Systematics, ecology and distribution of the mygalomorph spider genus Cteniza Latreille, 1829 (Araneae, Mygalomorphae, Ctenizidae) Author Decae, Arthur Author Mammola, Stefano Author Rizzo, Pierluigi Author Isaia, Marco text Zootaxa 2019 2019-01-29 4550 4 499 524 journal article 27469 10.11646/zootaxa.4550.4.2 6f2bd978-b3da-4c20-8211-a020355831e5 1175-5326 2625572 6371FA34-C296-4798-9C05-D52DAE0E8F9F Cteniza sauvagesi Rossi 1788 ( Figs. 1 , 3, 6 , 9, 12 , 15, 18 , 22 , 26 , 30 , 34, 36 , 37, 40–46 , 49–65 ) Aranea Sauvagii Rossi 1788 : 141 , figs VII–X (female); 1790: 218, plate X fig.11, (female holotype , failure to recover Rossi’s type material led us to consider it to be lost). Aranea Sauvagesii Latreille 1799 : 125 , plate 6, figs 2A–D (short description female probably on material collected by Rossi); 1802: 49; Westwood 1840: 170 (discussion of taxonomic placings). Mygale Sauvagesii Latreille 1804: 165 , plate LXIII, figs 7–10 (description female and natural history); 1818: 125–126 (ibid.); Dufour 1820: 102, plate 73, fig. 3 (ibid. ) new combination. Mygale sauvagesii Latreille 1806: 84 (bibliography). Mygale pionière ( M. fodiens ) Walckenaer 1805: 5 new synonymy. Mygale pionnière ( M. fodiens ) Olivier 1811(Ibid): 86; Lamarck 1818: 107 (ibid.); Walckenaer 1826: 4, plate 2, figs 1–2 (female, figs Mygale de Sauvage ); Latreille 1829 : 232 (description female); Dugès & Milne Edwards 1836: 35 (description female, burrow and mygale anatomy); Walckenaer 1837: 237–239 (burrow after Audouin). Ctenize pionière Latreille 1831: 507 new combination . Mygale fodiens Griffith 1833: 400 (discussion synonyms); Lucas 1840: 339–343, plate 2, fig. 1 (behavior); Saunders 1842: 160, 163 (note). Mygalodonte pionnière Simon 1864: 79 –80 (description burrow) new combination. Mygalodonta fondiens Simon 1864: 453 (list); Canestrini & Pavesi 1868: 22 (list); 1870: 3 (list). Nemesia fodiens Carruccio 1871: 55 , plate1–3 (first record from Sardinia ); Garneri 1902: 64 (list). Aëpycephalus brevidens Ausserer 1871 : 151 (description female after Pachyloscelis brevidens Doleschall undated, unpublished, manuscript); Simon 1892 : 95 (ibid.); Wunderlich 1995 : 358 new synonymy. Cteniza sauvagei Ausserer 1871 : 152 –153 (description female, indication type species, original designation); Simon 1873: 34– 35 (first description male, nomination type species), plate1,1 and 5; 1892: 70 (note), 95 (short description); 1903: 981 (key). C. sauvagesi : Simon 1914 : 4 , figs 1–5 (species diagnostics); Decae 1996 : 162 , figs 1–3 (genus diagnostics); Le Peru 2011: 72, fig. 32 (distribution); Opatova et al. 2013 : 1142 (phylogenetics); Colombo & Manunza 2013: 99–100, figs 1–2 (malacophagy). FIGURES 15–20. Top row left to right: 15, Ct. sauvagesi (female, Riz.006, Calabria); 16, Cy. cunicularium (female, 20/04/ 90.1, Crete); 17, Stasimopus sp. (female, AR4117, Bloemfontijn). Note open groups of “dagger-type” spines on the prolateral distal segments of leg I in Cteniza + Cyrtocarenum (15–16) versus the compact groups of short “curvy-spines” in Stasimopus (17). Bottom row left to right: 18, Ct. sauvagesi (female, Bra.005, Corsica); 19, Cy. cunicularium (female, 20/04/90.1, Crete); 20, Stasimopus sp. (female, AR4117, Bloemfontijn), note the cylindrical tibia III and proximally concave dorsal femur in Cteniza + Cyrtocarenum (18–19) versus the distally inflated tibia III and convex dorsal femur in Stasimopus (20).). Points of attention are marked with arrows. Scale bars= 1.0 mm Material examined. 1 female ( NEOTYPE ) id. Isaia.137, France Corsica , Cap Corse , Oreta , ( 42.86°N , 9.43°E ), leg. S. Mammola and A. Giresi ; 1 male (specimen described) id. NHMB-011, France , Corsica, Marine de Saint Ambrogio ( 42.6°N , 8.82°E ), leg. B. Salmini , B. and F. Giomi. 