Leiurus arabicus, 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 : 48-60

Leiurus arabicus sp. n.

( Figs. 25–46 , 58C , 87B , 88B , 89B , 90B , 91F , 92B–C , 93A–B , 94D , 95 , 98–100 , Tabs. 3A , 4 ) http://zoobank.org/urn:lsid:zoobank.org:act:1EAD22 25-BC00-46F3-B053- F59637 AAEF44

REFERENCES

Leiurus quinquestriatus: Vachon, 1979a: 49–50 (in part); Kinzelbach, 1985: map II (in part); El- Hennawy, 1992: 101, 125–126 (in part); Kovařík, 1998: 112 (in part); Fet & Lowe, 2000: 155 (in part); Hendrixson, 2006: 84–91 (in part), fig. 18 (in part), figs 20b, 20c, pl. 14; Kaltsas et al., 2008: 218– 219 (in part); El-Hennawy, 2009: 122 (in part); El- Hennawy, 2014: 45 (in part).

Leiurus quinquestriatus hebraeus: Levy & Amitai, 1980: 48–53 (in part).

HOLOTYPE. Adult ♀, Saudi Arabia, Kushm Dibi, 19.X.1979, leg. A. Barkham ( NHMB 17aq).

PARATYPES. Bahrain: 2 ♀, Al Amar, 25°59'N 50° 32'E, 20 m a.s.l., XI.1985, leg. D. Lickfold ( NHMB 17bg). Saudi Arabia: 1 ♀, 1 juv ♀, Kashm Khafs, 25°36'N 46°27'E, 720 m a.s.l., leg. W. Büttiker ( NHMB 17ac2); 1 juv ♂, Khushm Dibi, 24°17'N 46°09'E, 750 m a.s.l., leg. W. Büttiker ( NHMB 17ac); 1 juv ♂, Wadi Durmah, Stn 11, végétation riche, 24°37'N 46°06'E, 580 m a.s.l., 2.I.1976, leg. W. Büttiker ( NHMB 17m); 1 ♀, Wadi Tumair, Stn 13, oasis, désert de pierres sèches , 25°43'N 45°51'E, 650 m a.s.l., 20.II.1976, leg. W. Büttiker ( NHMB 17o); 2 ♀, 1 juv ♂, 3 juv ♀, Wadi Mizbil, Stn 18, désert de pierres sèches , 24°30'N 46°25'E, 700 m a.s.l., 25.II.1977, leg. W. Büttiker ( NHMB 17l); 1 juv ♀, Wadi Mutaiwijah, Stn 21 , Mecca Road , végétation relativement riche, 24°34'N 46°11'E, 600 m a.s.l., 18.III.1977, leg. W. Büttiker ( NHMB 17p); 1 ♂, Jeddah, 21°31'N 39°13.15'E, 1.IX.1977, leg. W. Büttiker ( NHMB 17q); 1 juv ♀, Wadi Mizbil, 24°29'N 46°25'E, 610 m a.s.l., 24.XII.1977, leg. W. Büttiker ( NHMB 17ao); 1 ♀, Jubail, 27°00.28'N 49°39.68'E, 8 m a.s.l., III.1978, leg. W. Büttiker ( NHMB 17v); 1 juv ♂, Jebel Banban, 25°23'N 46°36'E, 660 m a.s.l., 16.III.1978, leg. W. Büttiker ( NHMB 17n); 1 juv ♂, Khushm Dibi, 24°17'N 46°09'E, 750 m a.s.l., 19.V.1978, leg. W. Büttiker ( NHMB 17t); 1 ♂, Riyadh, 23°20'N 45°20'E, IX.1978, leg. B. Vincett ( NHMB 17ap); 1 ♂, 2 ♀, Wadi Khumrah, Station 12 , 24°55'N 46°11'E, 9.XI.1978, leg. W. Büttiker ( NHMB 17r); 1 ♀, Wadi Khumrah, 24°56'N 46°08'E, 830 m a.s.l., 17.VIII.1979, leg. W. Büttiker ( NHMB 17w); 1 ♀, Riyadh, 23°20'N 45°20'E, Sep 1979, leg. W. Büttiker ( NHMB 17ar); 1 ♂, Kushm al Buwayhiyat , 25°12'N 46°52'E, 720 m a.s.l., 23.IX.1979, leg. A. Barkham ( NHMB 17an); 1 juv ♂, 1 juv ♀, Khushm Dibi, 24°13.61'N 46°06.5'E, 19.X.1979, leg. A. Barkham ( NHMB 17ae); 3 ♂, Khushm Dibi, 24°13.61'N 46°06.5'E, 19.X.1979, leg. A. Barkham ( NHMB 17aq); 2 juv ♀, Jeddah, 21°31'N 39°13.15'E, 1.XI.1979, leg. W. Büttiker ( NHMB 17ak); 1 ♂, Riyadh, 23°20'E 45°20'E, 3.XI.1979, leg. W. Büttiker ( NHMB 17ai); 1 juv ♀, Jebel Buwayb, 25°21'N 46°45'E, 700 m a.s.l., 29.XI.1979, leg. W. Büttiker ( NHMB 17aa); 1 ♂, 1 ♀, Wadi Tuweig, 