Leiurus heberti, Lowe & Yağmur & Kovařík, 2014

Lowe, Graeme, Yağmur, Ersen Aydın & Kovařík, František, 2014, A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula, Euscorpius 191, pp. 1-129 : 61-73

Leiurus heberti sp. n.

( Figs. 47–57 , 58B , 59G–H , 87E , 88E , 89E , 90E , 91C , 92D , 93E , 95B–D , 98–100 , Tabs. 2 , 3B ) http://zoobank.org/urn:lsid:zoobank.org:act:2E5906 DA-1108-4669-B2B7-8831B60EA4D1

HOLOTYPE. Adult ♂, Oman, Wadi Andur, UV detection, rocky boulder wadi, along base of cliff, east edge of wadi , 17°40.08'N 54°39.27'E, 405 m a.s.l., 26.IX.1995, 22:50 h, leg. G. Lowe & M.D. Gallagher ( NHMB).

PARATYPES. Oman: 1 adult ♂, 1 immature ♂, Wadi Andur, UV detection, rocky boulder wadi, along base of cliff, east edge of wadi , 17°40.08'N 54°39.27'E, 405 m a.s.l., 26.IX.1995, 22:50 h, leg. G. Lowe & M.D. Gallagher ( NHMB); 1 adult or subadult ♀, Jabal Samhan , 15 km W of Jufa, 17°11.169’N 54°56.571'E, 380 m a.s.l., 24.IX.2011, leg. W. Grosser ( FKCP) .

DIAGNOSIS (adults). Medium to large Leiurus , 73–83 mm in length, carapace L 8.4–8.9 mm; base color yellow, carapace darkly pigmented on interocular area and carinae, tergites with weak, variable fuscosity or pale; pace between anterior median carinae with sparse medium or fine granules; area between posterior median carinae with shallow median furrow flanked by lateral arcs of medium or fine granules; medial intercarinal surfaces of tergites II–III sparsely granular, finely shagreened or smooth; posterior margin of coxa III smooth; metasoma very long, slender, metasoma II L/W ♂ 1.91– 1.95, ♀ 1.74, metasoma III L/W ♂ 2.19–2.30, ♀ 1.92, metasoma IV L/W ♂ 2.70–2.94, ♀ 2.30; ventromedian carinae of metasoma II and III with 36–46 (♂) or 20–25 (♀) denticles; ventrolateral carinae of metasoma V with small triangular denticles in male, enlarged subtriangular or lobate denticles in female; pedipalps very slender, patella L/W ♂ 4.28–4.48, ♀ 3.09; leg III patella L/D ♀ 4.35–4.54, ♂ 3.23; pectine teeth ♂ 36–37, ♀ 33–34; pectines long, narrow, pectine L/ carapace L ♂ 1.38– 1.44, ♀ 1.05, mid- pectine sensillar margin L/metasoma I W ♂ 0.198 –0.225, ♀ 0.114; basal 1–3 pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; fulcra mostly with 3–4 setae; pectine basal piece smooth except for anterior median patches of fine granules; leg III basitarsus with 8–11 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db distal to est; sternite VII with area between median carinae smooth or very faintly shagreened anteriorly; sternite carination: males, sternite III with median carinae strong, sternites IV–V with lateral carinae strong, median carinae moderate to strong; females, sternite III with median carinae obsolete, sternites IV–V with lateral carinae weak, median carinae obsolete.

COMPARISONS. L. heberti sp. n. appears most closely related to L. haenggii sp. n. from Yemen, southwestern Oman, and western Saudi Arabia. The two species share similar carination and granulation of the carapace, tergites and sternites, with transverse arcs of granules between posterior median carinae on the carapace, and moderately developed basitarsal bristle combs. L. haenggii sp. n. differs from L. heberti sp. n. as follows: (i) males with less elongated pedipalps, legs and metasoma II–V; (ii) carinae on metasoma I–IV more robust with coarser granulation; (iii) males with larger, lobate dentition on ventrolateral carinae of metasoma V; (iv) dark pigmentation typically extends over posterior area of carapace and most of tergites; (v) lower range of pectinal tooth counts. L. jordanensis Lourenço, Modrý et Amr, 2002 , from Jordan and northern Saudi Arabia also has long, slender pedipalps and metasomal segments, but can be differentiated from L. heberti sp. n. as follows: (i) different color pattern, with dark brown carapace, pedipalps, tergites and metasoma, and contrasting yellow telson and pedipalp fingers; (ii) larger size, adult female carapace length up to 11.4 mm; (iii) telson with vesicle elongated, not bulbous; (iv) lower range of pectinal tooth counts: ♂ 33–37, ♀ 27–31; (v) basitarsi I–III with 10–20 retrosuperior setae.