1 female id. TH.402, Calvi , Campo di a Veta ( 42.57°N , 8.72°E ) leg. K. Thaler and B. Knoflach ; 1 female id. TH.414A, Calvi , Forêt de Bonifacio ( 42.46°N , 8.85°E ) leg. K. Thaler and B. Knoflach ; 1 female id. Bra. 05, Corsica (no precise location) leg. B. Rast ; 2 females id. NHMP .020 and NHMP .021, Corsica (no precise location) leg. Unknown ; 3 females id. AR4215a–c, Corsica , region Ajaccio ( 41.93°N , 8.74°E ) leg. Unknown ; 2 females id. AR4211a–b, Corsica region Ajaccio leg. Dehant ; 1 female id. AR4216, Corsica Bastia , Ravin du Lupino ( 42.7°N , 9.45°E ) ; 1 female id. SMF3481 , Corsica (no precise location) leg. Unknown ; 1 male id. Colo. 015 Italy , Sardinia , Sassari , Maristella ( 40.59°N , 8.21°E ) leg. M.Colombo ; 1 male id. Colo. 018 Italy , Sardinia , Maristella ( 40.6°N , 8.22°E ) leg. M.Colombo ; 1 female id. Shu. 180, Sardinia , west of Cagliari ( 39.15°N , 8.64°E ) leg. S. Huber ; 1 female id. Colo. 17, Sardinia Maristella ( 40.6°N , 8.22°E ) leg. M. Colombo ; 1 female id. Colo. 16, Sardinia Collinas ( 39.64°N , 8.84°E ) leg. M. Colombo ; 1 juv. id. NHMB .03, Italy , Ponza , NW Mt. Guardia ( 40.9°N , 12.96°E ) leg. V. Cotterelli. 2 males id. Isaia. 133, 134, and 1 female id. Isaia-135, Italy , Calabria , Cosenza , Terranova da Sibari ( 39.66°N , 16.34°E ) leg. P. Rizzo ; 6 males id. Riz-01, 0 3, 0 4, 0 8, 0 9, 0 10, Italy , Calabria , Tarsia ( 39.61°N , 16.32°E ) leg. P. Rizzo ; 2 females id. Riz-02, 0 5 Italy , Calabria , Cosenza , Terranova da Sibari ( 39.66°N , 16.34°E ) leg. P. Rizzo ; 2 females id. Riz-02, 0 5 Italy , Calabria , San Lorenzo Bellizzi ( 39.8°N , 16.33°E ) leg. P. Rizzo. Type material and reference material from all locations will be deposited in MCSNEC (Bergamo). FIGURES 21–24. Top left to right: 21, Ct. moggridgei (female, Isaia.020 Liguria); 22, Ct. sauvagesi (female, Colo.017, Sardinia) Note the irregular distribution of macro-spigots pointed by arrows. Bottom left tyo right: 23, Cy. cunicularium (female, 15/04/90.4, Crete); 24, Cy. grajum (female, Isiai.016a, Kefalonia). Note the compacte groups of macro-spigots in Cy. cunicularium and the more slender PLS with conical distal segment in C. grajum . Scale bars= 1.0 mm FIGURES 25–28. Genus level diagnostic differences between Cteniza and Cyrtocarenum in the male bulb structure (standard ventral orientation). Left to right: 25, Ct. moggidgei (SHu.19.06.14, Alpes Maritimes); 26, Ct. sauvagesi (Isaia.134, Calabria); 27, Cy. cunicularium (GR80.11, Tinos); 28, Cy. grajum (GR.8.82.5, Kithira). Note the regular curve and tiny apical scoop (SC) in the embolus of the two Cteniza species and the robust distally bent and twisted embolus (tw) in the two Cyrtocarenum species. Scale bars= 0.5 mm FIGURES 29–32. Genus level diagnostic differences between Cteniza and Cyrtocarenum as seen in the armature and shape of the retrolateral male tibia and proximal metatarsus of leg I. 29, Ct. moggridgei (Shu-19.06.14 Alpes Maritimes); 30, Ct. sauvagesi (NHMB.011, Corsica); 31, Cy. cunicularium (GR80.11, Tinos); 32, Cy. grajum (GR.8.82.5, Kithira). Note strong spines (ss) in ventral half of the tibia and metatarsus as the sole armature in Cteniza and the strong ventral apical clasper-hooks mounted on a distinct process (ch) in Cyrtocarenum . Note also the difference in tibiae shape between the genera and the proximal bent and sclerotized metatarsus (sb) in Cyrtocarenum (absent in Cteniza ). Scale bars= 1.0 mm FIGURES 33–36. Living spiders of the genus Cteniza . 