24°32.6'N 46°32.7'E, 680 m a.s.l., 7.II.1980, leg. W. Büttiker, NHMB Sc 495); 1 ♀, Wadi Khumrah, 24°56'N 46°08'E, 830 m a.s.l., Jun 1980, leg. W. Büttiker ( NHMB 17bj); 1 ♂, Hudenah, 25°34.09'N 39°21.66'E, 800 m a.s.l., 1981, leg. W. Büttiker ( NHMB 17bk); 3 ♀, 3 juv ♀, Khushm Dibi, 24°17'N 46°09'E, 750 m a.s.l., 23.I.1981, leg. W. Büttiker ( NHMB 17ag2); 1 juv ♂, Wadi Khumrah, 24°56'N 46°08'E, 830 m a.s.l., 4.IV.1981, leg. W. Büttiker ( NHMB 17bl) ; 1 ♂, Kashm al-Atash , 24°07'N 46°19'E, 630 m a.s.l., 10.IX.1981, leg. W. Büttiker ( NHMB 17aa2); 1 ♀, 2 juv ♂, 1 juv ♀, Khushm Dibi, 24°17'N 46°09'E, 30.IX.1981, leg. W. Büttiker ( NHMB 17af2); 1 ♀, 1 juv ♂, 3 juv ♀, Riyadh, 24°18.6'N 46°28.2'E, 5.XI.1981, leg. W. Büttiker ( NHMB 17bm); 1 ♂, Jeddah, Al Odaybi , 21°31.01'N 39°13.15'E, 20 m a.s.l., 13.XI.1983, leg. M.F. Ismar ( NHMB 17ai2); 2 ♀, Jeddah, 21°31'N 39°13.15'E, 4.XII.1984, leg. K. Yip ( NHMB 17bi); 1 ♀, halfway between Ranja and Qalat Bishah , 20°37.81'N 42°39. 57'E, 1080 m a.s.l., 25.II.1985, leg. J. Dobricek ( NHMB 17ao2).

DIAGNOSIS (adults). Medium to large Leiurus , 74–100 mm in length, carapace L 8.9–11.6 mm; base color yellow to orange-yellow, with variable dark pigmentation on carapace and tergites, either uniformly dark on carapace and tergites I–VI, or yellow to brown with dark color around median ocular tubercle and light interocular triangle; metasoma V dark except for posterior end; area of carapace between anterior median carinae smooth or with scattered medium to fine granules, area between posterior median carinae with shallow to moderately deep median furrow flanked by arcs of fine or coarse granules; medial intercarinal surfaces of tergites II–III usually smooth or with sparse granulation; posterior margin of coxa III smooth or with sparse fine granules; metasoma moderately slender, metasoma II L/ W 1.65 – 1.89, metasoma III L/ W 1.86 –2.12, metasoma IV L/ W 2.31 –2.58; ventromedian carinae of metasoma II and III with 17–27 denticles (41/44 carinae); metasoma V with enlarged subtriangular or triangular denticles on ventrolateral carinae; pedipalps slender, patella L/ W 3.35 – 3.91; leg III patella L/D 3.82–4.38; pectine teeth ♂ 34– 40, ♀ 28–33; pectines long, narrow, pectine L/ carapace L ♂ 1.27, ♀ 1.03–1.19, mid-pectine sensillar margin L/metasoma I W ♂ 0.159, ♀ 0.093 –0.120; 1–2 basal pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; pectine basal piece smooth in females, smooth or slightly shagreened in males; leg III basitarsus with 7–19 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db usually distal to est; sternite VII with area between median carinae smooth or very faintly shagreened anteriorly, more heavily in males; sternite carination: males, sternite III with median carinae moderate to strong, sternites IV–V with lateral carinae strong, median carinae weak to moderate; females, sternite III with median carinae weak to moderately strong, sternites IV–V with lateral carinae strong, median carinae weak to moderate.

