Leiurus brachycentrus (Ehrenberg, 1829)

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 : 5-18

Leiurus brachycentrus (Ehrenberg, 1829) stat. n.

( Figs. 1– 11 , 59E–F , 87C , 88C , 89C , 90C , 91E , 92G , 93G , 95 , 98–100 , Tab. 3B )

Androctonus (Leiurus) quinquestriatus brachycentrus Ehrenberg, 1829: 353 –354.

SYNONYMS

Buthus beccarii Simon, 1882: 246–248 , pl. VIII, fig. 19.

Leiurus nasheri Kovařík, 2007: 137–141 , tab. 1, figs. 1– 8; syn. n.

REFERENCES

Androctonus (Leiurus) quinquestriatus brachycentrus: Gervais, 1844: 46 ; Braunwalder & Fet, 1998: 33.

Buthus beccarii: Simon 1890: 122 .

Buthus quinquestriatus: Kraepelin, 1891: 58–60 (in part); Thorell, 1893: 359–360 (in part); Kraepelin, 1899: 27–28 (in part); Werner, 1934: 269 (in part);

Buthus quinquestriatus brachycentrus: Birula, 1908: 126 ; Birula, 1910: 118; Borelli, 1915: 461; Birula, 1937: 102–105; Moritz & Fischer, 1980: 311.

Buthus (Buthus) 5- striatus brachycentrus: Birula, 1910: 171 ; Birula, 1917a: 228.

Leiurus quinquestriatus brachycentrus: Vachon 1966: 211 ; Levy, Amitai & Shulov, 1970: 231–233, 240; Pérez, 1974: 24; El-Hennawy, 1992: 126.

Androctonus quinquestriatus brachycentrus: Vachon, 1979a: 50 ; Lourenço, Modrý & Amr, 2002: 637; Lourenço, Qi & Cloudsley-Thompson, 2006: 98.

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

Leiurus quinquestriatus: El-Hennawy, 1992: 101 , 125– 126 (in part); Sissom, 1994: 20–23 (in part); Braunwalder & Fet, 1998: 33; Kovařík, 1998: 112 (in part); Fet & Lowe, 2000: 155 (in part); El- Hennawy, 2009: 122 (in part).

Leiurus nasheri: Yağmur et al., 2009: 1 , 5, 7–8.

TYPE MATERIAL. Holotype ♂, Yemen, “Lohaie” (= Al Luhayyah: 14°42.53'N 43°12'E), 1824–1825, leg. F.W. Hemprich & C.G. Ehrenberg ( ZMHB No. 141) (examined); 4 ♀ (holotype and paratypes of Leiurus nasheri ), 10 km W of Al Mansuriah, Al Hudaydah, 14°43’N 43°12’E, 110 m a.s.l., 8.IV.2007, leg. P. Kabátek & D. Král ( FKCP) (examined).

OTHER MATERIAL EXAMINED. Saudi Arabia: 1 ♂, Ad Darb, 17°43'22''N 42°15'09''E, 70 m a.s.l., 10.IV.1979, leg. W. Büttiker ( NHMB 17 ag). Yemen: 2 ♀ Ta’izz gov., 28.X.2007, N of Al Makha by road, 13°23'37"N 43°16'22"E, 5 m a.s.l. (locality No. 7) , 2 ♀, leg. D. Král ( FKCP) .

ETYMOLOGY. The name refers to the telson which is relatively robust compared to that of L. quinquestriatus , with an aculeus that in males is clearly shorter than the vesicle.

DIAGNOSIS. Small to medium sized Leiurus , 60–72 mm in length, carapace L 6.3–8.0 mm; base color yellow; legs, pedipalps, metasoma and telson with pattern of fuscous markings; metasoma V without dark pigment except on carinae; carapace with anterior median carinae extending to anterior marginal granule row; area of carapace between anterior median carinae lacking dark pigment, with sparse, fine granules, area between posterior median carinae with shallow median furrow, nearly flat, flanked by lateral arcs of fine granules; medial intercarinal surfaces of tergites II–III with dense, coarse to fine granulation; posterior margin of coxa III with row of small granules; metasoma moderately slender, metasoma II L/ W 1.51 –1.66, metasoma III L/ W 1.66 –1.83, metasoma IV L/ W 2.03 –2.19; ventromedian carinae of metasoma II and III with 14–16 denticles; metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps moderately slender, patella L/W ♂ 3.00–3.25, ♀ 2.72; leg III patella L/D 2.72–3.25; pectine teeth ♂ 31–35, ♀ 25–28; pectines medium sized, very broad in males, pectine L/ carapace L ♂ 1.18, ♀ 0.96, mid- pectine sensillar margin L/ metasoma I W ♂ 0.210 –0.234, ♀ 0.115; basal 6 pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; pectine basal piece very lightly, finely shagreened, nearly smooth; leg III basitarsus with 11–15 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db proximal to est; sternite VII with area between median carinae smooth in females, densely, finely shagreened in males; sternite carination: males, sternite III with median carinae strong, sternites IV–V with lateral carinae strong, median carinae weak; females, sternite III with median carinae obsolete, sternites IV–V with lateral carinae very weak, median carinae obsolete.

