Leiurus haenggii, 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 : 34-48

Leiurus haenggii sp. n.

( Figs. 23–34 , 46 , 58C , 59C–D , 87D , 88D , 89D , 90D , 91A , 92E–F , 93C–D , 94D–E , 95 , 98–100 , Tabs. 3B , 4 ) http://zoobank.org/urn:lsid:zoobank.org:act:E2D4F1 37-9FEA-4A9D-A0B5-07100870F6AA

REFERENCES

Buthus quinquestriatus: Pocock, 1895: 292 ; Finnegan, 1932: 91–92 (in part; record from Ain al Riqat (= Ayn Arzat)); Werner, 1936: 173.

Buthus quinquestriatus brachycentrus: Birula 1937: 102–105 .

Buthus (Buthus) quinquestriatus: Whittick, 1941: 43 (record from Yemen); Roewer, 1943: 206 (in part, record from Yemen).

Leiurus quinquestriatus voelschowi: Kovařík, 1997: 180 (in part, record from Yemen).

Leiurus quinquestriatus: Stahnke, 1972: 130 (in part); Lamoral & Reynders, 1975: 509–510 (in part); Vachon, 1979a: 49–50 (in part), figs. 37, 46-48, 64– 66; Kinzelbach, 1985: map II (in part); El- Hennawy, 1992: 101, 125–126 (in part); Sissom, 1994: 20–23 (in part), figs. 38–44; Kovařík, 1998: 112 (in part); Fet & Lowe, 2000: 155 (in part); Hendrixson, 2006: 84–91 (in part), fig. 18 (in part), fig. 19; Kovařík, 2007: 140, figs. 9–10; Kaltsas & 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, Ta’if, X.1975, leg. W. Büttiker ( NHMB 17k).