COMPARISONS. L. arabicus sp. n. differs from L. quinquestriatus , and resembles L. haenggii sp. n. in the characters indicated under comparison with the latter species. It differs from L. haenggii by having more slender pedipalp, leg and metasomal segments, and moderate to strongly granulated median carinae on sternites III–V of females ( Figs. 25B vs. 37B, 92B–C vs. 92E–F). These differences are more apparent when comparing adult females, and tend to be obscured in adult males which have as secondary sexual characteristics, elongation of pedipalp and metasomal segments and stronger carination or granulation of sternites. Morphometric ratios of L. haenggii and L. arabicus are overlapped in younger instars, but diverge with increasing age and separate in adults ( Figs. 46B–D ). Biometric separation of adult females of the two species was obtained from the product of three morphometric ratios quantifying slenderness of pedipalp, leg and metasomal segments: F s = (pedipalp patella L/W) × (leg III patella L/D) × (metasoma III L/W) ( Fig. 46A ). The interocular triangle was always dark or fuscous in L. haenggii , whereas in L. arabicus it was either pale (typical form in central Najd) or dark (Arabian Gulf coast and Jeddah).

ETYMOLOGY. The specific epithet refers to the Arabian Peninsula where this species is endemic.

DESCRIPTION (holotype female).

Coloration. Base color yellow; carapace with central mask of dark reddish-brown pigmentation around median ocular tubercle between central lateral and lateral ocular carinae, pigment extending forward to lateral eyes; interocular triangle yellow, with faint fuscosity along anterior median carinae and anterior carapace margin; posterior median area of carapace with weaker fuscosity; median and anterior areas of tergites with irregular faint fuscosity, pretergites mottled brown; metasoma V dark brown except for posterior end, dorsal surface less intensely pigmented but more fuscous medially; ventral aspect of metasoma IV with pair of dark longitudinal stripes along ventromedian carinae.

Carapace. Subrectangular, W/L 1.14, with steeply sloped lateral flanks; upper surface with nearly flat posterior and medial plateau, strongly raised ocular tubercle; interocular triangle convex laterally, depressed medially; anterior margin straight, medially microgranulated, laterally smooth, bordered by gently curved row of medium sized granules; 8 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 moderate to strong, coarsely granular; anterior median carinae not extending to anterior margin of carapace, separated from anterior marginal row of granules; central lateral and posterior median carinae fused into lyre configuration; central median carinae coarsely granular, anterior part straight, angled outward, posterior part outwardly curved; posterior lateral carinae strong, hind end with little or no lateral extension, projecting past posterior margin of carapace; lateral ocular carinae moderate, with small, spaced granules; granulation: sparse patches of 12–14 large granules on anterolateral corners of interocular triangle, 6–8 small granules in front of lateral ocular carinae; surface between anterior median carinae smooth, with one small granule and few microgranules; other intercarinal surfaces smooth except for few isolated small to medium granules; posterior median furrow shallow with broken median line of fine granules, flanked by lateral arcs of small to medium granules; posterior margin of carapace between posterior lateral carinae rimmed by irregular row of small to medium granules.