COMPARISONS. L. brachycentrus stat. n. is differentiated from other Leiurus by a combination of small size, distinctive color pattern, granular medial intercarinal surfaces of tergites II–III, proximal position of chela trichobothrium db relative to est, large male pectine teeth, morphometrics of pedipalp, legs and metasoma, and sternite carination. Among the Arabian species, L. macroctenus sp. n. is most similar in its morphometrics, large male pectine teeth, weak or obsolete carinae on sternites III–V in females, and db basal to est in a minority of specimens. It differs by its larger size, darker fuscous pigment on metasoma V, larger numbers of setae on basitarsal bristle combs, higher pectinal tooth counts, and larger female pectine teeth. L. savanicola from northern Cameroon is another small to medium sized species with maculate pigmentation. It differs in having darker fuscous pigment on metasoma V, distal position of db relative to est, and uniform serrate dentition on ventrolateral carinae of metasoma V.

REDESCRIPTION (adult male, Ad Darb).

Coloration. Base color yellow, with distinctive pattern of fuscous markings; carapace with dark pigment on median ocular tubercle, fuscosity extending forward on lateral interocular triangle external to anterior median carinae, and on central median, central lateral, lateral ocular, posterior median and posterior lateral carinae; medial areas between anterior median, central median and posterior median carinae yellow; pretergites lightly fuscous, with darker medial band flanked by pale spots in some segments; tergites I–VI with darkly marked median and lateral carinae, and fuscous anterolateral areas, posterior median areas yellow; tergite VII with fuscosity underlying median and inner lateral carinae; metasoma I–VI yellow with variable dark pigment on ventromedian and ventrolateral carinae, more intense posteriorly; metasoma V yellow to yellow-brown, with variable dark pigment on ventromedian, ventrosubmedian, ventrolateral and dorsolateral carinae; pedipalp femur and patella with faint to moderate fuscosity on dorsal and internal surfaces, external surface lighter; pedipalp chela with weak or very faint fuscosity on manus; femur of all legs with faint to moderate fuscosity on distal 2/3 of prolateral surface and retrosuperior carina; tibia of all legs with band of faint to moderate fuscosity on proximal 1/2 of inferior prolateral surfaces and superior prolateral carina.

Carapace. Subrectangular, W/L 1.03, with steeply sloped lateral flanks; upper surface with posterior and medial plateau areas, moderately raised ocular tubercle; interocular triangle convex laterally, weakly depressed medially; anterior margin straight, entire width bearing fine granules or denticles, bordered by row of large granules; 10 medium to long 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; granules of anterior median carinae extend forward continuously, contacting anterior marginal row of granules; central lateral and posterior median carinae fused into lyre configuration; central median carinae coarsely granular, anterior portion linear and angled outward, posterior portion outwardly curved; posterior lateral carinae strong, caudal end extended laterally and downward, projecting past posterior margin of carapace; lateral ocular carinae weak or moderate, with smaller granules; granulation: sparse patches of 10–15 large granules on each anterolateral corner of interocular triangle, 10–13 small granules on area on front of lateral ocular carinae; surface between anterior median carinae micro-shagreened with scattered small granules; lateral flanks with moderately dense cover of medium and fine granules; other intercarinal surfaces finely shagreened with scattered small granules; posterior median furrow shallow, almost flat, with median line of fine granules or microgranules, flanked by lateral arcs of small granules; posterior margin of carapace between posterior lateral carinae bordered by row of medium to small granules.