33, Ct. moggridgei (female, 20/04/05.4, Gorbio Alpes Maritimes); 34, Ct. sauvagesi (female, field recording by P. Rizzo Calabria); Ct. moggridgei (male, 20/04/05.5, Gorbio Alpes Maritimes); Ct. sauvagesi (male, field recording by P. Rizzo Calabria). Diagnosis. Cteniza sauvagesi differs from C. moggridgei in the larger size, as measured on the carapace length of both adult females and males. Maximal lengths recorded for C. moggridgei : females (n= 8) CL= 6.4, males (n= 8) CL= 6.2. Minimal lengths recorded for C. sauvagesi : females (n= 21) CL= 7.8, males (n= 11) CL= 7.3. The records for C. moggridgei include measurements given in Buchli (1968) . Males of C. sauvagesi differ from C. moggridgei in the shape of the carapace, anterior narrowing in C. sauvagesi ( Fig. 43 ) anterior rounded in C. moggridgei ( Fig. 44 ), and in the spine patterns on leg I, with a dense group of short spiky bristles on the prolateral femur in C. sauvagesi (absent in C. moggridgei ), and relative stronger development of spines on the prolateral patella and tibia in C, moggridgei ( Figs. 45–48 ). The two species are remarkably similar in sexual morphology (compare Figs. 59–62 and 66–69 ). Finally, the two species are geographically and ecologically separated (see section on SDM). Description. Female neotype ( Figs. 49–50 ). General appearance: robustly built, short legged spider with elevated, caudally truncated cephalic part of the carapace, deep procurved fovea and ovoid abdomen. Legs III and IV stronger than legs I and II, palps leg-like. Color: dorsal prosoma and legs almost uniform dark somewhat greenish brown, ventrally lighter brown, chelicerae darker reddish brown, abdomen dark grey dorsally lighter. Carapace (as Fig 41 ): shiny, glabrous except for some fine bristles around the eye-group and on the crest of the cephalic part, posterior edge concave. Eye-group (as Fig. 37 ): slightly longer than half the its width (EL/PR= 0.6), trapezoidal with posterior row slightly longer than anterior row (AR/PR= 0.9). ALE largest, PME larger than PLE, AME further than their diameter apart. Chelicerae: very strong, fine bristles dorsally in longitudinal zone, distally in triangular group, rastellum compact group of strong rigid short spiky spines on the inner apex of the basal segment, partly placed on distinct rastellar process, cheliceral furrow lined on both sides with irregular row of teeth (prolateral row extending forward), few smaller denticles on furrow bottoms, fangs, strong, proximally hooked, ventrally flattened between two sharp ridges. FIGURES 37–40. Top row: variable, sub-rectangular to trapezoidal, eye-formations in male and female of the genus Cteniza . 37, Ct. sauvagesi (male, Colo.018, Sardinia); 38, Ct. moggridgei (females, Isaia.020, Liguria). Bottom row: Patterns of sternal sigilla in male and female of the genus Cteniza . 39, Ct. moggridgei (male, Shu.19.06.14, Alpes Maritimes); 40, Ct. sauvagesi (female, Riz.006, Calabria). Note the large central sigilla (cs) and the small lateral sigilla (ls). Points of attention are marked with arrows. Scale bars= top-row= 0.5 mm, bottom row 1.0 mm Sternum (as Fig. 40 ): evenly spread fine erect bristles, large sub-central sigilla, small lateral sigilla. Labium (as Fig. 40 ): dome shaped with few distal cuspules, separated from sternum by wide, shallow labial furrow. Maxillae: short, rounded, distal apophyses, pinkish scopula along the ventral-prolateral edge, irregular double row of cuspules along proximal edges. Palps: leg-like, groups of strong lateral dagger-type spines on tarsus, tibia and a single prolateral patella spine. Legs I and II: with dagger-type spines in the ventral halves of the prolateral and retrolateral distal three segments (as Fig. 15 ). Leg III (as Fig. 18 ): shortest and strongest of legs with spine-groups on distal tarsus, dorsal half metatarsus, and prolateral patella and a single distal spine on prolateral tibia, ventral tarsus with thin scopula. Leg IV: longest of legs with few fine spines on the prolateral tarsus and metatarsus, retrolateral spines absent, ‘rake- like’ structure around the dorsal femur-patella joint. Claws: palp-claw one large curved proximal bifid tooth ( Fig. 51 ), Leg I ATC slightly sigmoid, STC with large proximal side teeth, few tiny subsidiary teeth ( Fig. 52 ), Leg IV ATC regularly curved, STC with bifid side tooth on posterior claw ( Fig. 53 ). Abdomen: ovoid, dense cover of fine bristles. FIGURES 41–42. Sexual dimorphic carapace structure in lateral view within Cteniza . 