Chelicera. Dorsal surface of manus smooth, with 6 short, pale microsetae, 4 near apical margin, 2 subapical, each surrounded by granules; dorsointernal carina at base of fixed finger very strong, well granulated, terminating anteriorly with prominent granules projecting over front of manus; single macroseta in middle of 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. Coxae smooth; coxal endite II with weak, finely granulated carina; coxae II–III with coarsely granular anterior carinae, distal margins bearing medium to coarse granules; proximal 1/3 of anterior carina of coxa III almost smooth; 3 long macrosetae along anterior carina of coxa II, 4 macrosetae (3 long, 1 short) along anterior carina of coxa III; anterior carina of coxa IV with regular small to medium granulation, with single proximal macroseta; posterior margin of coxa IV with finely granulated carina on proximal half; sternum smooth, subtriangular with indented lateral margins, deep posteromedian longitudinal sulcus and pit, two short macrosetae; genital opercula smooth with 5–6 short macrosetae.

Pectines. Basal piece with concave anterior margin divided by small median groove, lacking granules, bearing 6 macrosetae; pectines narrow, tips not extending past distal end of coxa IV; both combs with 3 marginal lamellae (right comb with small accessory lamella distal to first marginal lamella), 7–8 middle lamellae, 31–31 teeth; marginal and middle lamellae with sparse to moderate cover of short reddish macrosetae; fulcra with 3–5 setae; pectine teeth relatively small, mid-pectine sensillar margin L/ pectine L 0.056, mid-pectine sensillar margin L/ metasoma I W 0.107.

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, strongly raised with enlarged posterior granules, hind ends projecting past posterior margins of tergites, without lateral extensions; medial intercarinal surfaces smooth, with transverse anterior series of small or medium granules; lateral flanks steeply sloped with sparse granulation; tergites III–VI with 3 straight or slightly curved carinae with medium granules; medial intercarinal surfaces smooth, with short transverse anterior series of granules; very fine granulation on anterior median patch and in transverse strips on either side; lateral surfaces moderately sloped, heavily granulated, with short longitudinal rows of granules; tergite VII with 5 strong, granular carinae; inner and outer lateral carinae joined anteriorly by transverse granule rows; intercarinal surfaces smooth, with few isolated fine granules; fine granulation on anterior median patch and transverse strips on either side; posterior margins of tergites I–VI rimmed with linear rows of medium sized granules; posterior margin of tergite VII smooth except for sparse granules between inner lateral carinae; sternites: sternite III with moderately strong, broad, finely granulated median carinae; sternites IV–V with strong, finely granulated lateral carinae, weak, lightly shagreened median carinae; sternite VI with strong, coarsely granulated lateral carinae, moderate, finely granulated median carinae; sternite VII with strong, coarsely granulated median and lateral carinae; lateral margins of sternites IV–VII armed with regular denticulate granules; medial intercarinal surfaces of all sternites smooth or very faintly shagreened anteriorly, lateral intercarinal surfaces smooth posteriorly, lightly shagreened anteriorly on IV– VI; setation: sternite III with 4–5 macrosetae on median carinae, 5–6 on areas external to median carinae; sternites IV–VII with 2 paired macrosetae on median carinae, one pair in middle of sternite, other on posterior margin; lateral carinae on IV–VI with posterior marginal macrosetae, mid-carinal seta present or absent; intercarinal macrosetae: one anterior pair of lateral marginal setae on sternites IV–VI; two outer pairs of mediolateral setae on IV–V, one pair on VI; one pair of lateromarginal setae on VII; intercarinal posterior marginal macrosetae on III–VII: 9, 6, 4, 1, 0.

