Ischnocolus valentinus, (DUFOUR, 1820)

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 : 866-870

ISCHNOCOLUS VALENTINUS ( DUFOUR, 1820)

( FIGS 5A–B , 7 , 8 , 9A , 10A , 13A , 18A–N , 19A–G, 20A–F)

Mygale valentina Dufour (1820: 101 ; description of male).

Mygale valenciana: Walckenaer (1837: 228) .

Trechona valentina: Thorell (1870: 168) .

Ischnocolus holosericus L.Koch , in Ausserer (1871: 186; description of juvenile); Simon (1892: 136, fig. 119); Bacelar (1932: 171); Smith (1990: 129, figs 819–829; description of female).

Ischnocolus triangulifer Ausserer (1871: 186 ; description of juvenile); Bacelar (1932: 171).

Avicularia andalusiaca Simon (1873: 197 , pl. 1, fig. 2; description of male and female); Bacelar (1932: 171, figs. 3–5).

Ischnocolus algericus Thorell (1875: 123, description of male). Simon (1903: 925, figs. 1070–1071, m).

Ischnocolus fuscostriatus Simon (1885: 41 ; description of male).

Leptopelma cavicola Simon (1889: 396 ; description of male and female, burrow structure; 1909: 8); Reimoser (1919: 7); Roewer (1942: 222); Bonnet (1957: 2395); Benoit (1964: 414, figs 1, 2; male and female).

Ischnocolus maroccanus Simon (1873: 199 ; description of male and female).

Ischnocolus numidus Simon (1909: 9 ; description of male).

Ischnocolus tripolitanus Caporiacco (1937: 57 ; description of female).

Ischnocolus valentinus: Ausserer (1871: 186) ; Guadanucci & Wendt (2014: 391; description of male and female); Zonstein (2018: 114; description of male) (synonymy of Nemesia cavicola ); Montemor et al. (2020: 89, fig. 9C, D); Tamajón Gómez et al. (2020: 165).

Type material: Type locality Moixent , Valencian Community, Spain. Female neotype deposited at BMNH, examined .

Material examined: Spain: 1♂, 1♀ ( CRBAMM000941 , CRBAMM000930), province of Cádiz , El Bosque, 36°26 ′ 50 ″ N, 5°22 ′ 19 ″ W, 27.iii.2010 (M. A. Ferrández leg.) . 3♀♀ ( CRBAMM000965 , CRBAMM000966, CRBAMM000967), province of Cádiz , Grazalema, 36°27 ′ 28 ″ N, 5°17 ′ 34 ″ W, 28.iii.2010 (M. A. Ferrández leg.) . 1♀ ( CRBAMM000948 ), province of Sevilla , Castaño, 36°33 ′ 58 ″ N, 5°15 ′ 14 ″ W, 27.iii.2010 (M. A. Ferrández leg.) . 1♂, 1♀ ( CRBAMM 0 0 0 9 8 0, CRBAMM000986), province of Sevila , Coripe, 36°35 ′ 20 ″ N, 5°17 ′ 34 ″ W, 27.iii.2010 (M. A. Ferrández leg.) . 1♀ ( CRBAMM000141 ), province of Almería , La Rambla del Aljibe, 37°16 ′ 20 ″ N, 2°02 ′ 24 ″ W, 21.XI.2009 (E. Planas & V. Opatova leg.) . 1 juv. ( CRBAME000858 ), province of Málaga , Torcal de Antequera, 36°57 ′ 43 ″ N, 4°31 ′ 06 ″ W, 12.VI.2011 (E. Mora, V. Opatova & P. Sousa leg.) . Morocco: 2♀♀, 5 juv. ( CRBAMM000386 , C R B A M M 0 0 0 3 8 7, C R B A M M 0 0 0 3 8 8 C R B A M M 0 0 0 3 8 9, C R B A M M 0 0 0 3 9 0 CRBAMM000391, CRBAMM000392), province of Azilal , near Ouzoud falls, 31°57 ′ 35 ″ N, 6°46 ′ 05 ″ W, 10.IV.2010 ( V. Opatova & M. Arnedo leg.) . 2♀♀, 1♂, 3 juv. ( CRBAMM000393 , CRBAMM000395, C R B A M M 0 0 0 3 9 6, C R B A M M 0 0 0 3 9 8 CRBAMM000399, CRBAMM 000400), province of Azilal , Tilouguite, 32°05 ′ 04 ″ N, 6 °20 ′ 00 ″ W ,

10.IV.2010 ( V. Opatova & M. Arnedo leg.). 1 juv. ( CRBAMM000450 ), province of Fés , Djebel Zalach, 34°06 ′ 23 ″ N, 4°58 ′ 10 ″ W , 12.IV.2010 ( V. Opatova & M.Arnedo leg.). 1 juv. ( CRBAMM000485 ), province of Oujda , Ain Sfa, 34°49 ′ 28 ″ N, 2°05 ′ 12 ″ W , 14.IV.2010 ( V. Opatova & M. Arnedo leg.). 3 juv. ( CRBAMM000564 , CRBAMM000565, CRBAMM000566), province of Tetuán , Beni Yder Cherki, 35°23 ′ 08 ″ N, 5°31 ′ 20 ″ W , 17.IV.2010 ( V. Opatova, M. Arnedo leg.). 2 juv. ( CRBAMM000493 , CRBAMM000583), province of Berkane , Beni Snassen, 34°48 ′ 12 ″ N, 2°23 ′ 48 ″ W , 1 4.IV.2 0 1 0 ( V. Opatova & M. Arnedo leg.). 1♀ ( CRBAMM 000366), province of Al Haouz, Asni, 31°11 ′ 22 ″ N, 8°03 ′ 27 ″ W, 9.IV.2010 ( V. Opatova & M. Arnedo leg.). 1♂ (SMNS-Aran-1376), province of Murcia, Mula, 38°03 ′ 41 ″ N, 1°31 ′ 05 ″ W, 2005. 1♂ ( CRBA) province of Berkane, Taforalt, Grotte des Pigeons , 34°48 ′ 52 ″ N, 2°24 ′ 10 ″ W, 15.iv.2002 (M. Arnedo & C. Hernando leg.) .