41, Ct. sauvagesi (female, AR4224, Corsica); 42, Ct. sauvagesi (male, Riz.008). Note high elevated cephalic part steeply sloping to the fovea and smooth, shining cuticle in female (41) and finely sculptured, dull cuticle with thick lateral/caudal rim and reduced elevation of the cephalic part in male. Scale bars= 2mm Spinnerets (as Fig. 22 ): PMS club shaped, apical spigot field, PLS three segmented, basal segment longer than median plus distal segment, spigots concentrated in the distal parts of all segments. Spigot differentiation distinct with a single larger spigot on apical PMS and a few conspicuously larger spigots on all PLS segments, particularly prominent on the distal parts of the median segment. Spermathecae ( Fig. 54 ): “mushroom-shaped”, tripartite receptacles with distally narrowing, membranous, flattened proximal part, sclerotized medial part and “donutshaped” glandular distal part. Measurements. Female neotype (id. Isaia-137). TBL= 28.8, CL= 9.2, CW= 7.9, SL= 5.7, SW= 4.3, LL= 1.2, LW= 1.6, AR= 1.51, PR= 1.75, EL= 0.96, dALE= 0.44, dPLE= 0.25, dAME= 0.15, dPME= 0.22, disAME= 0.27, disPME= 0.75, PaL= 12.9, LegI= 16.0 LegII= 14.6, LegIII= 14.8, LegIV= 20.3, Int= 0.39, Ext= 0.89, LRL= 0.46, LRR= 0.55. Description. Male ( id. NHMB-011). General appearance ( Fig. 36 ): robustly built, long legged spider with small ovoid abdomen and slender, strongly elongated palps. Color (in ethanol): dorsal view prosoma and legs almost uniform dark brown, chelicerae slightly lighter brown, abdomen purplish, mottled, spinnerets yellow. Ventral sternum yellow with slightly darker edges and sigilla, labium and ventral coxae light brown, maxillae light brown with anterior yellow zone, legs and chelicerae slightly darker brown, anterior abdomen creamy color, posterior abdomen greyish, spinnerets white. Carapace: finely sculptured, almost glabrous, few thin bristles around eye-group (partly on narrow clypeus), thick rim (as Fig. 43 ), cephalic part only slightly elevated, gradually curving down to fovea (as Fig. 42 ), fovea deep pro-curved. Eye-formation: as in female. Chelicerae: weaker with smaller rastellum than in female, Sternum: with more pronounced curvature of the edge compared to female. Labium and Maxillae without cuspules. FIGURES 43–44. Species level diagnostics, differences in the shape of the male carapace. 43, C. sauvagesi (Colo.015, Sardinia); 44, C. moggridgei , (Shu.19.06.14, Alpes Maritimes). Note the anterior distinctively narrowing carapace in C. sauvagesi versus the rounded carapace in C. moggridgei . Palp ( Fig. 55 ): glabrous with only locally groups of short hairs, strongly elongated tibiae, patellae, femora, trochanters. Cymbium ( Fig. 56 ): anterior rounded with distal group of dorsal spines (mostly broken-off in the described specimen). Legs: first leg ( Fig. 45–46 ) stronger than other legs with numerous strong spines on pro- and retrolateral faces of tibia, and on prolateral patella. Prolateral femur with a field of short strong spiky bristles ( Fig. 45 ). Claws: STC with single sigmoid curved rows (combs) of teeth ( Figs. 57–58 ). Abdomen: small ovoid, dense cover of fine bristles. Spinnerets: differentiation of spigots less pronounced than in females. Bulb ( Figs. 59–62 ): simple pyriform with regularly curved slender, distally narrowing embolus with a tiny ‘scoop’ at the embolus tip. Measurements. TBL= 15.8, CL= 7.7, CW= 6.8, SL= 4.7, SW= 3.5, LL= 0.7, LW= 1.1, AR= 1.29, PR= 1.45, EL= 0.74, dALE= 0.32, dPLE= 0.23, dAME= 0.15, dPME= 0.23, disAME= 0.22, disPME= 0.66, PaL= 16.2, LegI= 20.8, LegII= 