Leiurus Ehrenberg, 1828

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 : 3-5

Genus Leiurus Ehrenberg, 1828

Androctonus (Leiurus) Ehrenberg in Hemprich & Ehrenberg, 1828, pl. I, fig. 5; Hemprich & Ehrenberg, 1829: 353 (in part).

REFERENCES

Androctonus (Liurus) : Hemprich & Ehrenberg, 1831 (in part); Sundevall, 1833: 33.

Leiurus: Vachon, 1949: 83–88 ; Vachon, 1950a: 199– 200, 202, figs. 550–552; Vachon, 1950b: 393, fig. 589; Vachon, 1952: 203–208, 372–374, 399, figs. 550–552; Bücherl, 1964: 57; Stahnke, 1972: 130; Lamoral & Reynders, 1975: 509; Levy & Amitai, 1980: 14, 46–47; Francke, 1985: 9, 15; Vachon & Kinzelbach, 1987: 92; Sissom, 1990: 101; Nenilin & Fet, 1992: 17; Kovařík, 1998: 112; Fet & Lowe, 2000: 155; Soleglad & Fet, 2003a: 26–27, tab. 2, fig. 44; Soleglad & Fet, 2003b: 88, 91, tab. 9; Fet et al., 2005: 10–12, tab. 1, fig. 23, 29; Prendini & Wheeler, 2005: 481; Hendrixson, 2006: 81–82; Dupré, 2007: 7; Kovařík, 2009: 24; El-Hennawy, 2009: 122.

TYPE SPECIES. Androctonus (Leiurus) quinquestriatus Ehrenberg, 1828 .

ETYMOLOGY. Two subgenera of Androctonus were created by Ehrenberg: Leiurus (= ‘slender tail’), including species with more elongated metasomal segments, and Prionurus (= ‘saw tail’) including species with heavier metasomal segments bearing serrated carinae.

DIAGNOSIS. Medium to large buthids ( Sissom, 1990), adults 51–115 mm; carapace and tergites with granulated carinae; carapace with distinct anterior, superciliary, central median, central lateral, posterior median and posterior lateral carinae; central lateral and posterior median carinae fused into a lyre configuration; tergites I–II, VII with 5 carinae, III–VI with 3 carinae; metasoma elongate, metasoma I–III with 10 carinae, median lateral carinae complete on I, reduced on II–III; metasoma IV with 8 carinae; metasoma V with 7 carinae, dorsolateral carinae weak, ventrolateral carinae strong with serrate or lobate dentition; telson with bulbous vesicle lacking subaculear spine or tubercle; pectines with fulcra; chelicerae with characteristic buthid dentition ( Vachon, 1963), two denticles on ventral aspect of fixed finger; pedipalps orthobothriotaxic, type A  ( Vachon 1974, 1975); pedipalp femur with trichobothrium d 2 on dorsal surface, pedipalp patella with d 3 internal to dorsomedian carina; chela smooth with carinae reduced or obsolete, dentate margins of fingers armed with linear subrows of primary denticles; subrows flanked by internal and external accessory denticles; movable finger with two enlarged subdistal internal denticles; males without scalloping at base of pedipalp fingers; tergites without macrosetae; tibial spurs present on legs III–IV; basitarsi I–III with regular series of macrosetae on retrosuperior, retroinferior and inferior margins; ventral surfaces of telotarsi with paired rows of macrosetae; prolateral and retrolateral tarsal spurs present on all legs. Sexual dimorphism: compared to females, males have a narrower mesosoma, more robust carination on tergites and sternites III–V, more slender pedipalps and metasoma, longer pectines with larger teeth, and weaker dentition or granulation on ventromedian carinae of metasoma II–III.

