ 27 ″ W , 11.iv.2010 ( V . Opatova & M. Arnedo leg.) . 3♀♀ ( CRBA005022 , CRBA005023, CRBA005024), province of Larache , Ksar el Kebir , 35°02 ′ 03 ″ N , 6°01 ′ 50 ″ W , 26.ii.2010 ( A. Calatayud-Mascarell & A. Sánchez-Vialas leg.) . 3♀♀ ( CRBA005027 , CRBA005028, CRBA005029), province of Benslimane , Benslimane , 33°39 ′ 06 ″ N , 7°05 ′ 23 ″ W , 26.ii.2020 ( A. Calatayud-Mascarell & A. SánchezVialas leg.) . 5♀♀ ( CRBA005032 , CRBA005033, CRBA005034, CRBA005035, CRBA005036), province of Sidi Bennour , Oualidia , 32°36 ′ 38 ″ N , 9°00 ′ 34 ″ W , 25.ii.2020 ( A. Calatayud-Mascarell & A. SánchezVialas leg.) . 1♀ ( CRBAMM000222 ), province of Al Haouz , Tahanaoute , 31°21 ′ 15 ″ N , 7°57 ′ 06 ″ W , 9.iii.2007 ( M. Arnedo & C. Ribera leg.) . 4♀♀ (deposited in Department of Zoology , Charles University ), province of Larache , 35°12 ′ 08 ″ N , 6°06 ′ 03’’W , 12.ix.2021 ( J. Korba & V . Opatova leg.) Remarks: Originally described as Cyrtauchenius elongatus Simon, 1873 , this species has been recently transferred to the genus Ischnocolus by Zonstein (2018) based on Ausserer’s description of Leptopelma africana , which clearly pointed to Theraphosidae (‘… two toothless claws bearing two tufts of hairs in each tarsi’). The three samples from the type locality analysed in our study ( JK 82, JK83, JK84, see Fig. 3 ) were recovered within the elongatus clade, which is clearly defined by its distinct morphology. Similarly, we included individuals from the type locality of H. insidiosa ( JK 87, JK88, see Fig. 3 ), which were shown to belong to the same clade as the remaining species identified as I. elongatus . In the case of I. hancocki , we could not examine the holotype female nor add samples from Larache (type locality) to the molecular analyses. However, we examined the morphology of specimens from the type locality, which fit the redescription by Guadanucci & Wendt (2014) and turned out to be indistinguishable from the rest of the elongatus morphotype samples. Moreover, the type locality Larache is located within the estimated ( SDM ) range of occurrence, only 30 km north-west from the topotype locality of I. elongatus . The putative synonymy of I. elongatus and I. hancocki was already suggested by Zonstein (2018) . Based on these arguments and the detailed examination of topotypes of each taxon, we herein propose I. elongatus as senior synonym of both I. hancocki and Harpactirella insidiosa . Diagnosis: Ischnocolus elongatus differs from all its congeners by the following combination of characters: robust appearance ( Fig. 23 ), apical segment of PLS triangular ( Figs 9C, D , 10C–F , 22D), cephalic region and eye tubercle elevated ( Figs 11B , 22E), presence of black bristles on cheliceral margin (Fig. 22D) and tarsus IV without pseudosegmentation. Females further differ from other Ischnocolus species , except I. vanandalae , by possessing 0–2 apical lobes on spermathecae ( Figs 6A–L , 22F). Males differ from other Ischnocolus species by having reduced spination on ventral tibia I ( Figs 8 , 21B–D) and having a shorter palpal tibia in comparison to the tarsus and patella (Fig. 21F–H). The lifestyle of I. elongatus is unique among its congeners [see Montemor et al. (2020) for natural history of Middle Eastern species], as it excavates deep tube-like burrows (see Natural history section). Description: Male, ( CRBAMM 000397, Tilougguite): Total length 14.12. Colour pattern: Colour in ethanol: legs and carapace light yellow-brown. Carapace with silver hairs (Fig. 21A). Abdomen darker brown with dorsal light striped pattern. Carapace: 5.43 long, 4.72 wide (Fig. 21A); cephalic region raised from lateral view; eye tubercle elevated, 0.68 long, 1.08 wide; fovea slightly procurved (Fig. 21A). Clypeus 0.23 wide. Eyes (Fig. 21E): AME 0.18, PME 0.17, ALE 0.23, PLE 0.20; PME-PME 0.67, ALE-AME 0.26, ALE-PLE 0.34, AMEPLE 0.38, AME-AME 0.35, ALE-ALE 0.81. Sternum, labium and maxillae: sternum 2.60 long, 2.47 wide, setose; labium 0.58 long, 1.02 wide, with approx. 