Leiurus hebraeus ( Birula, 1908 )

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 : 73-79

Leiurus hebraeus ( Birula, 1908) stat. n.

( Figs. 60–71 , 87F , 88F , 89F , 90F , 91G , 92K , 93I–J , 94D–E , 95 , 98–99 , Tabs. 3A , 5)

Buthus quinquestriatus hebraeus Birula, 1908: 124–129 .

REFERENCES

Scorpio occitanus : Audouin in Savigny, 1826: 173, pl. VIII, fig. 1 (in part); Audouin in Savigny, 1827: 410-411, pl. VIII, fig. 1 (in part).

Buthus quinquestriatus: Kraepelin, 1891: 58-60 , pl. I, fig. 7, pl. II, fig. 38 (in part); Pocock, 1891: 242– 243 (in part); Pavesi, 1895: 4; Kraepelin, 1899: 27- 28 (in part); Simon, 1892: 83 (?); Davydov, 1898: 41–43; Schenkel, 1932: 380; Werner, 1934: 269 (in part); Werner, 1935: 211; Bodenheimer, 1937: 235; Whittick, 1955; Whittick, 1970.

Buthus quinquestriatus hebraeus: Birula, 1910: 118 .

Buthus (Buthus) 5- striatus hebraeus: Birula, 1910: 170 .

Buthus (Buthus) quinquestriatus: Birula, 1917a: 23 , 213 (in part); Whittick, 1941: 43 (in part).

Leiurus quinquestriatus: Vachon, 1950a: 197 (in part); Stahnke, 1972: 130 (in part); Lamoral & Reynders, 1975: 509–510 (in part); Wahbeh, 1976: 89; Levy & Amitai, 1980: 47–48 (in part); Warburg et al., 1980: 206; Kinzelbach, 1984: 100, Fig 2 ; Kinzelbach, 1985: map II (in part); Amr et al., 1988: 373; El- Hennawy, 1988: 14–15, 18; El-Hennawy, 1992: 101, 125–126 (in part); Amr & El-Oran, 1994: 186; Kovařík, 1998: 112 (in part); Kabakibi et al., 1999: 80, fig. 1; Amr & Abu Baker, 2004: 237–238, 241; Yağmur et al., 2009: 2–3 (in part), fig. 10–11; Shehab et al., 2011: 335–337, fig. 2D, tabs. 1–2.

Leiurus quinquestriatus hebraeus: Vachon, 1966: 212 ; Levy, Amitai & Shulov, 1970: 231–242 (in part); Levy & Amitai, 1980: 48–53, fig. 47–51, map 3, Appendix (in part); El-Hennawy, 1992: 101, 126; Sissom, 1994: 23; Fet & Lowe, 2000: 156–157; Yağmur et al., 2009: 12; Khalil & Yağmur, 2010: 2.

Leiurus quinquestriatus voelschowi: Pohl, 1967: 209– 215 , figs. 1–4; Kovařík, 1997: 180 (in part, record from Jordan).

Leirus (sic) quinquestriatus hebraeus : Levy & Amitai, 1980: Appendix.

TYPE MATERIAL. Lectotype adult ♀, 4 immature paralectoptypes, Jordan, Wadi ‘ Arrud ( ZISP 578 ) (not examined, Figs. 70–71 ).