SPECIES COMPOSITION. The broad geographic range and morphological variation of Leiurus populations have caused difficulty for taxonomists. For a long time, the genus was assumed to be represented by a single polytypic species spread over a vast area, although different authors disagreed about the definition and validity of various subspecies. More recently, several distinctive species were described from widely disparate localities: L. jordanensis Lourenço et al., 2002 , from sandstone cliffs isolated by dunes in Jordan and northern Saudi Arabia; L. savanicola Lourenço et al., 2006 , from Sahel/ savannah in Cameroon; L. nasheri Kovařík, 2007 , from the humid Tihamah coastal plain in Yemen (= L. brachycentrus (Ehrenberg, 1829) stat. n.); and L. abdullahbayrami Yağmur et al., 2009 , from semi- arid rocky steppe in Turkey and northern Syria. Here we add four new species from the Arabian Peninsula: L. haenggii sp. n., from mountain ranges along the Red Sea coast (Hijaz and Asir) in Saudi Arabia, the Hadramaut in Yemen, and Dhofar mountains of Oman; L. arabicus sp. n., from arid plains and wadis of the central Najd plateau, extending to the east coast of Saudi Arabia; L. macroctenus sp. n., from coastal fog desert of Jiddat al Harasis in Oman; and L. heberti sp. n. from rocky wadis of the Jabal Samhan mountains in Dhofar, Oman. In addition, we propose revised diagnoses for: L. hebraeus ( Birula, 1908) stat. n. from semi-arid/ arid steppe habitats in Israel, Jordan and Syria; L. brachycentrus stat. n. from the Tihamah plain along the Red Sea coast of western Yemen and southwestern Saudi Arabia; and for the type species L. quinquestriatus from the Nile Valley and surrounding deserts in Egypt and Sudan, bringing the total number of included species to ten. Our findings show that, like many other scorpion genera, Leiurus is comprised of an assemblage of allopatric or parapatric species spread across different regions separated by physiographic barriers, each adapted to local environments and substrates. Additional species diversity may emerge when other local populations are analyzed in more detail, for example those in southern Sinai, and in more central parts of North Africa.

Key to adult Leiurus species examined in this study

1 Medial intercarinal surfaces of tergites II–III smooth or sparsely, lightly shagreened .....……………...... 2 Medial intercarinal surfaces of tergites II–III heavily or densely, finely shagreened .......……….............. 4

2 Males with metasoma III L/W> 2.10, metasoma IV L/W> 2.50; metasoma III ventromedian carinae with> 35 denticles ......................... L. heberti sp. n. Males with metasoma III L/W <2.10, metasoma IV L/W <2.50; metasoma III ventromedian carinae with <35 denticles ...............…………….............. 3

3 Females with pedipalp patella L/W> 3.20, F s> 23; female sternites III–IV with weak to moderate median carinae .........……............ L. arabicus sp. n. Females with pedipalp patella L/W <3.20, F s <23; female sternites III–IV with weak to obsolete median carinae ...…..…............... L. haenggii sp. n.

4 Male pectines with mid-pectine sensillar margin L/ metasoma I W> 0.20 .....…...……......................... 5 Male pectines with mid-pectine sensillar margin L/ metasoma I W <0.20 ......………........................... 6

5 Female pectines with mid-pectine sensillar margin L/ metasoma I W> 0.15 …... L. macroctenus sp. n. Female pectines with mid-pectine sensillar margin L/ metasoma I W <0.15 ........……........................... ............. L. brachycentrus (Ehrenberg, 1829) stat. n.

6 Metasoma III ventromedian carinae with> 30 denticles; metasoma I–IV uniformly fuscous ……......... ….... L. jordanensis Lourenço, Modrý et Amr, 2002 Metasoma III ventromedian carinae with <30 denticles; metasoma I–IV yellow to yellow-orange…. 7

7 Sternite VII with medial intercarinal surface smooth ..... L. abdullahbayrami Yağmur, Koç et Kunt, 2009 Sternite VII with medial intercarinal surface shagreened .........…..…................................................ 8

8 Carapace with anteromedian intercarinal surface smooth or sparsely shagreened; posterior median furrow of carapace shallow or flat, lateral granule arcs reduced or absent; females with pectine basal piece heavily shagreened, median carinae of sternites IV–V moderate, usually granular; males with pedipalp patella L/W> 3.1, metasomal segment L/W: II> 1.55, III> 1.7, IV> 2.05 .....…….......... …………...… L. quinquestriatus (Ehrenberg, 1828) Carapace with anteromedian intercarinal surface shagreened; posterior median furrow of carapace moderately deep, flanked by lateral granule arcs; females with pectine basal piece smooth or lightly shagreened, median carinae of sternites IV–V weak, smooth or obsolete; males with pedipalp patella L/W <3.1, metasomal segment L/W: II <1.55, III <1.7, IV <2.05 ... L. hebraeus ( Birula, 1908) stat. n.