Ischnocolus mogadorensis, SIMON, 1909

Korba, Jan, Opatova, Vera, Calatayud-Mascarell, Arnau, Enguídanos, Alba, Bellvert, Adrià, Adrián, Silvia, Sánchez-Vialas, Alberto & Arnedo, Miquel A, 2022, Systematics and phylogeography of western Mediterranean tarantulas (Araneae: Theraphosidae), Zoological Journal of the Linnean Society 196 (2), pp. 845-884 : 860-866

ISCHNOCOLUS MOGADORENSIS SIMON, 1909 ,

SP. REVAL.

( FIGS 5C , 7 , 8 , 9B , 10B , 13B , 14A–N , 15A–G , 16A–D , 17A–F )

Ischnocolus mogadorensis Simon (1909 , description of female, p. 10).

Ischnocolus valentinus: Guadanucci & Wendt (2014: 391 , as senior synonymy of I. mogadorensis ).

Type material: Female holotype. Morocco: Mogador ( Essaouira), Martínez de la Escalera leg., presumably deposited in MNHN in Paris, lost ( Guadanucci & Wendt, 2014). Not examined.

Material examined: Morocco: 1♂, 2♀♀ (CRBA004988, CRBA004989, CRBA004990), province of Agadir, Ankrime, 30°37 ′ 25 ″ N, 9°34 ′ 50 ″ W, 26.ii.2020 (J. Korba leg.). 1♂, 3 juv. (CRBA004981, CRBA004982, CRBA004983, CRBA004984), province of Tiznit, Bou Tazlaft, 29°37 ′ 40 ″ N, 9°52 ′ 52 ″ W, 24.ii.2020 (J. Korba leg.), 1 juv. (CRBAMM000346), province of Agadir, Imoza, 30°38 ′ 34 ″ N, 9°42 ′ 20 ″ W, 7.iv.2010 ( V. Opatova & M. Arnedo leg.). 1♀ (CRBAMM000342), province of Essaouira, Tamanat, 31°00 ′ 21 ″ N, 9°35 ′ 49 ″, 7.iv.2010 ( V. Opatova & M. Arnedo leg.). 1 juv. (CRBAMM000348), province of Taroudant, Cova Maravilles, 30°10 ′ 40 ″ N, 8°17 ′ 38 ″ W, 8.iv.2010 ( V. Opatova & M. Arnedo leg.). 1♀, 1 juv. (CRBAMM000370, CRBAMM000366), province Al Haouz, Asni, 31°11 ′ 22 ″ N, 8°03 ′ 27 ″ W, 9.iv.2010 ( V. Opatova, M. Arnedo leg.). 2 juv. (CRBAMM000358, CRBAMM000359), province of Taroudant, Tafingoult, 30°44 ′ 31 ″ N, 8°24 ′ 06 ″ W, 9.iv.2010 ( V. Opatova, M.Arnedo leg.). 1 ♀ (CRBA005025), province of Taroudant, Tizi n ′ Test 30°49 ′ 40 ″ N, 8°23 ′ 57 ″ W, 24.ii.2010 (A. Calatayud-Mascarell & A. Sánchez-Vialas leg.). 1 juv. (CRBA005026), province of Agadir, Assaka, 30°50 ′ 18 ″ N, 9°45 ′ 53 ″ W, 28.xii.2018 (A. Sánchez-Vialas leg.). 1♂, 1 juv, (CRBA005030, CRBA005031), province of Guelmim, Playa Blanca, 28°57 ′ 39 ″ N, 10°33 ′ 03 ″ W, 23.iv.2019 (A. Sánchez-Vialas leg.). 1 juv. (CRBA005037), province of Taroudant, Alto Atlas, 30°53 ′ 59 ″ N, 8°19 ′ 35 ″ W, 24.ii.2020 (A. Calatayud-Mascarell & A. SánchezVialas leg.), 1♂ (SMNS-Aran-3824), province of Azilal, Demnate, 31°45 ′ 03 ″ N, 7°05 ′ 33 ″ W.