Metasoma. Long, slender; total metasoma and telson L/ carapace L 5.9; carination: segment I with 10 complete carinae; segments II–III with 8 complete carinae, median lateral carinae restricted to posterior 0.28 of II, posterior 0.23 of III; metasoma IV with 8 carinae, V with 7 carinae; carinae on segments I–IV with crenulate granulation; dorsosubmedian carinae moderate on I–II, weak on III–IV; dorsolateral and ventrolateral carinae moderate on I–IV; median lateral carinae moderate on I–II, moderate posteriorly, weak anteriorly on III; ventromedian carinae moderate on I and IV, moderate anteriorly, strong posteriorly on II–III, with posterior granules taller but shorter; 20–21 granules on ventromedian carinae of metasoma II–III; metasoma V with dorsolateral carinae very weak, faintly granulated, ventrolateral carinae strong with dentate granules increasing in size posteriorly, with several large triangular denticles, ventrosubmedian carinae marked by series of small to medium dentate granules on anterior 2/3 of segment, ventromedian carina strong, armed with small to large dentate granules; lateral anal margin with 3 lobes, ventral anal margin with 10 irregular, narrow to wide transverse crenulations; intercarinal surfaces: segments I–IV smooth, segment V smooth dorsally, finely shagreened laterally, smooth to slightly shagreened ventrally; setation: segments I–IV: ventromedian carinae with 3 macrosetae (one posterior marginal), ventrolateral carinae with 2 macrosetae slightly external to carina; metasoma V with 5 macrosetae on lateral surface (2 lateral anal), 4 pairs on ventral surface.

Telson. Vesicle smooth, bulbous; ventral surface bearing scattered fine microsetae and several short macrosetae with associated shallow indentations; aculeus approximately same length as than vesicle.

Pedipalp. Femur: slender, L/ W 3.82; dorsoexternal, dorsointernal and ventrointernal carinae strong with coarse, closely spaced dentate granules; internal carina strong, with large dentate granules spaced well apart; external carina weak to obsolete, a smooth ridge with isolated dentate granules distally; all intercarinal surfaces smooth; linear cluster of 18–21 accessory macrosetae on lower distal external surface; patella: slender, L/ W 3.60; dorsointernal carina moderate with coarse granulation; dorsomedian carina weak with fine granulation; dorsoexternal, external and ventroexternal carinae weak, smooth; ventromedian carina weak, with fine granules; ventrointernal carina weak, with well spaced medium to small granules and ventral patellar spur; internal carina moderate, with closely spaced small granules and dorsal patellar spur; all intercarinal surfaces smooth; chela: slender, L/ W 6.93, movable finger L/ manus ventral L 2.3; dorsal marginal and dorsal secondary carinae weak, smooth; digital carina weak, smooth, obsolete on manus; external secondary and ventroexternal carinae weak, smooth; other carinae obsolete; all intercarinal surfaces smooth; manus and fixed finger with sparse short macrosetae; movable finger with numerous short macrosetae on ventral aspect, culminating in dense subapical brush; 13 primary denticle subrows on movable fingers, 11–13 on fixed fingers; all subrows except proximal flanked by internal and external accessory denticles. Trichobothriotaxy: orthobothriotaxic, type A ( Vachon, 1974), db on fixed finger distal to est.

Legs. Moderately long, slender, femur III L/ carapace L 1.14, patella III L/D 4.12; 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 tarsal spurs simple, non-setose; prolateral tarsal spurs basally bifurcate, bearing 1–3 macrosetae; basitarsi I–III with well developed bristle-combs, at least as wide as basitarsal segment; basitarsus III setal counts (left/ right): retrosuperior 15/17, retroinferior 13/13 (including basal accessory seta), inferior 15/13; ventral surface of telotarsi with robust, short tapered macrosetae.

Measurements of holotype female (mm). Total L 89.00; metasoma + telson L 58.00; carapace L 9.85, W 11.18, carapace preocular L 4.97; metasomal segments (L/ W /D) I 7.35/ 5.59/ 4.95, II 8.82/ 4.72/ 4.33, III 9.29/ 4.59/ 4.23, IV 10.30/ 4.05/ 3.90, V 11.51/ 4.13/ 3.84; telson L 9.73; vesicle L 5.15, W 3.80, D 3.53; pedipalp chela L 20.24, manus ventral L 6.22, manus W 2.92, manus D 3.47, fixed finger L 12.36, movable finger L 14.29; pedipalp femur L 10.70, W 2.80, patella L 11.71, W 3.25; pectine L 10.68, mid-pectine sensillar margin L 0.60; leg III femur L 11.19; leg III patella L 9.45, D 2.29.

