Leiurus quinquestriatus (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 : 80-88

Leiurus quinquestriatus (Ehrenberg, 1828)

( Figs. 72–82 , 87H , 88H , 89H , 90H , 91D , 92J , 93K , 94– 95 , 96C , 98–99 , Tabs. 3A , 5)

Androctonus (Leiurus) quinquestriatus Ehrenberg in Hemprich & Ehrenberg, 1828, pl. I, fig. 5.

SYNONYMS

Androctonus (Liurus) quinquestriatus aculeatus Ehrenberg in Hemprich & Ehrenberg, 1831.

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).

Androctonus (Leiurus) 5- striatus : Hemprich & Ehrenberg, 1829: 353.

Androctonus (Liurus) quinquestriatus aculeatus: Ehrenberg in Hemprich & Ehrenberg, 1831.

Androctonus quinquestriatus: C. L. Koch, 1841a: 43–44 , pl. CCLXV, fig. 622; C. L. Koch, 1850: 90.

Androctonus (Liurus) quinquestriatus aculeatus: Gervais, 1844: 46 .

Scorpio (Androctonus) quinque-striatus : Gervais, 1844: 46, pl. XXIV, fig. 2.

Androctonus troilus: C. L. Koch, 1850: 90 .

Buthus quinquestriatus: Koch, 1875: 7 ; Simon, 1879: 100; Karsch, 1881: 89; Pavesi, 1883: 96; Kraepelin, 1891: 58–60, pl. I, fig. 7, pl. II, fig. 38 (in part); Pocock, 1891: 242–243 (in part); Thorell, 1893: 359–360 (in part); Pocock, 1895: 299; Birula, 1898: 134; Kraepelin, 1899: 27–28 (in part); Kraepelin, 1901: 266–267; Werner, 1902: 598; Tullgren, 1909: 3; Birula, 1910: 118, 119–120 (in part); Simon, 1910: 70–71, fig. 9; Borelli, 1915: 461; Birula, 1917b: 4–5, figs. 1a, 2; Lampe, 1917: 191; King, 1925: 82; Borelli, 1927: 351–352; Gough & Hirst, 1927: 5, fig. 8; Borelli, 1929: 297; Schenkel, 1932: 381; Táborsky, 1934: 40 (in part); Werner, 1934: 269 (in part); Moriggi, 1941: 84; Abd-el-Wahab, 1957: 111–121, figs. 1–5.

Buthus (Buthus) quinquestriatus quinquestriatus: Birula, 1908: 124–129 ; Birula, 1917a: 223.

Buthus quinquestriatus quinquestriatus: Birula, 1910: 118 .

Buthus (Buthus) quinquestriatus: Birula, 1917a: 23 , 213 (in part); Birula, 1927: 79–80; Whittick, 1941: 43 (in part); Roewer, 1943: 206 (in part).

Leiurus quinquestriatus: Vachon, 1949: 83 , 88–93 (in part); Vachon, 1950a: 197 (in part); Vachon, 1952: 208–213, 369, 411, 441, 445, 449–452, figs. 639, 678, 686 (in part); Vachon, 1966: 211; Stahnke, 1972: 130 (in part); Probst, 1973: 329; Pérez, 1974: 24 (in part); Lamoral & Reynders, 1975: 509–510 (in part); Levy & Amitai, 1980: 47–48 (in part); Kinzelbach, 1985: map II (in part); El-Hennawy, 1987: 18; El-Hennawy, 1992: 98, 125-126 (in part); Kovařík, 1992: 183; Sissom, 1994: 20–23 (in part); Braunwalder & Fet, 1998: 32; Kovařík, 1998: 112 (in part); Fet & Lowe, 2000: 155 (in part); Lourenço et al. 2006: 98, 100 (in part), fig. 1–3; El-Hennawy, 2005: 76; El-Hennawy, 2008: 118, 121, tab IV; Lourenço & Cloudsley-Thompson, 2009: 126; Yağmur et al., 2009: 2–3 (in part); Kovařík & Ojanguren, 2013: 158, figs. 836–838.

Leiurus quinquestriatus quinquestriatus: Vachon, 1949: 88–93 (in part); Pérez, 1974: 24; Fet & Lowe, 2000: 155; Lourenço et al. 2002: 641; Lourenço et al. 2006: 97; Yağmur et al., 2009: 12.