Chelicera. Dorsal surface of manus smooth to slightly shagreened, with 4 short, pale microsetae, 2 near apical margin, 2 subapical, each surrounded by granules; dorsointernal carina at base of fixed finger moderately strong with small granules proximally, terminating distally in 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. Coxa I coarsely granulated, II with sparse medium or fine granules, III lightly shagreened or smooth except for proximal series of medium to small granules along posterior margin, IV lightly shagreened with several isolated medium granules; coxal endite II with strong, coarsely granulated carina; coxae II–III with medium to coarse granulation on anterior carinae, distal margins bearing coarse granules; proximal 1/3 of anterior carina of coxa III with medium tuberculate granulation; 3 macrosetae along anterior carinae of coxae II–III; anterior carina of coxa IV with regular medium to coarse granulation, with single proximal macroseta; posterior margin of coxa IV with finely granulated carina on proximal half; sternum weakly granulated, subtriangular with straight lateral margins, deep posteromedian pit, 2 short macrosetae; genital opercula smooth with 3 short macrosetae, posterolateral margins convex.

Pectines. Basal piece with concave anterior margin and small median groove and pit, slightly roughened, bearing 3 macrosetae; pectines broad, tips extending to proximal 1/3 to 1/2 of coxa IV; combs with 3 marginal lamellae, left comb with small accessory lamella distal to first marginal lamella, 8–9 middle lamellae, 31–33 teeth; basal middle lamella with proximal margin extended, protruding, 1.30–1.35 times longer than distal margin; marginal and middle lamellae with moderate cover of short reddish macrosetae; fulcra with 3–8 setae; pectine teeth relatively large, mid-pectine sensillar margin L/ pectine L 0.112, mid-pectine sensillar margin L/ metasoma I W 0.210.

Mesosoma. Tergites: pretergites smooth; tergites I– II with 5 granular carinae; median and inner lateral carinae linear with medium to coarse granules; outer lateral carinae aligned with posterior lateral carinae of carapace, angled outward, very strong with large, uniform granules, hind ends extending past posterior margins of tergites, without lateral extensions; medial intercarinal surfaces densely granulated, with small to medium granules, and shagreened; lateral flanks moderately sloped, heavily granulated; tergites III–VI with 3 straight coarsely granulated carinae; medial intercarinal surfaces finely granulated or shagreened; short transverse anterior series of granules present on III–IV, obsolete on V–VI; tergites III–VI with fine granulation on anterior median patch and short transverse strips on either side; lateral surfaces of III–VI moderately sloped, heavily granulated, III with short longitudinal rows of granules; tergite VII with 5 strong, granular carinae; inner and outer lateral carinae joined anteriorly by transverse granule rows; medial intercarinal surfaces smooth; patches of fine granulation on anterior ends of median and lateral carinae; posterior margins of tergites I–VI rimmed with linear rows of small to medium sized granules; posterior margin of tergite VII with 3 small granules between inner lateral carinae, and 1–2 small granules laterally; sternites: sternite III with median carinae strong, thick, finely granulated; sternites IV–VI with moderate to strong, finely granulated lateral carinae, weak, finely granulated median carinae; sternite VII with strong, crenulate-granulate median and lateral carinae; lateral margins of sternites IV–VII armed with fine, regular denticulate granules; medial intercarinal surface of III finely shagreened; medial and mediolateral intercarinal surfaces IV–V finely shagreened anteriorly, smooth posteriorly, VI finely shagreened except for posterior margin, medial intercarinal surface of VII finely shagreened anteriorly, smooth posteriorly; lateral intercarinal surfaces of III smooth or very weakly, finely shagreened, IV–VI shagreened anteriorly, smooth posteriorly; mediolateral and lateral surfaces of VII sparsely, finely shagreened; setation: sternite III with 5 macrosetae on median carinae (one shared), single small setae 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 single posterior marginal macroseta; intercarinal macrosetae: one pair of lateral marginal setae, one outer pair of mediolateral setae on IV–VI, one pair of lateral setae on VII; intercarinal posterior marginal macrosetae on III–VII: 4, 3, 3, 0, 0.