ETYMOLOGY. A patronym in honor of Blaine Hébert, California, for his friendship and support of arachnology, science education and music.

DESCRIPTION (holotype male).

Coloration. Base color pale orange-yellow; carapace with dark pigmentation on interocular triangle, and around median ocular tubercle; carinae of carapace and tergites darkened; faint intercarinal fuscosity on posteromedial area of carapace and medial areas of tergites; metasoma V black except for most distal portion; all appendages pale yellow.

Carapace. Subrectangular, W/L 1.03, with steeply sloped lateral flanks; upper surface with nearly flat posterior and medial plateau areas, well raised ocular tubercle; interocular triangle convex laterally, weakly depressed medially; anterior margin distinctly emarginate, microdenticulate except for lateral ends, bordered by row of large granules; 6 short macrosetae on anterior margin, carapace otherwise devoid of macrosetae; 5 lateral eyes (3 large, 2 small) on each side; carination: anterior median, superciliary, central lateral, posterior median and posterior lateral carinae strong, coarsely granular; anterior median carinae separated by clear gap from anterior marginal granules; central lateral and posterior median carinae fused into lyre configuration; central median carinae coarsely granular, anterior and posterior portions nearly collinear, not strongly curved, with only slight inward bend on posterior ends; posterior lateral carinae strong, hind end with slight lateral bend, projecting past posterior margin of carapace; lateral ocular carinae weak with medium granules; granulation: sparse patches of 8–10 medium or large granules on each anterolateral corner of interocular triangle, 8–9 small granules on area in front of lateral ocular carinae; surface between anterior median carinae smooth with 8 scattered small granules; lateral flanks almost smooth, with few sparse granules; other intercarinal surfaces smooth with few fine granules; posterior median furrow shallow, almost flat, with median line of several fine granules or microgranules, flanked by lateral arcs of small granules; posterior margin of carapace between posterior lateral carinae bordered by row of medium granules.

Chelicera. Dorsal surface of manus smooth, with 6 short, pale microsetae, 4 near apical margin, 2 subapical, each surrounded by clusters of granules; dorsointernal carina at base of fixed finger moderately strong, terminated anteriorly by large granules projecting over front of manus; single dark macroseta midway along dorsointernal carina; dorsal surface of movable finger smooth, with 4 pale microsetae; fingers with characteristic buthid dentition ( Vachon, 1963); movable finger dorsal margin with 5 teeth: dorsal distal tine, subdistal, median and 2 basal teeth fused in bicusp; ventral margin with 3 teeth: ventral distal tine, median and basal teeth; fixed finger margin with 4 teeth: distal tine, subdistal, median and basal teeth; ventral aspect of fixed finger with 2 teeth.

Coxosternal area. Coxa I smooth except for coarse granules on distal margin, II smooth with sparse fine granules along posterior margin, III smooth, IV smooth to faintly shagreened; coxal endite II with weak, finely granulated carina; coxae II–III with medium to coarse granulation on anterior carinae, distal margins bearing coarse granules; proximal part of anterior carina of coxa III with fine denticulation; 3 macrosetae along anterior carinae of coxae II–III; anterior carina of coxa IV with regular coarse granulation, becoming fine proximally, with single proximal macroseta; posterior margin of coxa IV with finely granulated carina on proximal half; sternum smooth, with anterior clusters of fine granules, scattered fine granules, one pair of macrosetae, and deep posteromedian furrow; genital opercula subtriangular with convex margins, sparse microgranulation in lateral areas, 5–6 short macrosetae.

Pectines. Basal piece with deep anterior median pit, with patches of fine granules anteromedially, otherwise smooth; 6 short macrosetae along anterior margin, 3 more in center; pectines long, tips extending past proximal 7/8 of trochanter IV; 3 marginal lamellae, small accessory lamella distal to first marginal lamella, 11 middle lamellae, 37–37 teeth; marginal lamellae, middle lamellae and fulcra with dense cover of short reddish macrosetae; fulcra with 3–4 setae; if anterior margins of left and right pectines aligned with posterior edges of coxae IV, only most basal pair of teeth slightly overlap, with wide gap between basal middle lamellae, and teeth contact only anterior half of median carinae of sternite III; pectine teeth of moderate size, mid-pectine sensillar margin L/ pectine L 0.080, mid-pectine sensillar margin L/ metasoma I W 0.198.