Diagnosis: Males can be distinguished from their congeners, except I. jickeli , by bearing only one spine apically along with dense concentration of spines on pro-ventral tibia I ( Figs 8 , 18B–D ). They further differ from all other Ischnocolus species by having straight embolus with narrowing tip ( Figs 7 , 18J–M ). Females differ from their congeners by having spermathecae as wide as long with several (> four) apical lobes ( Figs 5A, B , 19F ).

Description: A detailed redescription is provided by Guadanucci & Wendt (2014) with male and female reproductive organ drawings and male spine pattern on tibia I. Additional photos and drawings of male bulbus are shown by Zonstein (2018), Tamajón Gómez et al. (2020) and Decae (in: Nentwig et al., 2020).

Distribution: Following our circumscription of I. valentinus to the ‘north’ lineage, the distribution of the species is more restricted than previously thought, as it only includes the Iberian Peninsula and northern Morocco ( Fig. 13A ). The northernmost current localities are near Benidorm, in Alacant, Spain and the southernmost ones are located on the northern slopes of the High Atlas Mountains in Morocco. Previous studies have suggested the species occurrence extending as far east as Sicily, Tunisia and Libya ( Guadanucci & Wendt, 2014). Although the only Sicilian sample included in our phylogeny was recovered within the ‘north’ lineage (i.e. I. valentinus ), detailed morphological and molecular analyses of a more thorough specimen sampling in Algeria, Tunisia and Sicily will be required to confirm the status of the easternmost populations.

Natural history: Similar to I. mogadorensis , I. valentinus is an opportunistic species inhabiting natural cavities under and between rocks and tree roots. It usually covers the entrance with a dense, sheet web, sometimes resembling that of a funnelweb spider Macrothele calpeiana Walckenaer, 1805 or some Agelenidae . The burrow entrance can also sometimes resemble that made by I. elongatus , but the web of I. valentinus is less dense and more sheetlike ( Fig. 20E, F ). The burrow of I. valentinus never forms a proper tube, but continues as an irregular hole connecting one or several natural underground cavities. Habitat preferences range from Mediterranean open grasslands to dense Aleppo pine ( Pinus halepensis Mill. ) or Barbary thuja ( Tetraclinis articulata ) forests.

Figures

Figure 1. Map of sampled localities in Morocco and the Iberian Peninsula. Numbers correspond to localities as in the Supporting Information, Table S1.

Figure 2. Time-calibrated phylogeny of family Theraphosidae inferred by BEAST using six genes (12S, 16S, COI, 18S, 28S and H3). Boxes indicate node supports from BEAST (posterior probability, PP), maximum likelihood (ML, ultrafast bootstrap proportions UFBoot), MRBAYES (BI, posterior probability) and parsimony (MP, bootstrap proportions BS). Supports are summarized as follows: black = supported (PP> 0.95, UFBoot> 0.95, BS> 0.70), grey = recovered but support values below thresholds, white = not recovered. The axis is in millions of years.

Figure 3. Time calibrated phylogeny of western Mediterranean Ischnocolus inferred by BEAST using two concatenated genes (COI + ITS2) and values inferred from Theraphosidae analysis as COI rate priors. Boxes indicate node supports from BEAST (posterior probability, PP), maximum likelihood (ML, ultrafast bootstrap proportions UFBS) and MRBAYES (BI, posterior probability). Black = supported (PP> 0.95, UFBS> 0.95), grey = recovered with support values below thresholds, white = not recovered. Scale axis is in millions of years. Photo credits: valentinus morphotype – T. Romanov, elongatus morphotype – J. Korba. The Ischnocolus lineages are named and coloured as in Figure 2.

Figure 4. COI and ITS2 gene trees inferred by BEAST with bars representing three species delimitation methods: ABGD, mPTP and GMYC, respectively. Each individual block represents a delimited unit. Split blocks within ABGD and mPTP analyses indicate incongruences among phylogenetic inference and species delimitation analysis. Circles indicate support levels for the corresponding split in mPTP: black> 0.60, grey = 0.59–0.20 and no circle <0.20.

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 6. Spermathecae shape variation across lineages of the elongatus morphotype. Lineages are grouped as follows: A-C, ‘north-east’; D-F, ‘north’; G-I, ‘central’; J-L, ‘south’. 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 18. Ischnocolus valentinus, male, A–N (CRBAMM000941). 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.

Figure 20. Ischnocolus valentinus, habitus of living specimens and burrow. A, adult male from Ouzoud, northern Morocco; B, adult female from Ouzoud, northern Morocco; C, adult female from Ouzoud, northern Morocco; D, adult female from Alhurin del Grande, Andalucia, southern Spain; E, burrow entrance under tree; F, burrow under the stone. Photo credits: J. Korba (A, B, C, E, F), T. Romanoff (D).

Figure 19. Ischnocolus valentinus, female, A–G (CRBAMM00387). 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.