PARATYPES. Oman: 1 ♂, Wadi Ghaiz , Dhofar, under rock, 16°58'N 53°52'E, 60 m a.s.l., 11.IV.1980, leg. J.N. Barnes, MDG 5995 ( NHMB); 1 ♀, Sarfait Area, Dhofar, 16°40'N 53°06'E, 1985, leg. Sultan of Oman Land Forces ( NHMB); 1 ♀, Sarfait, Jabal Qara , 16°43'N 53°07'E, 1300 m a.s.l., 4.IX.1989, leg. R.P. Whitcomb ( NHMB); 1 ♀, Jabal Qara, Dhofar, edge of dry Nejd , 17°19'N 54°16'E, 850 m a.s.l., 6.IX.1989, leg. M.D. Gallagher MDG 8146 ( ONHM) ; 1 ♂, Salalah , Dhofar, 17°05'N 54°10'E, X.1993, leg. F. Barter ( NHMB); 1 ♀, Jabal Qara; north slopes, Nejd , UV detection rocky wadi, rocky wadi & rocky slopes, 17°17.83'N 54°05.11'E, 800 m a.s.l., 16.X.1993, 22:38 h, leg. G. Lowe ( GL) ; 1 ♂, 1 ♀, 1 juv, Jabal Qara; Salalah- Thumrait road , UV detection, rocky wadi, edge of Nejd Desert , dry zone, 17°17.26'N 54°05.36'E, 800 m a.s.l., 17.X.1993, 20:53 h, leg. G. Lowe ( NHMB); 2 ♂, 2 juvs, Jabal Qara; Salalah-Thumrait road , UV detection, wide rocky wadi, nr edge of wadi & rocky flats , 17°17.58'N 54°04.97'E, 800 m a.s.l., 17.X.1993, 21:45 h, leg. G. Lowe ( NHMB); 1 ♀, main road above Khor Rori Beach , east of Taqah , UV detection on ground, densely vegetated wadi, warm & humid with many insects , 17° 03.22'N 54°25.33'E, 50 m a.s.l., 18.X.1993, 21:24 h, leg. G. Lowe ( NHMB); 2 ♀, Salalah Plain; road to Ayn Hamran , 3.5 km off main coast road, UV detection, open plain with scattered Prosopis , 17°03.54'N 54°16.4'E, 50 m a.s.l., 18.X.1993, 22:29 h, leg. G. Lowe ( GL) ; 1 ♀, Jabal Qara , UV detection, on rock by line of date palms, 17°01.89'N 54°05.46'E, 29.IV.1995, 21:30 h, leg. J. Dundon ( BMNH) ; 2 ♂, 1 ♀, 1 juv, Jabal Qara , UV detection, 17°07.54'N 54°08.97'E, 142 m a.s.l., 30. IV.1995, leg. J. Dundon ( GL) ; 1 ♀, Khor Rori Beach (site F/23), UV detection, rocky slope of densely vegetated wadi between rocks, 17°03.18'N 54°25.51'E, 22 m a.s.l., 24. XII.2001, 02:25-04:45 h, leg. A. Winkler ( ONHM) ; 2 ♀, Wadi Khor Rori , Dhofar, 17°02'59.7"N 54°25'28.4"E, 22 m a.s.l., IX.2013, leg. T. Mazuch and P. Novák ( FKCP) ; 1 juv., UV detection 22:00–23:00 h, Taqah , 17°03.26'N 54°25.50'E, 26 m a.s.l., 21. III.2014, leg. D. Hoferek ( FKCP) ; 1 ♂, 1 ♀, 2 juvs, UV detection 20:00–22:00 h, Mirbat , 17°02.19'N 54°38.75'E, 54 m a.s.l., Adenium steppe, 22. III.2014, leg. D. Hoferek ( FKCP) . Saudi Arabia: 1 ♀, leg. W. Büttiker ( NHMB 17ae2); 1 juv ♀, Wadi Turabah, camp 2, 20°28.6'N 41°10.05'E, 1580 m a.s.l., leg. W. Büttiker ( NHMB 17ah); 3 ♂, Wadi Wajj, 18 km SW Taif, 21°10'N 40°16'E, 1810 m a.s.l., leg. W. Büttiker ( NHMB 17as); 1 ♀, Wadi Wajj, 18 km SW Taif, 21°10'N 40°16'E, 1810 m a.s.l., leg. W. Büttiker ( NHMB 17x); 1 ♀, 1 juv ♀, Taif, 21°18'N 40°24'E, 1670 m a.s.l., X.1975, leg. W. Büttiker ( NHMB 17k); 1 juv ♀, Abha Gizan km 53 , Wadi Ad Dilla , 17°51'N 42°22.8'E, 300 m a.s.l., 21.IV.1976, leg. W. Büttiker & W. Witt ( NHMB 17s); 1 ♀, 15 km W of Khamis Mushayt, 18°18.4'N 42°35.7'E, 2050 m a.s.l., 18.II.1977, leg. W. Büttiker ( NHMB 17u); 1 ♀, Adnan, 20°26'N 41°31'E, 1650 m a.s.l., 21.IX.1978, leg. W. Büttiker ( NHMB 17am); 2 ♀, Wadi Maraba, 17°53.76'N 42°23.4'E, 320 m a.s.l., 1.X.1978, leg. W. Büttiker ( NHMB 17al); 2 ♂, Wadi Qatan, 18°06.86'N 44°09.03'E, 1350 m a.s.l., 25.XI.1979, leg. W. Büttiker ( NHMB 17ad, 17af); 1 ♀, Wadi Qatan, 18°06.86'N 44°09.03'E, 1350 m a.s.l., 25.XI.1979, leg. W. Büttiker ( NHMB 17z); 1 ♀, Wadi Jaddah, 25°52'N 48°48'E, 26.XI.1979, leg. W. Büttiker ( NHMB 17y); 1 ♀, Adama, 19°20.33'N 42°01.37'E, 1770 m a.s.l., 1980, leg. W. Büttiker ( NHMB 17as2); 1 ♂, Wadi Gaanah, 18°25.8'N 41°53.6'E, 470 m a.s.l., 13.II.1980, leg. W. Büttiker ( NHMB 17ab); 1 juv ♂, Wadi Yamaniyah, 21°39'N 40°19'E, 1100 m a.s.l., 31.III.1980, leg. W. Büttiker ( NHMB 17ax); 2 ♀, Wadi Turabah, 20°29'N 41°12'E, 1480 m a.s.l., 4.IV.1980, leg. W. Büttiker ( NHMB 17am2); 2 juv ♂, 4 juv ♀, Adama, 19°20.33'N 42° 01.37'E, 1770 m a.s.l., 17.IV.1980, leg. W. Büttiker ( NHMB 17ap2); 2 juv ♂, Adama, 19°20.33'N 42°01. 37'E, 1770 m a.s.l., 17.IV.1980, leg. W. Büttiker ( NHMB 17ar2); 1 ♀, Wadi Shuqub, 20°43'N 41°10'E, 1480 m a.s.l., 21.IV.1980, leg. W. Büttiker ( NHMB 17bb); 1 juv ♂, 2 juv ♀, Wadi Turabah, 20°29'N 41°12'E, 1510 m a.s.l., 15.IX.1980, leg. W. Büttiker ( NHMB 17ad2); 1 ♀, Wadi Qatan, 18°06.86'N 44°09. 03'E, 1350 m a.s.l., 23.IX.1980, leg. W. Büttiker ( NHMB 17bf); 1 juv ♀, Wadi Sanakhah, 18°01'N 44°07'E, 1320 m a.s.l., 21.IV.1980, leg. W. Büttiker ( NHMB 17az); 2 ♂, 10 km NE of Biljurshi, 19°54.57'N 41°34'E, 2050 m a.s.l., 7.X.1980, leg. W. Büttiker ( NHMB 17ak2); 1 juv ♀, Biljurshi, 19°50.06'N 41°32. 54'E, 1840 m a.s.l., 7.X.1980, leg. W. Büttiker ( NHMB 17ay); 2 ♂, 1 ♀, Hesua, camp 1, 18°05'N 42°21'E, 1930 m a.s.l., 27.IX.1981, leg. W. Büttiker ( NHMB 17ab2); 1 ♂, 1 ♀, 2 juv ♂, Al Faraah, 20°57'N 40°12'E, 700 m a.s.l., 15.II.1982, leg. W. Büttiker ( NHMB 17aj2); 1 ♂, Wadi Wajj, 18 km SW of Taif, 21°10'N 40°16'E, 1810 m a.s.l., 23.IV.1982, leg. W. Büttiker ( NHMB 17aw); 1 juv ♂, 1 juv ♀, Wadi Maharish, 21°21.5'N 40°13'E, 1000 m a.s.l., 7.I.1983, leg. W. Büttiker ( NHMB 17au); 1 ♂, 1 ♀, 1juv ♀, Wadi Bani Malek, 21°34.48'N 39°17. 15'E, 8.II.1983, leg. W. Büttiker ( NHMB 17ah2); 1 juv ♂, Wadi Ellah, 22°35'N 41°35'E, 1480 m a.s.l., 9.IX.1983, leg. W. Büttiker ( NHMB 17be); 1 ♂, 1 juv ♀, Wadi Asidah, 20°25'N 41°12'E, 1560 m a.s.l., 10.IX.1983, leg. W. Büttiker ( NHMB 17al2); 1 ♀, Wadi Tayyah, 18°32'N 42°14'E, 825 m a.s.l., 15.IX.1983, leg. W. Büttiker ( NHMB 17av); 1 juv ♀, Wadi Dhiyan, 19°54.5'N 41°30.4'E, 1050 m a.s.l., 8.III.1984, leg. W. Büttiker ( NHMB 17bc); 1 juv ♂, 1 juv ♀ exuvium, Baha, 20°00.65'N 41°28'E, 2170 m a.s.l., 3.VIII.1984, leg. W. Büttiker ( NHMB 17at); 1 juv, Al Foqah, 19°50'N 41°51'E, 1630 m a.s.l., 10.IX.1984, leg. W. Büttiker ( NHMB); 1 juv ♂, 2 juv ♀, Al Foqah, 19°59'N 41°51'E, 1630 m a.s.l., 20.X.1984, leg. W. Büttiker ( NHMB 17aq2); 1 juv, Jebel Lebaba, 18°01.65'N 42°01.93'E, 420 m a.s.l., 24.III.1985, leg. J. Grainger ( NHMB); 1 ♂, Jebel Warjan, 23°59'N 39°10'E, 2100 m a.s.l., 2.IV.1985, leg. S. Collenette ( NHMB); 1 ♀, Fare, 22°45'N 39°48.4'E, 850 m a.s.l., 25.IV.1985, leg. W. Büttiker ( NHMB 17ba); 1 ♀, Jebel Al Lawz, 28°41.52'N 35°17.88'E, 2200 m a.s.l., 5.VI.1985, leg. J. Grainger ( NHMB 17an2); 1 ♂, 1 juv ♀, Shumaisy, 25°06'N 38° 43'E, 740 m a.s.l., 12.XI.1986, leg. W. Büttiker ( NHMB 17bh). Yemen: 1 ♂, 1 ♀, captive bred (Sanáa), M. Heule ( GL) ; 1 juv ♀, Jebel Bagim , nr Sadah, 16°55.5'N 43°47'E, 2000 m a.s.l., 27.IX.1985, W. Büttiker ( NHMB 17bd); 1 ♂, Wadí Dhahr , 15 km NW of Sanáa, 15°24.52'N 44°07.39'E, leg. Šťastný ( FKCP) ; 1 ♂, Wadithar , 14°26.6'N 44°45.5'E, 17.XI.2003, leg. P. Kabátek ( GL) ; 1 ♀, 4 juvs, Jabal Lawz , SE Sanáa, 15° 23'N 44°29'E, 2828 m a.s.l., leg. P. Kabátek ( FKCP) ; 2 ♀, 5 juvs, Al Bayda gov., At Taghiq vill. env., NW of Al Bayda by road ( Locality No. 16), 14°08.43'N 45° 25.88'E, 1968 m a.s.l., 4.-5.XI.2007, leg. D. Král ( FKCP) ; 1 ♀ ( FKCP) .