Mesosoma. Tergites: pretergites smooth; tergites I– II with 5 granular carinae; median and inner lateral carinae linear with medium granules; outer lateral carinae aligned with posterior lateral carinae of carapace, angled outward, strong with large, granules, hind ends extending past posterior margins of tergites without (on I) or with short (on II) lateral extensions; medial intercarinal surfaces smooth with few medium granules, and linear transverse series of fine granules on anterior margin; lateral flanks steeply sloped with sparse microgranulation; tergites III–VI with 3 straight coarsely granulated carinae; medial intercarinal surfaces mostly smooth with traces of microgranulation; fine granules present in anterior median patch and short transverse strips on either side; lateral surfaces of III–VI moderately sloped, with numerous medium to coarse granules, some arranged in short longitudinal rows; tergite VII with 5 strong, granular carinae; inner and outer lateral carinae joined anteriorly by transverse series of granules becoming more irregular laterally; medial intercarinal surface smooth, sparse anterior median and anterior lateral patches of microgranules; mediolateral surface smooth with 2–3 isolated small granules; lateral surfaces smooth; posterior margins of tergites I–VI rimmed with linear rows of small to medium sized granules; posterior margin of tergite VII with 5 granules between inner lateral carinae, and 0–2 small granules laterally; sternites: sternite III with median carinae strong, thick, densely finely granulated; sternites IV–V with thick, densely granulated carinae, lateral carinae strong, median carinae moderate; sternite VI with median carinae strong, with dense fine granulation, lateral carinae strong with dense, fine granulation in anterior half, coarse serial granulation in posterior half; lateral margins of sternites IV–VII armed with regular medium to large denticulate granules; medial intercarinal surface of III lightly shagreened anteriorly, heavily on posterior margin, of IV densely shagreened, smooth along posterior margin, of V densely shagreened anteriorly, with broad, smooth posterior margin, of VI densely shagreened anteriorly, nearly smooth towards posterior margin; mediolateral surfaces on IV–VI heavily shagreened anteriorly, lightly in posterior area; lateral surfaces of IV–VI smooth; sternite VII with 4 strong, narrow, coarsely granular carinae, intercarinal surfaces smooth except for lightly shagreened area anterior to median carinae; setation: sternite III with 4 macrosetae on median carinae, 4 along external margins; sternites IV–VII with 2 paired macrosetae on median carinae, one in middle of sternite, other on posterior margin; lateral carinae on IV–VI with single posterior marginal macroseta; intercarinal macrosetae: one pair of lateral submarginal setae, one pair of outer mediolateral setae on IV–VI, one pair of lateral setae on VII; intercarinal posterior marginal macrosetae on III–VII: 4, 3, 2, 0, 0.

Hemispermatophore. Trunk elongate, slender; flagellum long, filiform, pars recta 0.4 times length of trunk, pars reflecta 0.6 times length of trunk; inner lobe a broad lamina, gently tapering with blunt apex; median and outer lobes shorter, partially fused, sharply tapering to thin, flexed apical processes; basal lobe a prominent, digitate hook; measurements (topoparatype): trunk L (to base of flagellum) 7.6 mm, pars recta 2.9 mm, pars reflecta 4.5 mm, inner lobe (from base of flagellum) 837 m, median lobe (from juncture with inner lobe) 694 m, outer lobe (from juncture with median lobe) 300 m, basal lobe 216 m.