20 cuspules; maxillae with approx. 40 cuspules (Fig. 21I). Abdomen : 6.30 long, 3.23 wide; PLS basal segment 0.84 long, median segment 0.51 long, apical segment 0.52 long, triangular. Chelicerae: 2.27 long; basal article with nine teeth; intercheliceral tumescence present (Fig. 21N). Rastellum sensu Raven (1994) absent, but a group of strong black bristles in front of the fang base is present (Fig. 21N). Pedipalps: spination: femur (p)1ap., ventral furrow on tibia not sigmoid, broad. Length: 7.76 (femur 3.12, patella 1.82, ≤ tibia 2.09, tarsus 1.59). Copulatory bulb: bulb globular, embolus curved with pointed tip ( Figs 7 , 21F– H, J–M). Legs: scopula on all tarsi divided by a thick Figure 21. Ischnocolus elongatus , male, A –N ( CRBAMM 000397). A, prosoma, dorsal view. B, tibia I, prolateral view. C, tibia I, ventral view. D, tibia I, retrolateral view. E, eye tubercle, dorsal view. F, palpal bulbus, prolateral view. G, palpal bulbus, ventral view. H, palpal bulbus, retrolateral view. I, sternum, maxillae, labium and chelicerae, ventral view. J, bulbus, prolateral view. K, bulbus, ventral view. L, bulbus, retrolateral view. M, bulbus, dorsal view. N, chelicerae, prolateral view. Scale bar = 1 mm . band of setae. Scopula on ventral metatarsus I nearly totally occupied, II half occupied, III and IV <half occupied. Paired claws on tarsi I–IV, bipectinate, with two rows of five teeth. Leg measurement: length of legs IV> I> II> III. Leg I: 15.98 (femur 4.76, patella 2.62, tibia 3.39, metatarsus 3.01, tarsus 1.94), leg III: 14.30 (femur 3.54, patella 2.06, tibia 2.27, metatarsus 3.22, tarsus 2.04), leg IV: 19.15 (femur 4.98, patella 2.55, tibia 3.78, metatarsus 4.26, tarsus 2.49). Spines: I femur (p)ap1, (r)ap1, patella (r)1, tibia (r)1-1, (v)2- 2-2, metatarsus (v)1-1; III femur (p)ap1, patella (p)1, tibia (r)1-1, (p)2-2, (v)1-1-2, metatarsus (r)0-1-1, (p)2- 2-2, (v)1-1-1-1; IV femur (r)ap1, (d)ap1, patella (p)1, Figure 22. Ischnocolus elongatus, female, A –G ( CRBA 005027). Prosoma, dorsal view. B, sternum, maxillae and chelicerae, ventral view. C, eye tubercle, dorsal view. D, chelicerae, prolateral view (arrow indicates dense strong black bristles). E, whole body, lateral view. F, spermathecae. G, posterior lateral spinneret, retrolateral view. Scale bar = 1 mm . tibia (r)1-1-1-1-1-1, (p)2-1-1-1, (v)1-1-2, metatarsus (r)1-1, (p)1-2-1-1-2, (v)1-1. Tarsus IV without pseudosegmentation. Female ( CRBA 005027, Benslimane): Total length 25.07. Colour pattern: Colour in ethanol: carapace, chelicerae and legs dark orange-brown, abdomen grey-brown with light striped pattern. Colour of live specimens: northern populations have beige-golden setae on carapace, legs and chelicerae. Basal part of the chelicerae is black without setae, black stripe on the patella of each leg. Abdomen beige-golden with black spots ( Fig. 23C ). Southern populations have orange legs with black setae, carapace golden brown, chelicerae black with golden brown setae. Abdomen light to dark-brown with golden spot pattern ( Fig. 23A, B ). Carapace: 7.74 long, 6.39 wide (Fig. 22A); cephalic region raised from lateral view; eye tubercle strongly elevated (Fig. 22E), 0.99 long, 1.40 wide; fovea deep, straight to slightly procurved (Fig. 22A); clypeus 0.29. Eyes (Fig. 22C): AME 0.20, PME 0.21, ALE 0.26, PLE 0.23; PME-PME 0.89, ALE-AME 0.31, ALE-PLE 0.43, AME-PLE 0.54, AME-AME 0.47, ALE-ALE 1.01. Sternum, labium and maxillae: sternum 3.87 long, 3.44 wide; labium 1.14 long, 1.63 wide, with approx. 