DIAGNOSIS (adults). Medium to large Leiurus , 65–97 mm in length, carapace L 7.4–11.2 mm; base color yellow to orange-yellow, with varied fuscous pigmentation on carapace, tergites and metasoma V; interocular triangle fuscous or dark; metasoma V light brown to black except for posterior end; carapace with area between anterior median carinae bearing scattered medium to coarse granules, area between posterior median carinae with deep median furrow flanked by arcs of coarse granules; medial intercarinal surfaces of tergites II–III sparsely granulated, lightly shagreened or smooth; posterior margin of coxa III smooth or with sparse fine granules; metasoma robust to moderately slender, metasoma II L/ W 1.46 –1.80, metasoma III L/ W 1.55 –1.96, metasoma IV L/ W 1.90 –2.46; ventromedian carinae of metasoma II and III with 15–28 denticles (85/91 carinae); metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps moderately slender, patella L/W ♂ 3.01–3.52, ♀ 2.60–3.14; leg III patella L/D ♂ 3.51–4.02, ♀ 3.21–3.70; pectine teeth ♂ 28–39, ♀ 24–32; pectines long, narrow, pectine L/ carapace L ♂ 1.10–1.27, ♀ 0.91 –1.15, midpectine sensillar margin L/metasoma I W ♂ 0.138 – 0.175, ♀ 0.093 –0.116; basal 1–3 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 10–15 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db distal to est; sternite VII with area between median carinae smooth or with sparse fine granulation anteriorly, more heavily in males; sternite carination: males, sternite III with median carinae robust, sternites IV–V with lateral carinae robust, median carinae moderate or weak; females, sternite III with median carinae weak or obsolete, sternites IV–V with lateral carinae moderate, median carinae weak or obsolete.