Metasoma. Very long, slender, total metasoma and telson L/ carapace L 6.1; carination: segment I with 10 complete carinae; segments II–III with 8 complete carinae, median lateral carinae restricted to posterior 1/4 of II, posterior 1/7 of III; metasoma IV with 8 carinae, V with 7 carinae; carinae on segments I–IV finely crenulate-granulate; dorsosubmedian carinae moderately granulate on I, weakly granulate-crenulate on II–III, with irregular fine granulation on posterior 2/3 of IV; dorsolateral carinae moderately serrate or crenulate on I–II, weakly granulate-crenulate on III–IV, weak and finely granular on V; ventrolateral carinae moderate, finely granulate-crenulate on I–IV; median lateral carinae moderate, granulate on I–III; ventromedian carinae moderate to strong on I–IV, crenulations on II–III uniform, not enlarged posteriorly; 42–43 granules on ventromedian carinae of metasoma II–III; metasoma V with dorsolateral carinae weak, finely granular; ventrolateral carinae strong, with small and large triangular dentate granules, larger posteriorly, forming scalloped margin; ventrosubmedian carinae on V weak, confined to anterior 2/3 of segment, marked by small and large dentate granules; ventromedian carina strong, with small to large dentate granules, increasing in size posteriorly; lateral anal margin with 3 large rounded lobes; ventral anal margin with 11 irregular, transverse crenulations; intercarinal surfaces: smooth on segments I–III, smooth to minutely shagreened on IV, sparsely, finely shagreened on V; setation: ventromedian and ventrolateral carinae on segments I–IV each bearing 2 exterior macrosetae; metasoma V with 4 macrosetae on lateral surface, 2 on ventrosubmedian carinae.

Telson. Vesicle smooth, bulbous; ventral surface with scattered fine microsetae and several short macrosetae; aculeus slightly shorter than vesicle.

Pedipalp. Femur: slender, L/ W 4.3; dorsoexternal, dorsointernal and ventrointernal carinae strong with regular large conical granules; internal carina strong, with irregular large dentate granules; external carina obsolete, a smooth ridge with isolated large dentate granules; dorsal and internal surfaces finely, sparsely shagreened, ventral and external surfaces nearly smooth; linear group of 15–17 short macrosetae on distal 1/3 of segment along lower external surface, nearly level with or distal to trichobothrium e 2, part of total of 20–21 distal external accessory macrosetae; patella: slender, L/ W 4.48; dorsointernal carina moderate, with medium to fine granules; dorsomedian carina weak, finely granular; dorsoexternal and external carinae weak, smooth; ventroexternal and ventromedian carinae weak, with fine granules; ventrointernal carina obsolete, marked by noncontiguous coarse conical granules; internal carina moderate, with widely spaced large conical granules separated by fine granulation; both internal and ventrointernal carinae with proximal patellar spur; all patellar surfaces smooth; chela: slender, L/ W 8.24, with tenuous upwardly curved fingers, movable finger L/ manus ventral L 2.34; dorsal marginal and ventroexternal carinae faint, smooth, other carinae obsolete; all chela surfaces smooth; manus and fixed finger with sparse short macrosetae; movable finger with numerous short macrosetae on ventral aspect, culminating in dense subapical brush; 12 primary denticle subrows on fixed and movable fingers (left fixed finger regenerated, with 8 subrows); all denticle subrows flanked by internal and external accessory denticles, except proximal subrow of fixed finger without internal accessory denticle; Trichobothriotaxy: orthobothriotaxic, type A ( Vachon, 1974), db on fixed finger distal to est (db missing on right fixed finger, et missing on left fixed finger, both present in paratypes).

Legs. Long, slender, femur III L/ carapace L 1.27, patella III L/D 4.54; inferior carinae strongly denticulate on femur I–IV and patella I–III, very weakly denticulate, almost smooth on patella IV; tibia III–IV with long spurs; retrolateral pedal spurs simple, non-setose; prolateral pedal spurs basally bifurcate, bearing 2–3 macrosetae; basitarsi I–III with weakly developed bristlecombs, basitarsus III setal counts (left/right): retrosuperior 9/8, retroinferior (including basal accessory seta) 12/13, inferior 14/12; ventral surface of telotarsi with sparse paired rows of stout, finely tapered macrosetae. Right leg I of holotype malformed, vestigial.

Measurements of holotype male (mm). Total L 83.00; metasoma + telson L 54.00; carapace L 8.79, W 9.34, carapace preocular L 4.00; metasomal segments (L/ D /W) I 7.00/ 4.30/ 4.96, II 8.36/ 3.93/ 4.37, III 8.87 / 3.67 / 3.86, IV 9.95 / 3.31 / 3.38, V 10.57 / 3.08 / 3.27; telson L 8.75; vesicle L 4.94, D 3.27, W 3.48; pedipalp chela L 18.21, manus ventral L 5.69, manus W 2.21, manus D 2.57, fixed finger L 11.36, movable finger L 13.29; pedipalp femur L 9.98, W 2.32, patella L 11.12, W 2.48; pectine L 12.17, mid-pectine sensillar margin L 0.98; leg III femur L 11.18; leg III patella L 9.37, D 2.06.