Androctonus (Leiurus) quinquestriatus: Braunwalder & Fet, 1998: 33 .

NOTE: The above synonymy and reference listing omits records from Libya, Algeria and the southern Sinai, as we did not revise the status of those populations.

TYPE MATERIAL. 8 original syntypes (2 ♂, 2 ♀, 4 juv): Egypt, Sinai; Arabian Desert; Sudan, Dongola (= Dunqulah, Nubia); 1823–1824, leg. F.W. Hemprich & C.G. Ehrenberg ( ZMHB No. 140) (examined, Fig. 82 ).

OTHER MATERIAL EXAMINED. Egypt: 2 juvs, Lisht , 29°33'N 31°08'E, 54 m a.s.l., leg. A. Hrdlička ( USNM) ; 1 ♂, 1♀, Kena (= Qena), 26°09.77'N 32° 42.69'E ( NHMB 17g); 2 ♀, Girga, 26°19.74'N 31°52.8'E ( NHMB 17h); 2 ♂, 2 ♀, Kharga Oasis, 25°15'N 30°35'E ( NHMB 17i); 2 ♂, 2 ♀, 1 juv ♀, Cairo, 30°03'N 31°15'E, leg. Zschokke ( NHMB 17c); 1 ♂, 5 ♀, 2 juvs ♂, 2 juvs ♀, Cairo environs, 30°03'N 31°15'E, leg. E. Graeter ( NHMB 17d); 3 ♀ ( NHMB 17f); 1 ♂, 3 ♀, 2007 (GL). Sudan: 1♂, 4 ♀, Malha , West Darfur, 15°04.63'N 26°09'E, 900 m a.s.l., 20.I.1986, leg. H. Dumont ( MEB 364 ) ; 2 ♂, 7 ♀, 12 juvs, Sabaloka Mt. , ca 16°20'N 32°30'E, 24.X.–14.XI.2011, leg. P. Pokorný ( FKCP) .

DIAGNOSIS (adults). Medium to large Leiurus , 69–90 mm in length, carapace L 7.8–11.0 mm, color uniformly yellow or with variable fuscous markings on carapace and tergites; metasoma V yellow, or darkened except for posterior end; carapace with area between anterior median carinae lacking dark pigment, smooth or slightly roughened, with few sparse granules, area between posterior median carinae flat or with shallow median depression, lateral arcs of granules either very fine or absent; medial intercarinal surfaces of tergites II–III densely shagreened with fine granules, granulation weaker and sparser in females ( Figs. 93K , 94 ); posterior margin of coxa III smooth, or with scattered fine granules; metasoma moderately slender, metasoma II L/ W 1.51 –1.96, metasoma III L/ W 1.67 –2.22, metasoma IV L/ W 2.07 –2.63; ventromedian carinae of metasoma II and III with 19–26 denticles (90/92 carinae); metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps slender, patella L/W ♂ 3.17–3.85, ♀ 2.88–3.61; leg III patella L/D 3.24–4.38; pectine teeth ♂ 30–39, ♀ 26–38; pectines medium sized, pectine L/ carapace L ♂ 1.11–1.40, ♀ 0.89–1.38, midpectine sensillar margin L/ metasoma I W ♂ 0.142 – 0.184, ♀ 0.078 –0.156; basal 1–3 pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; pectine basal piece densely, shagreened, usually with more coarse medial granules in males; leg III basitarsus with 8–16 retrosuperior setae; pedipalp chela fixed finger with trichobothrium usually db distal to est; area of sternite VII between median carinae with dense, fine granulation or shagreened anteriorly, more heavily in males; sternite carination: males and females: sternite III with median carinae strong, densely granular or shagreened, sternites IV–V with lateral carinae strong, median carinae moderate.

MEASUREMENTS.