20 cuspules; maxillae with approx. 70 cuspules (Fig. 22B). Abdomen: oval, 12.87 long, 6.77 wide (Fig. 22E); PLS basal segment 1.30 long, median segment 0.63 long, apical segment 0.84 long, triangular (Fig. 22G). Vulva: formed by two widely separated triangular receptacles without lobes (Fig. 22F). Chelicerae : robust, 4.20 mm long; basal article with nine teeth; rastellum sensu Raven (1994) absent, but a group of strong black bristles is present on the margin (Fig. 22D). Pedipalps: length: 11.61 (femur 4.26, patella 2.66, tibia 2.52, tarsus 2.17). Spination: femur (p)ap1, tibia, (v)1-1-2, (p)1-2-1. Legs: scopula on all tarsi divided by a thick band of setae.Scopula on ventral metatarsus Ientirely occupied, II four-fifths occupied, III three-quarters occupied, IV three-quarters occupied. Leg measurement: length of legs IV> I> II> III. leg IV: 21.1 (femur 6.05, patella 3.36, tibia 4.58, metatarsus 4.45, tarsus 2.66), leg III: 14.52 (femur 4.59, patella 2.61, tibia 2.60, metatarsus 2.72, tarsus 2.00), leg II: 17.19 (femur 5.21, patella 3.07, tibia 3.59, metatarsus 3.08, tarsus 2.24), leg I: 19.44 (femur 5.99, patella 3.65, tibia 4.13, metatarsus 3.34, tarsus 2.33). Spines: I femur (p)ap1, tibia (v)1-1, tarsus (v)1-0; II femur (p)ap1, tibia (r)0-0-1, (v)1-1-2, tarsus (v)1-1; III femur (r)ap1, (p)ap1, patella (r)1, (p)2, tibia (r)1-1, (p)2-2, (v)1-1-2, metatarsus (r)1, (p)2-2-2, (v)1-1-2; IV femur (r)ap1, patella (r)ap1, tibia (r)1-1- 1, (v)2-1-2-1-2 metatarsus (r)1-2-1, (p)1-1, (v)2-2-2-3. Tarsus IV without pseudosegmentation. Figure 23. Habitus of I. elongatus from different regions of Morocco. A, adult female from Tafraout (south); B, subadult female from Ksar el Kebir (north); C, adult female from Imgoune (south); D, adult female from Benslimane (north); E, adult female from Ounagha (central); F, adult male from Tafraout (south). Scale bar = 10 mm. Photo credits: J. Korba (A, C, E), A. Sánchez-Vialas (B, D), S. Vogler (F). Distribution: The species is currently known only from Morocco , where it ranges from Larache in the north to Mesti in the south. Distribution in Algeria is highly probable. Natural history: Ischnocolus elongatus constructs a 20–30 cm deep tube burrow with an open entrance and a palisade build from surrounding material resembling that of wolf spider genus Lycosa , but with more dense and compact silk lining ( Fig. 24E– H ). Burrows in southern populations (~20 observed) consist of a single, slightly inclined underground tube, sometimes connected to the surface by side exits. Burrows in northern populations seemed to be more complex (~ten burrows observed). A tube vertically extending from the entrance ends after approximately 5–10 cm , forming a chamber where the spider deposits prey remnants and old exuviae. The main burrow connects laterally to the vertical tube few cm below the entrance. The opening to the lateral tube is small and often closed by a dense web. After a short distance, the horizontally oriented side tube turns vertical and continues approx. for another 20 cm . The species occurs in a wide variety of habitats and climates, also in sympatry with either I. valentinus or I. mogadorensis . It has been found in humid localities in northern Morocco , in Aleppo pine ( Pinus halepensis ) or cork-oak ( Quercus suber L.) forest with Mediterranean fan palm ( Chamaerops humilis ) ( Fig. 24C ), in the central coast on light sandy soil with Berber thuja ( Tetraclinis articulata ) ( Fig. 24B ) and in semi-arid localities in southern Morocco among argan stands [ Argania spinosa (L.) Skeels] ( Fig. 24D ) or in Macaronesian vegetation with Euphorbia balsamifera Aiton and E. officinarum L. on hard sandstone bedrock (24A). In Atlas and Anti-Atlas Mountains, the localities do not exceed 2000 m a. s. l.