ETYMOLOGY. A patronym in honor of Dr. Ambros Hänggi, Naturhistorisches Museum, Basel, for his many contributions to science and his long support of the authors studies of Arabian scorpions.

COMPARISONS. L. haenggii sp. n. differs from L. quinquestriatus and resembles L. arabicus sp. n. in the following characters: posterior medial area of carapace with shallow to moderately deep median furrow, not flat, flanked by lateral granule arcs; medial intercarinal surfaces of tergites II–III between granule clusters smooth to sparsely shagreened or granulated ( Figs. 94D– E ); medial intercarinal surfaces of sternites smooth or lightly, finely shagreened; pectine basal piece smooth or very slightly granulated. It differs from L. arabicus sp. n. by having more robust leg, pedipalp and metasomal segments ( Fig. 46 ), and smooth or weakly granulated median carinae on sternites III–V of females ( Figs. 92E– F ).

DESCRIPTION (holotype female).

Coloration. Base color yellow or yellow-orange, carapace and tergites with extensive dark pigmentation; carapace dark on anterior interocular area, carinae, and posterior margin, light around posterior median furrow, lateral flanks outside lateral carinae pale; pretergites I– VI dark posteriorly, maculate anteriorly, III–VI with pair of pale median spots; tergites I–VI fuscous on medial and mediolateral intercarinal surfaces, with pair of pale anterior median spots, lateral flanks pale; tergite VII with slight fuscosity in anterior median area; ventromedian carinae of metasoma III–IV with dark stripes; metasoma V dark except for posterior end.