OTHER MATERIAL EXAMINED. Israel: 3 ♂, 7 ♀, 15 juvs, Palaestina (ev. Jordanien) ( NHMB 17a); 1 ♀, Palaestina (ev. Jordanien) , Jericho , 31°30.6'N 35°16.8'E, IV.1927, leg. O. Wohlberedt ( NHMB 17-Ia); 2 ♀, ca. 11 km S of Beersheba, Hadarom , 31°06.19'N 34°49.41'E, 350 m a.s.l., 15.IV.1983, leg. A.M. de Saint Michel ( MEB 034 ) ; 1 ♂, 2 juvs, between Nesher & Yagur, Haifa District, 32°45.22'N 35°03.69'E, 27 m a.s.l., 20.IX.1984, leg. M. R. Warburg ( MEB 197 ) ; 1 ♂, 1 ♀, Haifa area, 32°48'N 34°59'E, 300 m a.s.l., 30.XI.1984, leg. M. R. Warburg ( MEB 198 ) ; 1 ♂, 3 ♀, 3 juvs, Negev desert, Vadi Hazaz near Sede Boqer (Haluqim Ridge), XI.- XII.2004, leg. J. Král ( FKCP); 1 ♂, 2 ♀, Negev desert, Sede Boqer , IX.2007, leg. J. Král and M. Forman ( FKCP) . Jordan: 1 ♂, 4 ♀, Madaba, 31°43'N 35°48'E, 764 m a.s.l., 22. III.1983, leg. M.A. Jafar ( MEB 346 ) ; 1 ♀, Tabaqat , 32°27'N 35°37'E, – 200 m a.s.l., 4. V.1995, leg. V. Sejva (GL) ; 1 ♀, Kurayyima Udoli , 32°16.75'N 35°35.87'E, 23. VII.2000 (GL). Syria: 4 ♂, Bosra, V.1994, leg. D. Modrý ( FKCP); 1 ♀, Seydmaya, V. 1994, leg. D. Modrý ( FKCP).

DIAGNOSIS (adults). Small to medium sized Leiurus , 58–77 mm in length, carapace L 6.5–9.3 mm; base color yellow to orange-brown, with variable fuscosity on carapace and tergites; metasoma V either clear or fuscous except for posterior end; carapace with area between anterior median carinae finely shagreened, with scattered medium to fine granules, area between posterior median carinae with shallow median furrow flanked by arcs of medium or fine granules; medial intercarinal surfaces of tergites II–III shagreened, with variable medium to fine granulation; posterior margin of coxa III smooth or with sparse fine granules; metasoma robust, metasoma II L/ W 1.38 –1.64, metasoma III L/ W 1.49 –1.69, metasoma IV L/ W 1.71 –2.03; ventromedian carinae of metasoma II and III with 14–19 denticles (46/48 carinae); metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps relatively stout, patella L/W ♂ 2.81–2.95, ♀ 2.48–2.83; leg III patella L/D 3.13– 3.71; pectine teeth ♂ 33–37, ♀ 27–32; pectines medium sized, pectine L/ carapace L ♂ 1.14–1.29, ♀ 0.92–1.21, mid-pectine sensillar margin L/ metasoma I W ♂ 0.169 – 0.180, ♀ 0.100 –0.136; basal 1–2 pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; pectine basal piece very lightly, finely shagreened, almost smooth in females, almost smooth or weakly granulated in males; leg III basitarsus with 7–12 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db either distal or proximal to est; sternite VII with area between median carinae densely, finely microgranulated or shagreened; 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 or obsolete, sternites IV–V with lateral carinae weak to moderate, median carinae obsolete.

MEASUREMENTS.

Male, Israel ( NHMB 17 a) (mm). Total L 58.0; metasoma + telson L 36.0; carapace L 6.74, W 7.22, carapace preocular L 3.05; metasomal segments (L/ D /W) I 4.98/ 4.01/4.34, II 5.82 / 3.63/3.97, III 6.08 / 3.56/3.85, IV 6.92/ 3.44/3.59, V 7.67 / 2.89/3.49; telson L 7.00; vesicle L 3.85, D 2.78, W 2.87; pedipalp chela L 11.62, manus ventral L 3.51, manus W 1.93, manus D 2.14, fixed finger L 6.79, movable finger L 7.98; pedipalp femur L 5.98, W 2.03, patella L 7.36, W 2.55; pectine L 7.83, mid-pectine sensillar margin L 0.735; leg III femur L 6.87; leg III patella L 5.79, D 1.825; chela db–est (left/ right) distance 0.292 / 0.958; pectine teeth (left/ right) 35/ 35.