Paratype male (Riyadh). Differs from holotype female as follows: less intense fuscosity on carapace and metasoma V, tergites yellow, metasoma IV without dark markings on ventromedian carinae; body narrower, carapace W/L 1.04; stronger carinae on carapace, tergites and coxae; medial intercarinal surfaces of tergites very faintly shagreened; pectine basal piece narrower, anterior margin more strongly incised, with deep median pit; pectines wider, longer, tips extending to basal 1/3 of trochanter IV, teeth larger with longer sensillar margins, mid-pectine sensillar margin L/ pectine L 0.078, midpectine sensillar margin L/ metasoma I W 0.159; 37–38 pectine teeth; 1–2 basal pectine teeth overlap if anterior pectine margins aligned to posterior margins of coxae IV; sternite III with stronger, thicker, granulated median carinae; sternites IV–V with stronger, thicker, more heavily granulated lateral and median carinae; sternites VI–VII with all carinae stronger, more coarsely granulated; intercarinal surfaces of sternites III–VI roughened, micro-shagreened, anterior lateral areas more heavily, densely shagreened; ventromedian carinae on metasoma II–III more weakly crenulate, with smaller posterior denticles; metasoma V with dorsal surface slightly roughened, ventral surface more densely shagreened; pedipalp patella more slender, L/ W 3.82, with larger denticles and spurs on ventrointernal and internal carinae; pedipalp chela more slender, L/ W 7.18, with slightly stronger carination, dorsal marginal and dorsal secondary carinae weakly granulated on manus.

Measurements of paratype male (NHMB 17ap) (mm). Total L 84.00; metasoma + telson L 56.00; carapace L 9.23, W 9.58, carapace preocular L 4.34; metasomal segments (L/ W/ D) I 7.37/ 5.76/ 4.92, II 8.90/ 5.00/ 4.52, III 9.20/ 4.72/ 4.29, IV 10.30/ 4.30/ 3.99, V 10.70/ 4.10/ 3.76; telson L 9.47; vesicle L 4.87, W 3.80, D 3.43; pedipalp chela L 18.75, manus ventral L 6.27, manus W 2.61, manus D 2.98, fixed finger L 11.14, movable finger L 13.03; pedipalp femur L 9.89, W 2.70, patella L 11.19, W 2.93; pectine L 11.73; mid-pectine sensillar margin L 0.918; leg III femur L 11.05; leg III patella L 9.06, D 2.07.

Variation (females). Color: the holotype represents a light color phase from the central plateau region of Saudi Arabia. A dark color phase is represented by material from the Arabian Gulf coast and from Jeddah on the Red Sea Coast: carapace and tergites I–VI dark except for narrow band along lateral margins, tergite VII dark on anterior median area, dark stripes on ventromedian and ventrolateral carinae of metasoma II–IV, diffuse intercarinal fuscosity may be present on IV. Morphosculpture: area between anterior median carinae of carapace mostly smooth but sometimes with scattered fine granules; proximal 1/3 of anterior carina of coxa II smooth to finely granulated (weaker granulation than distal 2/3); basal piece of pectines smooth to lightly shagreened; anterior mediolateral intercarinal surfaces of sternites weakly to moderately shagreened; intercarinal surfaces of pedipalp femur and patella smooth to lightly, finely shagreened; inferior carina of leg IV patella smooth to weakly denticulate; morphometrics and meristics: see Table 4 .

DISTRIBUTION. The studied material is mostly from the Najd plateau region of central Saudi Arabia and wadis around the Tuwayq escarpment (light color phase), with a few samples from the east coast (Al Amar and Jubail on the Arabian Gulf) and from Jeddah on the Red Sea coast (both dark color phases). Jeddah lies at the northern end of the Tihamah plain, a distinctive ecological region separated from the Najd by escarpments of the Hijaz and Asir mountain ranges. It is possible that the seemingly disjunct record in Jeddah is due to artificial dispersal from dark phase populations in the east via the heavily traveled Mecca Road. The elevation range of records is 20 – 800 m, with the lowest on the east coast, and the highest on the Najd plateau.