Paratype female (Jabal Samhan). Smaller than holotype male, presumably sub-adult; differs as follows: metasoma, pedipalps and legs not conspicuously elongated: metasoma II L/ W 1.74, metasoma III L/ W 1.92, metasoma IV L/ W 2.30, metasoma V L/ W 2.47, pedipalp femur L/ W 3.25, pedipalp patella L/ W 3.09, pedipalp chela L/ carapace L 1.72, pedipalp chela L/ manus W 6.12, leg III patella L/D 3.23; wider mesosoma; pectines shorter, tips extending to base of trochanter IV, teeth smaller with shorter sensillar margin, 33–34 in number; basal pectine teeth not overlapped when anterior pectinal margins aligned with posterior margins of coxae IV; median carinae obsolete on sternites III–V, weak on VI; lateral carinae moderate, smooth on sternites IV–V, moderate, finely granular on VI; sternite VII with finer median and lateral carinae; sternites with smooth intercarinal surfaces; coarser granules and crenulation on ventromedian and ventrolateral carinae on metasoma II–III, with more enlarged denticles on posterior half of segments; metasoma II–III with 20–25 granules on ventromedian carinae; metasoma V with larger denticles on ventrolateral carinae, posterior denticles enlarged, triangular; pedipalp femur with 27 distal external accessory macrosetae.

Measurements of sub-adult female (mm). Total L 73.00; metasoma + telson L 48.00; carapace L 8.43, W 9.55, carapace preocular L 4.10; metasomal segments (L/ D/ W) I 5.76/ 4.11/ 4.77, II 7.01/ 3.87/ 4.03, III 7.33/ 3.86/ 3.82, IV 8.15/ 3.54/ 3.54, V 9.28/ 3.12/ 3.76; telson L 8.70; vesicle L 4.38, D 3.12, W 3.30; pedipalp chela L 14.50, manus ventral L 4.31, manus W 2.37, manus D 2.72, fixed finger L 9.08, movable finger L 10.40; pedipalp femur L 7.96, W 2.45, patella L 9.04, W 2.93; pectine L 8.86; mid-pectine sensillar margin L 0.54; leg III femur L 8.88; leg III patella L 7.11, D 2.20. Meristics: pedipalp movable fingers with 12/ 12 (left/ right) subrows of primary denticles, fixed fingers with 11/ 8 subrows (anomalous fusion of rows on right finger); basitarsus III seta counts (left/ right): retrosuperior 15/11, retroinferior 14/13, inferior 13/14.

Variation. Comparison of the holotype to two smaller paratype males showed an increasing slenderness of pedipalps, legs and metasoma with increasing size ( Tab. 2 ). The smaller of the two paratype males is more similar to L. haenggii sp. n. in morphometrics and the presence of 4 enlarged denticles on ventrolateral carinae of metasoma V. Median carinae on sternites III– V are weak or obsolete on the smaller paratype male. The larger paratype male, like the holotype, has weakly developed bristle combs on basitarsi I–III, with seta counts (left/right): retrosuperior 9/10, retroinferior 12/12, inferior 13/13.

DISTRIBUTION. Known only from the Jabal Samhan mountains of Dhofar Province, Oman.

ECOLOGY. The male types were collected by ultraviolet detection among rocks and boulders along the edge of a lightly vegetated wadi on the northern side of the Jabal Samhan mountains. All were very agile and aggressive when captured. The only known female paratype was collected near the base of the Jabal Samhan escarpment facing the southern coastal plain. Both sites are at lower elevations (ca. 400 m a.s.l.), and it is not known if this species occurs at higher elevations in these mountains whose peaks reach over 2,000 m a.s.l. The compressed body, elongated legs, pedipalps and metasoma, and weakly developed basitarsal bristle combs are consistent with a lithophilous ecomorphotype.

Figures

Figure 47: Leiurus heberti sp. n., holotype male. Habitus, dorsal aspect. Wadi Andur, Oman.

Figure 48: Leiurus heberti sp. n., holotype male. Habitus, ventral aspect. Wadi Andur, Oman.

Figure 49: Leiurus heberti sp. n., holotype male. A. Carapace and tergites. B. Coxosternal area and sternites. Wadi Andur, Oman.