Female, Israel ( NHMB 17 a) (mm). Total L 76.0; metasoma + telson L 44.0; carapace L 8.09, W 8.91, carapace preocular L 3.82; metasomal segments (L/ D /W) I 5.70/ 4.30/4.83, II 6.44 / 4.15/4.30, III 6.48 / 4.05/4.10, IV 7.53/ 3.82/3.84, V 9.06 / 3.49/4.06; telson L 8.41; vesicle L 4.53, D 3.28, W 3.49; pedipalp chela L 14.34, manus ventral L 4.28, manus W 2.41, manus D 2.56, fixed finger L 8.20, movable finger L 10.03; pedipalp femur L 6.88, W 2.23, patella L 8.32, W 3.05; pectine L 8.83, mid-pectine sensillar margin L 0.653; leg III femur L 8.15; leg III patella L 6.92, D 2.00; chela db–est (left/ right) 0.354 / 0.417; pectine teeth (left/ right) 30/ 30.

DISTRIBUTION. Israel, Jordan, Syria, Lebanon.

REMARKS. Birula (1908) separated L. q. hebraeus from the nominotypic subspecies by 11 characters. We assessed these characters by comparing 20 adult L. hebraeus from Israel and Jordan with 26 adult L. quinquestriatus from Egypt and Sudan:

(1) weaker carapace carination and granulation, (2) granulated ocular tubercle, (3) more coarsely shagreened tergites, (4) non-granulated intercarinal surfaces on the metasoma: (1) – (4) were unreliable when granulation was visualized in fine detail by UV fluorescence;

(5) ventromedian carinae of metasoma II – III with denticles increasing in size posteriorly (vs. uniform in size): this was confirmed for L. hebraeus , but also observed in some L. quinquestriatus , and hence was not diagnostic;

(6) smaller number of denticles (15 – 17) on ventromedian carinae of metasoma III (vs. 18 – 27): this was well supported with no overlap, i.e. 46/46 carinae of L. quinquestriatus with 19 or more denticles, 24/24 carinae of L. hebraeus with 18 or fewer denticles;

(7) lower range of denticles (18 – 31) on ventrolateral carinae of metasoma V (vs. 23 – 38): due to the broad overlap of ranges, this was not diagnostic;

(8) more stout pedipalp and metasomal segments: this was supported by smaller morphometric ratios, pedipalp patella L/W and metasoma III – IV L/W;

(9) metasoma I only barely wider than metasoma II (vs. significantly wider): this was not diagnostic, due to significant overlap of the ratio metasoma I W/ metasoma II W ( L. quinquestriatus 1.10 – 1.21, L. hebraeus 1.09 – 1.14);

(10) telson vesicle longer than aculeus (chord length), vesicle width equal to aculeus length (vs. about the same length or shorter, and width <aculeus length): there was a trend for L. quinquestriatus to have a relatively longer aculeus and a shorter, more bulbous vesicle than L. hebraeus , but there was significant overlap in the ratio of vesicle L/ telson L ( L. quinquestriatus 0.48 – 0.56, L. hebraeus 0.52 – 0.61);

(11) pedipalp movable finger short, less than twice manus length, with 12 denticle subrows (vs. long, slender,> twice manus length, and 13 subrows): this was not diagnostic, because although the movable finger was on average relatively shorter in L. hebraeus vs. L. quinquestriatus , there was broad overlap in the ratio movable finger L (chord)/ manus ventral L, and movable fingers had either 12 or 13 subrows in both species.

Levy et al. (1970) accepted characters (5) and (10) of Birula (1908), and proposed 3 additional characters to differentiate L. q. hebraeus :

(12) posterior median area of carapace granulated, with paired granule arcs (vs. smooth or sparsely granulated): this difference was confirmed for most specimens, although some L. quinquestriatus have weak arcs of fine granulation;

(13) lateral flanks of tergite V with irregular granulation (vs. short rows of granules, resembling carinae): this rather subjective character was quite variable, and short granule rows could be discerned or not in both L. hebraeus and L. quinquestriatus ;

(14) weak or obsolete carinae on sternites (slightly more distinct on posterior segments) (vs. distinct, elevated strong carinae on sternites IV–VII): this difference applies to males, however females have weaker carinae and some L. quinquestriatus females can have weak to obsolete carination on sternites IV–V.