Justification for the revalidation of the species: Integration of evidence provided in this work has revealed that the ‘south’ lineage of the valentinus morphotype clade represents a different species from the ‘north’ lineage, which corresponds to the nominal I. valentinus . Simon (1909) described the female of I. mogadorensis from the surroundings of Essaouira ( Mogador), but the type material could not be examined because it is presumably lost ( Guadanucci & Wendt, 2014). The original description lacks any diagnostic traits other than a ‘pinkish-brown coloration’ ( Simon, 1909: 10). The type locality is Mogador (today Essaouira) but can refer broadly to the surroundings of the city. We lacked samples from the direct vicinity of Essaouira, but our nearest locality, Tamanat, is just 60 km south. Additionally, the SDM of I. valentinus ‘south’ lineage predicted its distribution well into coastal plains up to Essaouira (see Fig. 13B ). Therefore, we propose restoring the name I. mogadorensis and revalidating the species.

Diagnosis: Males can be distinguished from their congeners by the presence of two short, strong spines apically on the ventral part of tibia I and a moderate number of remaining spines distributed equally across tibia I ( Figs 8 , 14B–D ). They further differ from I. valentinus by their smaller size (N = 3) and markedly S-shaped embolus ( Figs 7 , 14F–J, L, M ).

Females differ from all other Ischnocolus species, except I. jickelii , by the spermatheca shape, which is longer than wide, narrowing apically and having three cross-like shaped apical lobes ( Figs 5 , 15F ).

Description: Male (CRBA004987, Ankrime): total length 12.93. Colour pattern: Colour in ethanol:

legs and carapace light yellow-brown. Carapace with silver hairs ( Fig. 14A ). Abdomen darker brown with dorsal light, striped pattern. Colour of live specimen ( Fig. 16 ): variable, depending on the life stage. Legs and carapace golden brown, chelicerae golden grey, abdomen dark with lighter striped pattern ( Fig 14B, C ). After moulting, specimens can be dark. Carapace: 5.32 long, 4.46 wide; cephalic region almost flat from lateral view; eye tubercle low 0.49 long, 0.85 wide;

fovea with shallow depression, slightly recurved ( Fig. 14A ). Clypeus 0.15 wide. Eyes ( Fig. 14E ): AME 0.15, PME 0.16, ALE 0.27, PLE 0.17; PME-PME 0.46, ALE-AME 0.23, ALE-PLE 0.26, AME-PLE 0.34, AMEAME 0.27, ALE-ALE 0.67. Sternum, labium and maxillae: sternum 2.4 long, 2.02 wide, setose, with three inconspicuous sigilla on each margin opposite to coxa I, II and III. Labium twice as wide as long, 0.37 long, 0.86 wide, with two cuspules. Maxillae with approx. 40 cuspules ( Fig. 14K ). Abdomen: 6.30 long, 3.23 wide; long setae scattered across the dorsal part of abdomen with a group of long black setae in the anterior part. PLS basal segment 0.87 long, median segment 0.67 long, apical segment digitiform 0.75 long. Chelicerae: 2.31 long, basal article with nine teeth intercheliceral tumescence present, few long bristles along the margin. Pedipalps: length: 7.76 (femur 2.71, patella 1.80, tibia 2.12, tarsus 1.13). Furrow on ventral tibia short, slightly sigmoid. Spination: femur (p)1ap. Copulatory bulb: small bulb with remarkably S-shaped embolus with flattened tip ( Figs 7 , 14 ). Legs: scopula on all tarsi divided by a thick band of setae. Scopula on ventral metatarsus I nearly totally occupied, II half occupied, III and IV <half occupied. Leg measurement: length of legs IV> I> II> III; leg I: 15.98 (femur 4.72, patella 2.14, tibia 3.64, metatarsus 3.18, tarsus 2.23), leg II: 15.12 (femur 4.13, patella 2.09, tibia 3.61, metatarsus 3.16, tarsus 2.13), leg III: 14.30 (femur 3.78, patella 1.83, tibia 3.51, metatarsus 3.12, tarsus 2.04), leg IV: 19.15 (femur 4.93, patella 2.04, tibia 4.50, metatarsus 4.90, tarsus 2.76). Spines: I femur (p)1, tibia (p)2-2, (v)3-1- 1-ap2, tarsus(v)1-1. II femur (r)1, tibia (r)1-1-1, (v)1-1- 1-2, metatarsus(r)1-0-0, (p)1-0-0, (v)1-1; III femur (r)1, tibia (r)1-1,(v)1-1-1-3, metatarsus (r)1-0-1, (v)1-1; IV femur (r)1, (p)1, patella (p)2, tibia (r)1-2-1-1, (p)1-1-1, (v)1-1-2, metatarsus (r)1-1-2-1-1-1, (p)1-1-1-1-1, (v)1- 1-2; tarsus IV pseudosegmented.