Figure 50: Leiurus heberti sp. n., holotype male. Right pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, ventral aspect. E. Chela, dorsal aspect. F. Fixed finger dentition. G. Movable finger dentition. H. Chela, external aspect. Wadi Andur, Oman.

Figure 51: Leiurus heberti sp. n., paratype female. Habitus, dorsal aspect. Jabal Samhan, Oman.

Figure 52: Leiurus heberti sp. n., paratype female. Habitus, ventral aspect. Jabal Samhan, Oman.

Figure 53: Leiurus heberti sp. n., paratype female. A. Carapace and tergites. B. Coxosternal area and sternites. Jabal Samhan, Oman.

Figure 54: Leiurus heberti sp. n., paratype female. Right pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, ventral aspect. E. Chela, dorsal aspect. F. Chela, external aspect. Jabal Samhan, Oman.

Figure 55: Leiurus heberti sp. n. Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Male holotype, Wadi Andur, Oman. Female paratype, Jabal Samhan, Oman.

Figure 56: Leiurus heberti sp. n., holotype male. Tarsi. A-D. Left telotarsus and distal basitarsus, ventral aspect. A. Leg I. B. Leg II. C. Leg III. D. Leg IV. E. Right basitarsus III retrolateral aspect. Upper scale bar: A–D, lower scale bar: E. Wadi Andur, Oman.

Figure 57: Leiurus heberti sp. n., holotype male. Trichobothrial map of pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, external aspect. E. Chela, ventral aspect. Wadi Andur, Oman.

Figure 58: Hemispermatophores of new Leiurus species. A. L. macroctenus sp. n., paratype male, Duhai, Oman. B. L. heberti sp. n., holotype male, Wadi Andur, Oman. C. L. haenggii sp. n., paratype male, Jabal Qara, Oman. Distal portions hemispermatophores illustrated to show flagellum and lobes (right hemispermatophores, convex aspect). Scale bars: 1 mm.

Figure 59: Chelicerae of Leiurus species. A, B. L. macroctenus sp. n., paratype male, Thumrait, Oman. Right chelicera, ventral (A) and dorsal (B) aspect. C, D. L. haenggii sp. n., paratype male, Jabal Qara, Oman. Right chelicera, ventral (C) and dorsal (D) aspect. E, F. L. brachycentrus (Ehrenberg, 1829), female, Al Mansuriah, Yemen. Right chelicera, ventral (E) and dorsal (F) aspect. G, H. L. heberti sp. n., holotype male, Wadi Andur, Oman. Right chelicera, ventral (G) and dorsal (H) aspect. Scale bar in A: 1 mm (also applies to B–H).

Figure 87: Metasoma V and telson of Leiurus spp. Males, lateral aspect. A. L. abdullahbayrami Yağmur, Koç et Kunt, 2009. B. L. arabicus sp. n.. C. L. brachycentrus (Ehrenberg, 1829) stat. n.. D. L. haenggii sp. n. E. L. heberti sp. n. F. L. hebraeus (Birula, 1908) stat. n.. G. L. macroctenus sp. n.. H. L. quinquestriatus (Ehrenberg, 1828). Locality data as in Figs. 9, 20, 31, 43, 55, 67, 79. Scale bars: 2 mm.

Figure 88: Metasoma V and telson of Leiurus spp. Males, ventral aspect. A. L. abdullahbayrami Yağmur, Koç et Kunt, 2009. B. L. arabicus sp. n.. C. L. brachycentrus (Ehrenberg, 1829) stat. n.. D. L. haenggii sp. n.. E. L. heberti sp. n. F. L. hebraeus (Birula, 1908) stat. n.. G. L. macroctenus sp. n.. H. L. quinquestriatus (Ehrenberg, 1828). Locality data as in Figs. 9, 20, 31, 43, 55, 67, 79. Scale bars: 2 mm.

Figure 89: Metasoma V and telson of Leiurus spp. Females, lateral aspect. A. L. abdullahbayrami Yağmur, Koç et Kunt, 2009. B. L. arabicus sp. n. C. L. brachycentrus (Ehrenberg, 1829) stat. n. D. L. haenggii sp. n. E. L. heberti sp. n. F. L. hebraeus (Birula, 1908) stat. n. G. L. macroctenus sp. n. H. L. quinquestriatus (Ehrenberg, 1828). Locality data as in Figs. 9, 20, 31, 43, 55, 67, 79. Scale bars: 2 mm.