Metasoma. Moderately long, slender, total metasoma and telson L/ carapace L 6.29; carination: segment I with 10 complete carinae; segments II–III with 8 complete carinae, median lateral carinae restricted to posterior 0.34–0.50 of II, posterior 0.22 of III; metasoma IV with 8 carinae, V with 7 carinae; carinae on segments I– IV crenulate-granulate; dorsosubmedian carinae moderate on I–III, weak on IV; dorsolateral carinae moderate on I–IV, ventrolateral carinae moderate to strong; median lateral carinae granulate, moderate on I–III; ventromedian carinae moderate on I–IV, crenulations on II–III very weak anteriorly; 14–20 granules on ventromedian carinae of metasoma II–III; metasoma V with dorsolateral carinae very weak, faintly granulated, ventrolateral carinae strong, with blunt subtriangular denticles increasing in size posteriorly, ventrosubmedian carinae marked by series of non-contiguous small to large dentate granules along anterior 5/8 of ventral surface, ventromedian carina strong, with closely spaced medium to large rounded, dentate granules increasing in size posteriorly; lateral anal margin with 3 blunt lobes divided by deep incisions, ventral anal margin with 9 irregular, narrow to wide transverse crenulations; intercarinal surfaces: segments I–IV smooth to lightly shagreened, segment V lightly shagreened dorsally and laterally, more densely 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, rather elongate with steep posterior slope; ventral surface bearing scattered fine microsetae and several short macrosetae with associated shallow indentations; aculeus shorter than vesicle.

Pedipalp. Femur: moderately slender, L/ W 3.54; dorsoexternal, dorsointernal and ventrointernal carinae strong with coarse, closely spaced dentate granules; internal carina strong, with irregular small and large dentate granules; external carina weak to moderate, with well spaced coarse dentate granules; external surface smooth, other intercarinal surfaces lightly shagreened; linear cluster of 14–15 accessory macrosetae on lower distal external surface; patella: moderately slender, L/ W 3.25; dorsointernal carina strong with coarse granlation; dorsomedian carina moderate with fine granulation; dorsoexternal carina very weak, almost smooth; external carina weak, smooth; ventroexternal carina weak, almost smooth; ventromedian carina weak with fine granules; ventrointernal carina strong, with well spaced medium granules and ventral patellar spur; internal carina strong with large dentate granules interspersed with medium granules and dorsal patellar spur; external surfaces smooth, other intercarinal surfaces very lightly shagreened; chela: slender, L/ W 6.09, fingers relatively short, movable finger L/ manus ventral L 1.93; dorsal internal carina weak to obsolete, marked by few isolated granules on manus; dorsal marginal carina weak, with faint granulation at base of manus; external secondary carina weak, smooth; ventroexternal carina moderate, 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; 12 primary denticle subrows on movable fingers, fixed fingers damaged or with anomalous dentition; all subrows except proximal flanked by internal and external accessory denticles. Trichobothriotaxy: orthobothriotaxic, type A  ( Vachon, 1974), db on fixed finger proximal to est.

Legs. Moderately elongated, femur III L/ carapace L 1.09, patella III L/D 3.58; inferior carinae strongly denticulate on femur I–IV, moderately denticulate on patella I–III, weakly denticulate on patella IV; tibia III– IV with long spurs; retrolateral tarsal spurs simple, non- setose; prolateral tarsal spurs basally bifurcate, bearing 1–3 macrosetae (1 on I–II, 2 on III, 2–3 on IV); basitarsi I–III with well developed bristle-combs, clearly wider than basitarsal segment; basitarsus III setal counts (left/ right): retrosuperior 13/11, retroinferior 11/11 (including basal accessory seta), inferior 12/12; ventral surface of telotarsi with long, slender tapered macrosetae.

Measurements of male from Ad Darb (NHMB 17ag) (mm). Total L 59.50; metasoma + telson L 40.00; carapace L 6.36, W 6.54, carapace preocular L 2.75; metasomal segments (L/ W /D) I 4.99/ 4.00/ 3.53, II 6.02/ 3.63/ 3.27, III 6.28/ 3.44/ 3.12, IV 7.02/ 3.20/ 2.92, V 7.74/ 3.11/ 2.58; telson L 6.75; vesicle L 3.87, W 2.58, D 2.65; pedipalp chela L 11.27, manus ventral L 3.78, manus W 1.85, manus D 2.25, fixed finger L 6.49, movable finger L 7.31; pedipalp femur L 5.93, W 1.68, patella L 6.79, W 2.09; pectine L 7.52, mid-pectine sensillar margin L 0.841; leg III femur L 6.97; leg III patella L 5.89, D 1.65.

Measurements of holotype male (ZMHB No. 141) (mm). Total L 61.0; carapace L 6.8, W 7.0, metasomal segments (L/ W /D) I 5.30/ 4.30/ 3.50, II 6.15/ 3.90/ 3.45, III 6.50/ 3.75/ 3.40, IV 7.05/ 3.35/ 3.30, V 7.90/ 3.30/ 3.00; telson L 6.90; vesicle L 4.20, W 2.85, D 2.75; pedipalp chela L 10.80, manus ventral L 3.90, manus W 2.20, manus D 2.20, fixed finger L 6.60, movable finger L 7.90; pedipalp femur L 5.40, W 1.70, patella L 7.30, W 2.15; pectine L 8.55, mid-pectine sensillar margin L 1.025; leg III femur L 5.20; leg III patella L 6.00, D 1.65. Pectine teeth: 34–35.