Carapace. Subrectangular, broad, W/L 1.17, with moderately sloped lateral flanks; upper surface with nearly flat posterior and medial plateau, strongly raised ocular tubercle; interocular triangle convex laterally, depressed medially; anterior margin very slightly emarginate, nearly straight, medially microdenticulate, bordered by row of medium sized granules; 5 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 nearly straight, angled outward, posterior part outwardly curved; posterior lateral carinae strong, hind end without lateral extension, projecting only slightly past posterior margin of carapace; lateral ocular carinae moderate, with medium, spaced granules; granulation: sparse patches of 17–20 small to large granules on anterolateral corners of interocular triangle, 9–10 small to medium granules in front of lateral ocular carinae; surface between anterior median carinae smooth except for 10 small to medium granules dispersed in anterior area; other intercarinal surfaces smooth except for few isolated small to medium granules; posterior median furrow shallow, broad, with few median microgranules, flanked by lateral arcs of small granules; posterior margin of carapace between posterior lateral carinae rimmed by regular series of medium granules.

Chelicera. Dorsal surface of manus smooth, with 6 short, pale microsetae, 3 near apical margin, 3 subapical, each surrounded by granules; 2–3 finer microsetae on anterolateral margins; dorsointernal carina at base of fixed finger moderately strong, 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 5 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 moderately granular anterior carinae, distal margins with coarse granules; proximal 1/3 of anterior margin of coxa III with sparse microgranulation, almost smooth; 3 long macrosetae along anterior carina of coxa II, 4 macrosetae along anterior carina of coxa III; anterior carina of coxa IV with regular small to medium granulation, with 1–2 proximal macrosetae; posterior margin of coxa IV with finely granulated carina on proximal half; sternum smooth, subtriangular with slightly concave lateral margins, deep posteromedian longitudinal sulcus and pit, 2 short macrosetae; genital opercula smooth with 5–6 short macrosetae, posterolateral margins weakly concave.

Pectines. Basal piece with concave anterior margin divided by small median emargination, lacking granules, bearing 9 macrosetae; pectines narrow, tips not extending past distal end of coxa IV; both combs with 3 marginal lamellae and small accessory lamella distal to first marginal lamella, 9 middle lamellae, 30–30 teeth; marginal and middle lamellae with moderate cover of long reddish macrosetae; fulcra with 3–5 setae; pectine teeth relatively small, mid-pectine sensillar margin L/ pectine L 0.063, mid-pectine sensillar margin L/ metasoma I W 0.105.

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, strong, uniformly granular, hind ends extending past posterior margins of tergites, without lateral extensions on I, with short extensions on II; medial intercarinal surfaces smooth, with sparse medium or coarse granules; lateral flanks moderately sloped with sparse fine granulation; tergites III–VI with 3 straight carinae with medium granules; medial intercarinal surfaces smooth, with short transverse anterior series of granules; very fine granulation on anterior median patch and short transverse strips on either side; lateral surfaces moderately sloped, well granulated, III – IV 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 isolated lateral patches on either side; posterior margins of tergites I – VI rimmed with regular rows of small to medium sized granules; posterior margin of tergite VII with regular small granules; sternites: sternite III with median carinae very weak, smooth, almost obsolete; sternites IV–V with weak, finely granulated lateral carinae, obsolete median carinae; sternite VI with moderate, finely crenulate lateral carinae, weak finely granulated median carinae; sternite VII with strong, crenulategranulate median and lateral carinae; lateral margins of sternites IV–VII armed with regular denticulate granules; medial intercarinal surfaces of all sternites smooth, lateral intercarinal surfaces smooth posteriorly, smooth to faintly shagreened anteriorly on IV–VI; setation: sternite III with 5 macrosetae on median carinae, 6–7 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–V with mid-carinal and posterior marginal macrosetae, on VI with posterior marginal macrosetae; intercarinal macrosetae: one pair of lateral marginal setae on sternites IV–VI; 2 pairs of mediolateral setae on IV–V, one pair on VI; one pair of latero-marginal setae on VII; intercarinal posterior marginal macrosetae on III–VII: 7, 5, 4, 0, 0.