19,1, LegIII= 17.3, LegIV= 24.3, BuL= 1.91, EmL= 0.86, BuW= 0.80. Size variation. Although our samples of C. sauvagesi from geographically separated populations on Corsica , Sardinia and in Calabria are small, we did not find distinctive size differences between males (CL: av= 8.1, sd= 1.0, n= 11) and adult females (CL: av= 9.2, sd= 1.0, n= 21) collected in different regions. With respect to qualitative differences in morphological characters, Ausserer (1871: 153) reported substantial variation within the Corsican population of C. sauvagesi . We studied this variation in our samples to conclude that some of Ausserer’s characters are difficult to interpret, either because we have no information about his methods of measurement (e.g. length/width ratio of the rastellar process) or the ambiguity in interpretation of character states (e.g. fovea shape, shallower or deeper grooves in the carapace). Where we could follow Ausserer (1871) , we found the same variation in sternum shape and eye configurations in all populations ( Corsica , Sardinia Calabria), supporting the specific unity of different populations of C. sauvagesi . The only potentially diagnostic character we found to distinguish isolated populations of C. sauvagesi was the spine patterns on lateral faces of tarsus I in males ( Fig. 63–65 ). Here, prolateral and retrolateral spines were found in the Calabrian population (n= 8); such spines were absent in the Sardinian and Corsican populations (n= 3). The variation and differences in morphology found are regarded to be insufficient to justify the recognition of different species occurring on Corsica , Sardinia and Calabria. Hence, we regard C. sauvagesi as a single species distributed in at least three currently marine isolated geographical regions. FIGURES 45–48. Species level diagnostics, differences in spine patterns on leg I in males 45, C. sauvagesi (right, prolateral, NHMB.011, Corsica); 46, C. sauvagesi (right, retrolateral, NHMB.011, Corsica); 47, C. moggridgei (right, prolateral, 20.04.0 5.5, Alpes Maritimes); 48, C. moggridgei (right, retolateral, 20.04.0 5.5, Alpes Maritimes). Note the field of spikybristles (sb) on prolateral femur in C. sauvagesi (45) absent in C. moggridgei (47), and the relatively stronger spines on prolateral patella (ps) and tibia (ss) in C. moggridgei (47–48). Size variation measurements. Males (n= 5): TBL= 14.6–18.2, CL= 7.7–8.8, CW= 6.8–7.7, SL= 4.3–5.2, SW= 3.4–4.5. AR= 1.28–1.41, PR= 1.45–1.77, EL= 0.74–1.05, dAME= 0.15–0.19, disAME= 0.22–0.30, PaL= 15.6–17.4, LegI= 20.8–23.1, LegII= 19.1–21.7, LegIII= 17.1–20.1, LegIV= 24.3–28.2, BuL= 1.98–2.21, EmL= 0.89–1.13, BuW= 0.80–0.95. Females (n= 11): TBL= 17.2–27.8, CL= 7.8–10.2, CW= 7.0–9.3, SL= 5.1–6.8, SW= 3.7–5.2, AR= 1.43–2.10, PR= 1.49–2.37, EL= 0.93–1.26, dAME= 0.14–0.20, disAME= 0.23–0.42, PaL= 12.3– 16.3, LegI= 14.6–20.5, LegII= 13.4–18.4, LegIII= 14.0–18.4, LegIV= 18.3–25.5. Remarks. The validity of C. brevidens has never been confirmed. A study of detailed photographic material of the supposed holotype of C. brevidens , kindly provided by Christoph Hörweg (NHMW-Vienna, Inv. No. 11, “TYPUS”, leg. Koelbel) and a study of a specimen labeled C. brevidens by Costa in the collection of the MNHN- Paris (A.D. personal observation), indicated that only the shape of the eye-formation would distinguish C. brevidens from C. sauvagesi (see Decae 1996 : figs 1–2). Significant variations in the general shape of the eyeformation in C. sauvagesi were first reported by Ausserer (1871: 152-153) and are confirmed in this study (PR/EL, n= 16, range 1.49–2.26, av.= 1.86, sd.= 0.18, AR/PR, n= 16, range 0.77–0.90, av.= 0.85, sd. 0.04). Here we judge these differences insufficient to discriminate species and hence regard C. brevidens (Doleschall, 1871) as a junior synonym of C. sauvagesi ( Rossi, 1788 ) n. syn .