Female (paratype of L. nasheri ). Differs from male as follows: body wider, carapace W/L 1.11; carapace, tergites and coxae with weaker, more finely granulate carinae, intercarinal surfaces more finely granulated or shagreened; genital opercula with slightly sinuous posterolateral margin; pectine basal piece wider with small anteromedian notch; pectines narrower, shorter, 7–8 middle lamellae; proximal margin of basal middle lamella not extended, 1.15–1.28 times longer than distal margin; pectine tips not extending past distal end of trochanter IV, teeth smaller, with short sensillar margins, mid-pectine sensillar margin L/ pectine L 0.070, mid-pectine sensillar margin L/ metasoma I W 0.115; 25–28 pectine teeth; basal pectine teeth do not overlap if anterior pectine margins aligned to posterior margins of coxae IV; sternite III–VI with median carinae obsolete, medial surfaces smooth; sternites IV–V with lateral carinae weak, smooth, lateral surfaces weakly shagreened; sternite VI with moderate, finely granulated lateral carinae, sternite VII with moderate, crenulategranulate median and lateral carinae, intercarinal surfaces finely roughened; spiracles on VI with fine granules along anterior margins; ventromedian carinae on metasoma II–III more strongly crenulate, with larger denticles; telson with sparse, fine granules on ventral surface of vesicle; metasoma and pedipalp segments less slender; pedipalp fixed fingers with 11 primary denticle subrows; prolateral tarsal spurs with 1, 2, 2–3, 3 setae on legs I–IV.

Measurements of female (mm). Total L 61.00; metasoma + telson L 38.00; carapace L 7.04, W 7.84, carapace preocular L 3.39; metasomal segments (L/ W/ D) I 4.82/ 4.10/ 3.50, II 5.54/ 3.67/ 3.47, III 5.78/ 3.47/ 3.60, IV 6.59/ 3.25/ 3.12, V 7.47/ 3.56/ 2.97; telson L 7.16; vesicle L 3.95, W 3.12, D 2.88; pedipalp chela L 12.20, manus ventral L 4.25, manus W 2.36, manus D 2.80, fixed finger L 7.05, movable finger L 8.57; pedipalp femur L 5.79, W 1.89, patella L 7.11, W 2.61; pectine L 6.78; mid-pectine sensillar margin L 0.474; leg III femur L 7.08; leg III patella L 5.77, D 1.94.

Variation. Fuscous intercarinal markings were darker in female from Al Mansuriah, much weaker in male from Ad Darb. Chela trichobothrium db was always proximal to est, but its position was variable, db -est distance/ pedipalp fixed finger L -0.0497 – -0.0056 (- 0.0208 ± 0.0132; n = 11 fingers).

DISTRIBUTION. Known only from the Tihamah plain along the Red Sea coast of western Yemen and southwestern Saudi Arabia. In addition to the material examined, we have also included the type locality of Buthus beccarii (Moka = Al Mukhah, 13°19'11''N 43°14'24''E) ( Simon, 1882). Borelli (1915) listed a male and female from Aden as L. q. brachycentrus , but we have not confirmed this record.

ECOLOGY. All collections are from low elevation coastal sites (<110 m a.s.l.). The well developed tarsal bristle combs and long setation suggest adaptation to an arenicolous life on sandy soils of coastal dunes. The Tihamah plain is a hot environment with daily high temperatures of ca. 43° C, and 40–60 % relative humidity, and the southern sites lie in the zone of coastal fog desert.

REMARKS. There has been some ambiguity about the type locality. It was cited as ‘district of Gedda (Gumfudam)’ by Levy et al. (1970: 233), and ‘Jidda (Gumfudam)’ by Levy & Amitai (1980: 47), and later transcribed as ‘Jeddah, Arabia’ by Sissom (1994: 23). This is misleading because ‘Gumfudam’ (= Al Qunfudhah, Saudi Arabia) is ca. 325 km SE of Jeddah. In Symbolae Physicae, Ehrenberg (1831) described brachycentrus as a form of A. quinquestriatus from Arabian desert ‘prope Gumfudam’ (= near Al Qunfudhah), but then writes: ‘In Arabiae felitis prope Lohaiam 3 species collegimus: Androctonus (Liurus) tunetanum  intermedium cum quinquestriato  brachycentro et Prionurum liosoma.’ Thus the type locality is actually ‘Lohaie’ (= Al Luhayyah, ca. 867 km SE of Jeddah), in Arabia Felix (= Yemen) (see also Birula, 1908; Braunwalder & Fet, 1998). According to Moritz & Fischer (1980), and confirmed here, the label stored with the type of L. brachycentrus stat. n. reads ‘Arabia Felix’.