Thus, from the above list, only characters (6), (14) (males only), and perhaps (12), were potential diagnostic characters separating L. hebraeus from L. quinquestriatus . Four examined adult and subadult males from southern Sinai resembled L. quinquestriatus in having slender pedipalp and metasomal segments, and reduced granulation between the posterior median carinae of the carapace. However, they grouped with the northern populations from Israel and Jordan in other morphometrics (i.e. lower metasoma I W/ II W, lower pedipalp movable finger L/ manus ventral L, higher vesicle L/ telson L) and meristics (number of denticles on ventromedian carinae of metasoma II–III). Levy et al (1980) also noted that the southern populations of L. q. hebraeus had more slender appendages. Partial overlap of characters suggests that there could be two subspecies, L. q. quinquestriatus and L. q. hebraeus , with a hybrid zone in southern Sinai. However, there appears to be a physical barrier preventing gene flow between these populations because Leiurus is excluded from the sandy coastal plain connecting North Africa and the Sinai Peninsula ( Levy & Amitai, 1980). Genetic isolation is supported by divergent evolution of distinct polypeptide toxins in their venoms ( Smertenko et al., 2001) and different physicochemical profiles of venom proteins ( Nascimento et al., 2006). Here, we propose to diagnose the African and Middle Eastern populations as distinct species by a combination of morphological characters. The southern Sinai populations occur in a different habitat (i.e. rocky mountains), and may belong to other undescribed species.

Figures

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

Figure 60: Leiurus hebraeus (Birula, 1908) stat. n., male. Habitus, dorsal aspect. Israel (NHMB 17a).

Figure 61: Leiurus hebraeus (Birula, 1908) stat. n., male. Habitus, ventral aspect. Israel (NHMB 17a).

Figure 62: Leiurus hebraeus (Birula, 1908) stat. n., male. A. Carapace and tergites. B. Coxosternal area and sternites. Israel (NHMB 17a).

Figure 63: Leiurus hebraeus (Birula, 1908) 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. Chela, external aspect. Israel (NHMB 17a).

Figure 64: Leiurus hebraeus (Birula, 1908) stat. n., female. Habitus, dorsal aspect. Israel (NHMB 17a).

Figure 65: Leiurus hebraeus (Birula, 1908) stat. n., female. Habitus, ventral aspect. Israel (NHMB 17a).

Figure 66: Leiurus hebraeus (Birula, 1908) stat. n., female. A. Carapace and tergites. B. Coxosternal area and sternites. Israel (NHMB 17a).

Figure 67: Leiurus hebraeus (Birula, 1908) 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. Israel (NHMB 17a).

Figure 68: Leiurus hebraeus (Birula, 1908) stat. n.. Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Male and female, Israel (NHMB 17a).

Figure 69: Leiurus hebraeus (Birula, 1908) stat. n., 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. A–D: female, Israel (NHMB 17a). E: female, Jericho, Israel (NHMB 17I-a).

Figure 70: Leiurus hebraeus (Birula, 1908) stat. n., lectotype female, designated here (ZISP 578). Habitus. A. Dorsal aspect. B. Ventral aspect. Wadi ‘Arrud, Jordan.

Figure 71: Leiurus hebraeus (Birula, 1908) stat. n., lectotype female and four paralectotypes, designated here (ZISP 578). Wadi ‘Arrud, Jordan. Label text in Russian: “Det. A. Birula, 5 ex. Coll. 25.IV.1897, Petro-Arabia, Wadi-Arrud, Southern shore of the Dead Sea, Davydov.”

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).