Metasoma. Moderately long, with robust segments, total metasoma and telson L/ carapace L 5.61; carination: segment I with 10 complete carinae; segments II–III with 8 complete carinae, median lateral carinae restricted to posterior 1/2 of II, posterior 1/2 to 1/3 of III; metasoma IV with 8 carinae, V with 7 carinae; carinae on segments I–IV granulate or crenulate-granulate; dorsosubmedian carinae granulate, moderate on I–III, weak on IV; dorsolateral carinae granulate, moderate, ventrolateral carinae granulate-crenulate, moderate to strong on I–IV; median lateral carinae granulate, moderate on I–II, moderate posteriorly, weak anteriorly on III; ventromedian carinae moderate on I, strong on II–IV; granules on II–IV larger, taller posteriorly; ventromedian carinae with 20–21 granules on metasoma II, 25–26 on III; metasoma V with dorsolateral carinae very weak, faintly granulated, ventrolateral carinae strong with rounded dentate granules increasing in size posteriorly, with several large subtriangular, lobate denticles, ventrosubmedian carinae marked by prominent series of medium to large rounded, dentate granules along length of segment, ventromedian carina strong, with medium to large rounded, dentate granules increasing in size posteriorly; 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, lightly shagreened or finely granulate laterally and 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 slightly shorter than vesicle.

Pedipalp. Femur: moderately slender, L/ W 3.28; dorsoexternal, dorsointernal and ventrointernal carinae strong with coarse, closely spaced dentate granules; internal carina strong, with small and large dentate granules spaced well apart; external carina moderate, with well spaced coarse dentate granules; all intercarinal surfaces smooth; linear cluster of 29–30 accessory macrosetae on lower distal external surface; patella: moderately slender, L/ W 2.89; dorsointernal carina moderate with medium granulation; dorsomedian and dorsoexternal carinae moderate with fine granulation; external carina moderate, smooth; ventroexternal carina weak, with fine granulation; ventromedian carina moderate, with medium to fine granulation; ventrointernal carina moderate, medium to small dentate granules and ventral patellar spur; internal carina moderate, with several larger dentate granules interspersed with closely spaced small granules, and dorsal patellar spur; all intercarinal surfaces smooth or with sparse, fine granules; chela: slender, L/ W 5.94, movable finger L/ manus ventral L 2.10; dorsal marginal and external secondary carinae 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 left movable finger (right movable finger of holotype with anomalous dentition, 6 subrows), 11–12 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 elongated, femur III L/ carapace L 1.05, patella III L/D 3.61; inferior carinae strongly denticulate on femur I–IV, moderately denticulate on patella I–III, 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 2–3 macrosetae; basitarsi I–III with well developed bristle-combs, at least as wide as basitarsal segment; basitarsus III setal counts (left/ right): retrosuperior 11/10, retroinferior 13/12 (including basal accessory seta), inferior 12/13; ventral surface of telotarsi with robust, short tapered macrosetae.

Measurements of holotype female (mm). Total L 85.00; metasoma + telson L 54.00; carapace L 9.62, W 11.24, carapace preocular L 4.69; metasomal segments (L/ W /D) I 6.92/ 6.01/ 5.10, II 8.19/ 5.30/ 4.89, III 8.42/ 5.00/ 4.81, IV 9.64/ 4.65/ 4.55, V 10.81/ 4.60/ 4.18; telson L 10.48; vesicle L 5.58, W 3.11, D 3.92; pedipalp chela L 18.46, manus ventral L 6.13, manus W 3.11, manus D 3.50, fixed finger L 10.77, movable finger L 12.88; pedipalp femur L 9.37, W 2.86, patella L 10.53, W 3.64; pectine L 10.04, mid-pectine sensillar margin L 0.633; leg III femur L 10.06; leg III patella L 8.58, D 2.38.

Paratype male ( Wadi Asidah ). Differs from holotype female as follows: body narrower, carapace W/L 1.13; carapace, tergites and coxae with stronger, more coarsely granulate carinae; genital opercula with convex posterolateral margin; pectine basal piece narrower, with deeper anteromedian fold; pectines wider, longer, 11–13 middle lamellae, tips extending to proximal 1/3–1/2 of trochanter IV, teeth larger with longer sensillar margins, mid-pectine sensillar margin L/ pectine L 0.084, midpectine sensillar margin L/ metasoma I W 0.146; 36–38 pectine teeth; 3 basal pectine teeth overlap if anterior pectine margins aligned to posterior margins of coxae IV; sternite III with strong, thick, granulated median carinae; sternites IV–V with stronger, more heavily granulated lateral carinae, IV with moderate, V with weak granulated median carinae; sternites VI–VII with all carinae strong, coarsely granulated; spiracles on IV– VI with fine granules along anterior margins; intercarinal surfaces of sternites III–VI roughened, shagreened in anterior median areas; ventromedian carinae on metasoma II–III more weakly crenulate, with smaller denticles; dorsal surface of metasoma V more heavily shagreened; pedipalp femur and patella with stronger carination, pedipalp chela manus with weak dorsointernal carina with small granules.