Female (CRBA004989, Ankrime): Total length 15.86. Colour pattern: Colour in ethanol: carapace, chelicerae and legs light orange, abdomen greybrown with light striped pattern ( Fig. 15A, E ). Colour of live specimen: body uniformly dark-brown-pinkish. Abdomen with light, striped pattern ( Fig. 15A–C ). Carapace: 5.51 long, 4.34 wide; cephalic region almost flat from lateral view ( Fig. 15E ); eye tubercle low, 0.50 long, 0.87 wide. Fovea with shallow depression slightly recurved ( Fig. 15A ). Clypeus 0.22. Eyes ( Fig. 15C ): AME 0.14, PME 0.13, ALE 0.26, PLE 0.19; PME-PME 0.50, ALE-AME 0.23, ALE-PLE 0.29, AME-PLE 0.35, AME-AME 0.28, ALE-ALE 0.68. Sternum, labium and maxillae: sternum 2.69 long, 2.18 wide; labium twice as wide as long, 0.45 long, 0.97 wide, with four cuspules; maxillae with approx. 40 cuspules ( Fig. 15B ). Abdomen: 8.35 long, 4.14 wide; PLS basal segment 0.82 long, median segment 0.51 long, apical segment 0.75 long, digitiform ( Fig. 15G ). Vulva: spermathecae longer than wide, each with three apical lobes in a cross-like disposition ( Fig. 15F ). Chelicerae: 2.93 long; basal article with 10 teeth; few long bristles on the margin ( Fig. 15D ). Pedipalps: length: 9.06 (femur 3.05, patella 1.89, tibia 2.20, tarsus 1.92). Spination: without spines. Legs: Scopula on all tarsi divided by a thick band of setae. Scopula on ventral metatarsus I and II entirely occupied, III three-quarter occupied, IV half occupied. Leg measurement: length of legs IV> I> II> II. Leg IV: 17.93 (femur 4.63, patella 2.54, tibia 4.09, metatarsus 4.28, tarsus 2.39); leg III: 12.45 (femur 3.28, patella 1.92, tibia 2.50, metatarsus 2.89, tarsus 1.86), leg II: 12.65 (femur 3.62, patella 2.20, tibia 2.64, metatarsus 2.40, tarsus 1.79); leg I: 14.24 (femur 4.12, patella 2.69, tibia 3.16, metatarsus 2.58, tarsus 1.69). Spines: I femur (p)0-0-1, tibia (p)0-1-1, (v)1-1- 2, (p)1-2-1, (v)1-2-2; II femur (p)0-0-1, tibia (p)1-1-1, (v)1-1-2, metatarsus (v)1-1; III femur (d)0-0-1r., tibia (r)1-1, (p)2-2-1, (v)1-1-2, metatarsus (r)2-2, (p)2-2-2, (v)1-1-1, patella (p)2; IV femur (r)0-0-1, (p)0-0-1, tibia (r)1-2-2-1, (p)1-2-1-2, (v)1-1-2, metatarsus (r)1-1-1- 1-1-1-1, (p)3-2-2-2, (v)1-1-1, patella (p)2; tarsus IV pseudosegmented.