ECOLOGY. Most collections were made from vegetated wadis and oases in arid, stony desert in the region around Riyadh. The species is probably arenicolous, inhabiting burrows in sandy desert soils.

REMARKS. L. arabicus and L. haenggii appear to be closely related parapatric forms distributed in adjacent ecological regions and habitats of the Arabian Peninsula, i.e. rocky mountains along the Red Sea coast and Hadramaut vs. alluvial desert plains of the central Najd plateau and eastern plains extending to the Gulf coast, respectively. Our samples permitted a differential diagnosis based on a combination of pedipalp, leg and metasomal morphometrics, and sternite carination in adult females. However, there remains the possibility that additional sampling could reveal hybridization in transition zones between the two eco-regions. This can be tested in the future by analyzing additional collections from a wider area.

Figures

Figure 25: Leiurus haenggii sp. n., holotype female. A. Carapace and tergites. B. Coxosternal area and sternites. Ta’if, Saudi Arabia (NHMB 17k).

Figure 26: Leiurus haenggii sp. n., holotype female. Pedipalp. A. Right femur, dorsal aspect. B. Right patella, dorsal aspect. C. Right patella, external aspect. D. Left chela, dorsal aspect. E. Left chela, ventral aspect. F. Left movable finger dentition. G. Left fixed finger dentition. H. Left chela, external aspect. Ta’if, Saudi Arabia (NHMB 17k).

Figure 27: Leiurus haenggii sp. n., paratype male. Habitus, dorsal aspect. Wadi Asidah, Saudi Arabia (NHMB 17al2).

Figure 28: Leiurus haenggii sp. n., paratype male. Habitus, ventral aspect. Wadi Asidah, Saudi Arabia (NHMB 17al2).

Figure 29: Leiurus haenggii sp. n., paratype male. A. Carapace and tergites. B. Coxosternal area and sternites. Wadi Asidah, Saudi Arabia (NHMB 17al2).

Figure 30: Leiurus haenggii sp. n., paratype male. 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. Wadi Asidah, Saudi Arabia (NHMB 17al2).

Figure 31: Leiurus haenggii sp. n. Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Male paratype, Wadi Asidah, Saudi Arabia (NHMB 17al2). Female holotype, Ta’if, Saudi Arabia (NHMB 17k).

Figure 32: Leiurus haenggii sp. n., holotype female. 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. Ta’if, Saudi Arabia (NHMB 17k).

Figure 33: Leiurus haenggii sp. n., paratype 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 Asidah, Saudi Arabia (NHMB 17al2).

Figure 34: Leiurus haenggii sp. n.. A. Paratype female. B. Paratype male. Habitus in vivo, dorsal aspect. Captive bred from Sana'a, Yemen (M. Heule).

Figure 35: Leiurus arabicus sp. n., holotype female. Habitus, dorsal aspect. Kushm Dibi, Saudi Arabia (NHMB 17aq).

Figure 36: Leiurus arabicus sp. n., holotype female. Habitus, ventral aspect. Kushm Dibi, Saudi Arabia (NHMB 17aq).

Figure 37: Leiurus arabicus sp. n., holotype female. A. Carapace and tergites. B. Coxosternal area and sternites. Kushm Dibi, Saudi Arabia (NHMB 17aq).

Figure 38: Leiurus arabicus sp. n., holotype female. 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. Kushm Dibi, Saudi Arabia (NHMB 17aq).

Figure 39: Leiurus arabicus sp. n., paratype male. Habitus, dorsal aspect. Riyadh, Saudi Arabia (NHMB 17ap).

Figure 40: Leiurus arabicus sp. n., paratype male. Habitus, ventral aspect. Riyadh, Saudi Arabia (NHMB 17ap).

Figure 41: Leiurus arabicus sp. n., paratype male. A. Carapace and tergites. B. Coxosternal area and sternites. Riyadh, Saudi Arabia (NHMB 17ap).

Figure 42: Leiurus arabicus sp. n., paratype male. Left pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, dorsal aspect. E. Chela, ventral aspect. F. Movable finger dentition. G. Fixed finger dentition. H. Chela, external aspect. Riyadh, Saudi Arabia (NHMB 17ap).