Measurements of paratype male (NHMB 17al2) (mm). Total L 82.50; metasoma + telson L 55.00; carapace L 9.00, W 10.20, carapace preocular L 4.46; metasomal segments (L/ W/ D) I 6.90/ 6.42/ 5.47, II 8.39/ 5.73/ 5.10, III 8.68/ 5.36/ 4.93, IV 9.77/ 4.95/ 4.64, V 10.46/ 4.59/ 4.01; telson L 9.75; vesicle L 5.11, W 3.87, D 3.68; pedipalp chela L 18.59, manus ventral L 5.97, manus W 3.06, manus D 3.33, fixed finger L 11.13, movable finger L 13.02; pedipalp femur L 9.62, W 3.00, patella L 10.62, W 3.53; pectine L 11.17; midpectine sensillar margin L 0.939; leg III femur L 10.70; leg III patella L 8.81, D 2.38.

Variation. Color: the dark dorsal color pattern of the holotype was the most common form, but other variants were observed. There may be less intense, diffuse fuscosity on carapace and tergites, with brown color extending onto lateral flanks of tergites I–VI (Adnan, Adama). Intercarinal fuscosity on the carapace may be limited to the anterior area in front of the median ocular tubercle, with posterior areas light (Wadi Maraba, Jeddah), and tergites may be pale (Jeddah). Dark pigment on metasoma V and ventral carinae of metasoma II–IV can vary from faded yellow-brown to dark black, and the posterior yellow area on metasoma V may extend forward up to 1/3 the length of the segment, with the dark zone terminating in trident pattern towards the posteroventral area (Jeddah, Wadi Qatan). Juveniles usually exhibit darker color patterns. Morphosculpture: female sternite carination was typically weak or obsolete, with minor variation: median carinae of sternite III obsolete in 9 cases, very weak in 7, weak in 3, median carinae of sternites IV (or V) obsolete in 17 (or 16) cases, weak in 2 (or 3). Morphometrics and meristics: see Table 4 . Metasomal segments were typically robust, but were more elongated in a few examples: males from Wadi Ghaiz and Jeddah, and a female from Wadi Qatan had metasomal segments as slender as L. arabicus sp. n. The number of denticles on ventromedian carinae of metasoma III was significantly higher in males (22.93 ± 2.81) than females (20.16 ± 3.24) (P = 0.0085, t test).

DISTRIBUTION. Records indicate a wide distribution in the western Arabian Peninsula, along the chain of mountains running parallel to the Red Sea coast of Saudi Arabia and Yemen (Al Hijaz and Asir mountains). A series of specimens was also collected from the Dhofar mountains in Oman, and the species is likely to occur in the intervening Hadramaut ranges along the southern coast of Yemen. Records span a wide range of elevations (22 – 2,828 m a.s.l.), ranging from coastal plains to the Asir highlands.

ECOLOGY. Specimens in Oman were found at night by UV detection on rock and gravel substrates in densely vegetated wadis, from coastal plains to mountains. The species probably shelters in scrapes excavated in soil beneath rocks. Similar lapidicolous habitats may be occupied in the mountains of Yemen and western Saudi Arabia.

REMARKS. Birula (1937) referred a series of specimens from Hodeida (= Al Hudaydah, Yemen) to L. quinquestriatus brachycentrus . We consider these to represent L. haenggii , based on the following reported characters: larger size (adult female 86.4 mm, adult male 72.6 mm), dark coloration of carapace, mesosoma and metasoma V, and nearly smooth intercarinal surfaces on tergites.

Figures

Figure 23: Leiurus haenggii sp. n., holotype female. Habitus, dorsal aspect. Ta’if, Saudi Arabia (NHMB 17k).

Figure 24: Leiurus haenggii sp. n., holotype female. Habitus, ventral aspect. Ta’if, Saudi Arabia (NHMB 17k).

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 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 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 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.