Figure 90: Metasoma V and telson of Leiurus spp. Females, ventral aspect. A. L. abdullahbayrami Yağmur, Koç et Kunt, 2009. B. L. arabicus sp. n. C. L. brachycentrus (Ehrenberg, 1829) stat. n.. D. L. haenggii sp. n. E. L. heberti sp. n. F. L. hebraeus (Birula, 1908) stat. n. G. L. macroctenus sp. n. H. L. quinquestriatus (Ehrenberg, 1828). Locality data as in Figs. 9, 20, 31, 43, 55, 67, 79. Scale bars: 2 mm.

Figure 91: Pigmentation patterns of metasoma IV, V of Leiurus species, ventral aspect. A. L. haenggii sp. n., holotype female, Ta’if, Saudi Arabia (NHMB 17k). B. L. macroctenus sp. n., holotype male, Thumrait, Oman. C. L. heberti sp. n., holotype male, Wadi Andur, Oman. D. L. quinquestriatus (Ehrenberg, 1828), female, Cairo, Egypt (NHMB 17d). E. L. brachycentrus (Ehrenberg, 1829) stat. n., male, Ad Darb, Saudi Arabia (NHMB 17ag). F. L. arabicus sp. n., holotype female, Kushm Dibi, Saudi Arabia (NHMB 17aq). G. L. hebraeus (Birula, 1908) stat. n., female, Israel (NHMB 17a). H. L. abdullahbayrami Yağmur, Koç et Kunt, 2009, male, Eski Sarkaya Village, Turkey. Scale bars: 2 mm.

Figure 92: Sternite III of Leiurus spp., female. A. L. abdullahbayrami Yağmur, Koç et Kunt, 2009, Caybasi Village, Turkey. B. L. arabicus sp. n., holotype, Kushm Dibi, Saudi Arabia (NHMB 17aq). C. L. arabicus sp. n., paratype, Al Amar, Bahrain (NHMB 17bg). D. L. heberti sp. n., paratype, Jabal Samhan. Oman. E. L. haenggii sp. n., paratype, Wadi Turabah, Saudi Arabia (NHMB 17al). F. L. haenggii sp. n., paratype, Adnan, Saudi Arabia (NHMB 17am). G. L. brachycentrus (Ehrenberg, 1829) stat. n., Al Mansuriah, Yemen. H. L. macroctenus sp. n., paratype, Masirah Island, Oman. I. L. macroctenus sp. n., paratype, Thumrait, Oman. J. L. quinquestriatus (Ehrenberg, 1828), Kharga Oasis, Egypt (NHMB 17i). K. L. hebraeus (Birula, 1908) stat. n., Israel (NHMB 17a). L. L. jordanensis Lourenço, Modry et Amr, 2002, al-Tawil, Saudi Arabia.

Figure 93: Medial intercarinal areas of tergite III of female Leiurus spp. A. L. arabicus sp. n., paratype, Jeddah, Saudi Arabia. B. L. arabicus sp. n., paratype, Wadi Khumrah, Saudi Arabia (NHMB 17bj). C. L. haenggii sp. n., paratype, Yemen. D. L. haenggii sp. n., paratype, Wadi Maraba, Saudi Arabia (NHMB 17al). E. L. heberti sp. n., paratype, Jabal Samhan. Oman. F. L. abdullahbayrami Yağmur, Koç et Kunt, 2009, Caybasi Village, Turkey. G. L. brachycentrus (Ehrenberg, 1829) stat. n., Al Mansuriah, Yemen. H. L. macroctenus sp. n., paratype, Masirah Island, Oman. I. L. hebraeus (Birula, 1908) stat. n., Israel (NHMB 17a). J. L. hebraeus Birula, 1908, Kurayyima, Jordan. K. L. quinquestriatus (Ehrenberg, 1828), Kharga Oasis, Egypt (NHMB 17i). L. L. jordanensis Lourenço, Modry et Amr, 2002, al-Tawil, Saudi Arabia.