Figure 43: Leiurus arabicus sp. n. Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Male paratype, Riyadh, Saudi Arabia (NHMB 17ap). Female holotype, Kushm Dibi, Saudi Arabia (NHMB 17ap).

Figure 44: Leiurus arabicus sp. n., holotype female. 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. Kushm Dibi, Saudi Arabia (NHMB 17aq).

Figure 45: Leiurus arabicus sp. n., holotype female. Trichobothrial map of pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, external aspect. E. Chela, ventral aspect. Kushm Dibi, Saudi Arabia (NHMB 17aq).

Figure 46: Morphometric differentiation between female Leiurus arabicus sp. n. and L. haenggii sp. n. A. Scatter plot of the compound morphometric ratio (slenderness factor) Fs = a.b.c (where a = metasoma III L/W, b = pedipalp patella L/W, c = leg III patella L/D) vs. Carapace L (N = 30 L. arabicus, N = 43 L. haenggii). The distribution of ratios of the two species were overlapped for juveniles and young, and were increasingly separated in subadults and adults, as segments of L. arabicus become more slender by allometric growth. B, C, D. Log-log scatter plots of metasoma III L vs. W, pedipalp patella L vs. W, and leg III patella L vs. D, respectively. Linear regression was applied separately to data sets split into upper and lower size ranges for each species (red lines). Numbers show fitted slope ± standard deviation (SD), with SD ranges in brackets. Slopes were similar between L. arabicus and L. haenggii over the lower size range, and diverged over the upper size range. For all three segments, allometry was significantly diphasic in L. arabicus, and only weakly so, or monophasic, in L. haenggii.

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 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 94: Comparative analysis of granulation on tergite III of female Leiurus. A. UV fluorescence image of tergite III medial intercarinal surface of L. quinquestriatus (Egypt). B. Mask of granulation pattern on medial intercarinal surface from image in Figure 94A. C. Enlarged view of tergite III granulation in rectangular area indicated in Figure 94A. Granules were identified as local maxima of fluorescence intensity with contour shadows cast by directional UV illumination, and were modeled by elliptical regions-of-interest (ROIs). Parameters of granule ROIs were measured in ImageJ 1.44 (Rasband, 1997–2011). Area of granulometric analysis was restricted to surfaces bounded by the lateral and posterior marginal carinae, and a line passing through oblique, anterior transverse rows of enlarged granules (granules along carinae and bounding lines were omitted). D. Scatter plot of mean granule diameter vs. total granule area for 4 species of Leiurus (L. arabicus sp. n., L. haenggii sp. n., L. hebraeus stat. n. and L. quinquestriatus). Each point represents granulometric data derived from tergite III of one scorpion specimen (bilaterally, as shown in Figure 94B). Data were extracted from 18,423 ROIs from 36 scorpions. For each specimen, total granule area (a measure of density of granulation) was computed as sum of areas of all ROIs, and mean ROI diameter (a measure of coarseness of granulation) as the mean value of the maximum diameters of all granule ROIs. For comparative analysis, images from different size scorpions were resampled to equalize the distance between left and right posterior marginal granules of the lateral carinae (arbitrarily set to 4,000 units or pixels; linear dimensions expressed as [pixel], areas as [pixel2]). The total granule area separated the species into 2 groups: i.e. sparsely granulated (L. arabicus, L. haenggii) and densely granulated (L. hebraeus, L. quinquestriatus). In contrast, the distributions of ROI diameter were broadly overlapping. E. To obtain a more sensitive comparison of the coarseness of granulation, normalized cumulative distributions of single ROI areas for 3 species of Leiurus were computed (inset indicates number of scorpions analyzed). Relative horizontal positions of these curves indicated increasing coarseness of granulation, in rank order: L. haenggii <L. quinquestriatus <L. hebraeus (Ngranules = 2103, 5202, 2190 respectively). This ranking was confirmed by a Kolmogorov-Smirnov test which detected significant differences between the distributions of log (granule area) (P <0.001).

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.