The taxonomic status of L. brachycentrus has long been uncertain. Ehrenberg (1829, 1831) originally differentiated subspecies brachycentrus from the nominotypic form by two diagnostic characters: short aculeus, equal in length to telson vesicle, and absence of dark pigment on metasoma V. The subspecies was accepted by Birula (1908, 1917) and Borelli (1915). Birula (1937) published a more detailed redescription based on a series of specimens from Al Hudaydah, but did not study the type, and we consider his material to represent a different species, L. haenggii sp. n. Vachon (1979) has suggested that lighter coloration on metasoma V may also occur in other Leiurus populations. Levy et al. (1970) mentioned L. q. brachycentrus but did not render an opinion because they did not examine Arabian material. Later, Levy & Amitai (1980) synonymized L. q. brachycentrus with the nominate subspecies. However, they did not examine the type and assumed that L. brachycentrus was represented by specimens from Jeddah. We examined several Leiurus from Jeddah and found that they belong to different species. Our analysis of the holotype of L. brachycentrus , and similar specimens from the surrounding region on the Tihamah plain, demonstrates that this is a valid species.

Simon (1882) described Buthus beccarii based on a single male collected from Moka (=Al Mukha) on the southern Red Sea coast of Yemen. Kraepelin (1891, 1899) listed it as a synonym of L. quinquestriatus , and Birula (1908) considered it a synonym of Buthus quinquestriatus brachycentrus . This is consistent with its type locality on the southern Tihamah coastal plain, and with certain mentioned characters, i.e. relatively smaller size (total length 56.7 mm), tan yellow color with dark carinae, pectinal tooth count 34, and anterior median carinae extending continuously to anterior margin of carapace. As previously noted ( Kraepelin, 1891; Levy et al., 1970), the rostral limit of the anterior median carinae can vary in Leiurus , and is not by itself a reliable diagnostic character. Nevertheless, in specimens of L. brachycentrus that we examined, the carinae did form continuous arcs of granules extending to the anterior transverse marginal row of granules, with very little gap (typically less than one granule diameter; Figs. 3A , 7A ). In other species there was often a substantial gap, or the arrangement of granules became irregular near the anterior margin causing a break up of the carinae. Considering these characters and the locality, we regard B. beccarii to be a synonym of L. brachycentrus stat. n.

Figures

Figure 1: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., male. Habitus, dorsal aspect. Ad Darb, Saudi Arabia (NHMB 17ag).

Figure 2: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., male. Habitus, ventral aspect. Ad Darb, Saudi Arabia (NHMB 17ag).

Figure 3: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., male. A. Carapace and tergites. B. Coxosternal area and sternites. Ad Darb, Saudi Arabia (NHMB 17ag).

Figure 4: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., 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. FL. Fixed finger dentition, left chela. G. Movable finger dentition. H. Chela, external aspect. Ad Darb, Saudi Arabia (NHMB 17ag).

Figure 5: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., female. Habitus, dorsal aspect. Al Mansuriah, Yemen.

Figure 6: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., female. Habitus, ventral aspect. Al Mansuriah, Yemen.

Figure 7: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., female. A. Carapace and tergites. B. Coxosternal area and sternites. Al Mansuriah, Yemen.

Figure 8: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., 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. Al Mansuriah, Yemen.

Figure 9: Leiurus brachycentrus (Ehrenberg, 1829) stat. n.. Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Male, Ad Darb, Saudi Arabia. Female, Al Mansuriah, Yemen.

Figure 10: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., 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. Ad Darb, Saudi Arabia (NHMB 17ag).

Figure 11: Leiurus brachycentrus (Ehrenberg, 1829) stat. n., holotype male (ZMHB No. 141). A. Dorsal habitus. B. Ventral habitus. C. Right pedipalp, dorsal aspect. D. Right pedipalp, ventral aspect. E. Left pectine. F. Metasoma, lateral aspect. G. Right leg IV, telotarsus, basitarsus and distal tibia, ventral aspect. Scale bar: A, B, 10 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.