Distribution: The species is endemic to southern Morocco and northern Western Sahara. Actual distribution includes western High Atlas (either the southern and northern slopes east to Demnate) and Anti-Atlas Mountain ranges (up to 2000 m a.s.l.), the coastal region south from Essaouira and the coastal part of southern Morocco down to Western Sahara ( Fig. 13B )

Natural history: The lifestyle is similar to that of I. valentinus and could be described as opportunistic. Some specimens were found to excavate shallow chambers under rocks covered with loose, sheet web, whereas others were found in deeper holes under rocks, or they inhabited the cavities between rocks and stones ( Fig. 17E, F ). Habitat preferences range from coastal forests of Barbary thuja, Tetraclinis articulata (Vahl) Mast. , habitats with dwarf fan palm ( Chamaerops humilis L.) ( Fig. 17A ), coastal Euphorbia L. communities ( Fig. 17D ), High Atlas Juniperus L. communities ( Fig. 17C ) to semi-desert habitats. The distribution model for this species overlaps with the extension of the Mediterranean Acacia – Argania Dry Woodland and Succulent Thicket ecoregion (PA1212), defined by the WWF (2021).

Figures

Figure 5. Spermathecae shape variation across the valentinus morphotype. A, B, ‘north’ lineage (I. valentinus); C, ‘south’ lineage (I. mogadorensis). Scale bar = 1 mm.

Figure 7. Bulbus shape comparison between males. 1, I. mogadorensis; 2, I. valentinus; 3, I. elongatus. A, proventral; B, dorsal; C, prolateral; D, ventral; E, retrolateral. Scale bar = 1 mm.

Figure 8. Comparison of the spine pattern on tibia I among males. 1, I. mogadorensis; 2, I. valentinus; 3, I. elongatus. A, ventral; B, prolateral; C, retrolateral. Scale bar = 1 mm.

Figure 9. Lateral view of abdomen showing variability in PLS length. A, I. valentinus; B, I. mogadorensis; C, I. elongatus ‘north’; D, I. elongatus ‘south’. Scale bar = 1 mm.

Figure 10. Lateral view of PLS showing shape variability of apical segment in: A, I. valentinus; B, I. mogadorensis; C, I. elongatus ‘north-east’; D, I. elongatus ‘north’; E, I. elongatus ‘central’; F, I. elongatus ‘south’. Scale bar = 1 mm.

Figure 13. Suitability maps obtained with MAXENT for I. valentinus (A), I. mogadorensis (B), I. elongatus ‘north’ (C), I. elongatus ‘central’ (D) and I. elongatus ‘south’ (E) for Morocco and the Iberian Peninsula.

Figure 14. Ischnocolus mogadorensis, male, A–N (CRBA004987).A, prosoma, dorsal view.B, tibia I, prolateral view. C, tibia I, ventral view. D, tibia I, retrolateral view. E, eye tuberlce, 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.

Figure 15. Ischnocolus mogadorensis, female, A–G (CRBA004989). A, prosoma, dorsal view. B, sternum, maxillae and chelicerae, ventral view. C, eye tubercle, dorsal view. D, chelicerae, prolateral view. E, whole body, lateral view. F, spermathecae. G, posterior lateral spinneret, retrolateral view. Scale bar = 1 mm.

Figure 16. Ischnocolus mogadorensis, habitus of living specimens. A, subadult female from Souss Massa region; B, adult female from Imsoane; C, subadult male from Boutazlaft; D, adult female from Souss Massa NP. Photo credits: E. Hijmensen (A), P. Fabiánek (B), J. Korba (C), R. Cavalcante (D).

Figure 17. The habitats and burrows of I. mogadorensis. A, locality Alto Atlas, 1310 m a.s.l., shrubby vegetation with Chamaerops humilis. B, mountains near Ankrime, 1150 m a.s.l., shrubby vegetation with Juniperus phoenicea. C, locality Tizi n′Test, 2010 m a.s.l., rocky habitat with Juniperus oxycedrus. D, locality Bou Tazlaft, semi-arid Macaronesian vegetation with Euphorbia balsamifera and E. officinarum. E, burrow between stones. F, entrance to subterranean burrow with only slight web cover. Photo credits: A. Sánchez-Viallas (A, C), J. Korba (B, D), E. Hijmensen (E, F).