Figure 95: Variation in key diagnostic biometrics of adult Leiurus spp. A. Cumulative distributions of position of trichobothrium db relative to est on pedipalp fixed finger for: L. abdullahbayrami (53), L. brachycentrus stat. n. (11), L. hebraeus stat. n. (38), L. macroctenus sp. n. (176), L. quinquestriatus (61), L. haenggii sp. n. (158), L. arabicus sp. n. (87) (number of fingers measured in parentheses). Data for males and females were pooled. The db–est distances were normalized to pedipalp movable finger length (chord length from finger tip to external articular condyle), with positive values indicating db distal to est, negative values indicating db proximal to est. Distributions of L. haenggii (µ = 0.06367) and L. arabicus sp. n. (µ = 0.06671) were not significantly different (P = 0.405148, t test), indicating a close relationship between these species. Distribution of L. quinquestriatus (µ = 0.047313) was significantly different from those of L. haenggii and L. arabicus (P = 0.000102 and 0.000030 respectively). B. Scatter plot of morphometric ratios of mid-pectine sensillar margin L (MPSM, indicated in inset) to metasoma I W, and to carapace L, showing differences in relative pectinal tooth size in females of nine species of Leiurus. C. A subset of the female data in Figure 95B, plotted as cumulative distributions of morphometric ratio of mid-pectine sensillar margin L to metasoma I W. Distributions of L. haenggii (µ = 0.102268) and L. arabicus (µ = 0.10432) were not significantly different (P> 0.1, K-S test). D. Scatter plot of morphometric ratios of mid-pectine sensillar margin L to metasoma I W (relative size of pectine teeth), and metasoma II L/W (slenderness of metasoma II) for females of nine species of Leiurus. A, C, D: symbol key as in Figure 95B.

Figure 98: Selected morphometrics of Leiurus, Cicileiurus, Cicileus and Compsobuthus compared to other Buthus group scorpions. A–B. Scatter plots of the fraction of fixed finger length distal to db (A) and est (B) vs. the ratio of movable finger length to carapace length. Each point represents one sex of one species. Larger ordinate values correspond to more basal positions of the trichobothria, and larger abscissa values to longer pedipalp fingers. There was a significant inverse correlation between relative length of the portion of the fixed finger distal to db and est (R = -0.6534, -0.5488, respectively), and the relative length of the movable finger (the latter being a measure of elongation of both fixed and movable fingers). Highlighted symbols show that Leiurus (light magenta circles), Cicileiurus (red triangle), Cicileus (green squares) and Compsobuthus (yellow circles) are located in the lower right halves of the plots, i.e. all have relatively elongated fingers and more distal placement of both db and est. Gray circles are data from other Buthus group species. C. Scatter plot of the fraction of fixed finger length distal to db vs. the fraction distal to est. The strong positive correlation (R = 0.8052) indicates a tendency for db and est to move together towards more distal locations as the fixed finger becomes more elongated. Species above the diagonal (solid blue) have db proximal to est, and those below have db distal to est. Solid gray lines in A–C are fits by least squares regression through all points. D. Scatter plot of pedipalp femur L/W (a measure of pedipalp elongation) vs. carapace L (a measure of body size). These two variables were not significantly correlated (R = 0.094). Data were compiled from the literature and specimens in the authors collections for 38 genera and 203 species representing the majority of taxa in the Buthus group, including both males (N = 124) and females (N = 97). Genera (and number of species) included: Afghanobuthus (1), Androctonus (11), Apistobuthus (2), Baloorthochirus (1), Birulatus (2), Buthacus (13), Butheoloides (11), Butheolus (5), Buthiscus (1), Buthus (26), Cicileiurus (1), Cicileus (2), Compsobuthus (33), Congobuthus (1), Darchenia (1), Gint (1), Hemibuthus (1), Hottentotta (31), Leiurus (10), Liobuthus (1), Lissothus (3), Mesobuthus (7), Neobuthus (2), Odontobuthus (6), Orthochirus (12), Pantobuthus (1), Pectinibuthus (1), Plesiobuthus (1), Polisius (1), Razianus (3), Saharobuthus (1), Somalibuthus (1), Vachoniolus (4), Vachonus (1).

Figure 99: Geographic plot of locality data of material examined for nine species of Leiurus distributed over northeast Africa, the Levant and Arabian Peninsula. Map colored by elevation with shaded relief. Additional locality data for: L. abdullahbayrami from Yağmur et al., 2009; for L. brachycentrus Ehrenberg, 1829 stat. n., from Simon, 1882 (Buthus beccarii).

Figure 100: Geographic plot of locality data of material examined for five species of Leiurus from the Arabian Peninsula. Map colored by terrain with shaded relief, indicating major physiographic regions.