Male syntype ( Fig. 82C , subadult) (mm). Total L 63.0; metasoma + telson L 41.5; carapace L 6.6, W 7.8; metasomal segments (L/ W) I 5.4/ 4.2, II 6.3 / 3.6, III 6.3 / 3.5, IV 7.3 / 3.2, V 8.6 / 3.0; telson L 7.6; pedipalp chela L 13.1, manus W 1.9, movable finger L 8.9; pedipalp femur L 7.2, W 1.7, patella L 8.0, W 2.2; pectine L 8.5, mid-pectine sensillar margin L 0.85; pectine teeth 30–30. Adult female syntype ( Fig. 82A ) (mm). Total L 90.0; metasoma + telson L 56.7; carapace L 10.3, W 11.6; metasomal segments (L/ W) I 7.4/ 5.8, II 8.5 / 5.3, III 8.6 / 5.2, IV 10.1 / 4.7, V 11.4 / 4.4; telson L 10.7; pedipalp chela L 18.7, manus W 3.3, movable finger L 13.5; pedipalp femur L 10.1, W 2.8, patella L 10.8, W 3.5; pectine L 9.7, mid-pectine sensillar margin L 0.90; pectine teeth 30.

Male, Kena, Egypt (NHMB 17g) (mm). Total L 74.00; metasoma + telson L 48.00; carapace L 8.28, W 8.57, carapace preocular L 3.69; metasomal segments (L/ D /W) I 6.31/ 4.30/5.15, II 7.50 / 4.57/4.55, III 7.71 / 3.78/4.28, IV 8.62/ 3.41/3.87, V 9.28 / 3.22/3.68; telson L 7.62; vesicle L 3.76, D 3.04, W 3.04; pedipalp chela L 15.60, manus ventral L 4.49, manus W 2.35, manus D 2.57, fixed finger L 9.90, movable finger L 11.40; pedipalp femur L 8.22, W 2.31, patella L 9.50, W 2.81; pectine L 9.98, mid-pectine sensillar margin L 0.898; leg III femur L 8.77; leg III patella L 7.59, D 2.00; chela db–est distance (left/ right) 0.958 / 0.917; pectine teeth (left/ right) 35/ 35.

Female, Kena, Egypt (NHMB 17g) (mm). Total L 80.00; metasoma + telson L 53.00; carapace L 9.46, W 10.80, carapace preocular L 4.44; metasomal segments (L/ D /W) I 6.62/ 5.18/5.75, II 7.96 / 4.64/5.04, III 8.16 / 4.49/4.79, IV 8.99/ 3.93/4.35, V 10.11 / 3.59/4.34; telson L 9.33; vesicle L 4.63, D 3.51, W 3.62; pedipalp chela L 17.03, manus ventral L 5.14, manus W 2.63, manus D 3.14, fixed finger L 9.99, movable finger L 12.14; pedipalp femur L 8.88, W 2.75, patella L 10.12, W 3.44; pectine L 9.53, mid-pectine sensillar margin L 0.653; leg III femur L 9.71; leg III patella L 8.29, D 2.17; chela db–est distance (left/ right) 0.917 / 1.125; pectine teeth (left/ right) 28/ 28.

DISTRIBUTION. Egypt, Sudan.

REMARKS. The syntypes include assorted material from the Sinai, the Nile Valley in Egypt and Sudan, and the desert region of Egypt east of the Nile. These could represent more than one species if the populations in the Sinai are distinct from those of the Nile Valley. However, since individual specimens do not have locality labels, they cannot be referred to specific populations. We refrain from designating a lectotype until the taxonomic status of these populations is clari-fied by further study.

Figures

Figure 72: Leiurus quinquestriatus (Ehrenberg, 1828), male. Habitus, dorsal aspect. Kena, Egypt (NHMB 17g).

Figure 73: Leiurus quinquestriatus (Ehrenberg, 1828), male. Habitus, ventral aspect. Kena, Egypt (NHMB 17g).

Figure 74: Leiurus quinquestriatus (Ehrenberg, 1828), male. A. Carapace and tergites. B. Coxosternal area and sternites. Kena, Egypt (NHMB 17g).

Figure 75: Leiurus quinquestriatus (Ehrenberg, 1828), 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. Kena, Egypt (NHMB 17g).

Figure 76: Leiurus quinquestriatus (Ehrenberg, 1828), female. Habitus, dorsal aspect. Kena, Egypt (NHMB 17g).

Figure 77: Leiurus quinquestriatus (Ehrenberg, 1828), female. Habitus, ventral aspect. Kena, Egypt (NHMB 17g).

Figure 78: Leiurus quinquestriatus (Ehrenberg, 1828), female. A. Carapace and tergites. B. Coxosternal area and sternites. Kena, Egypt (NHMB 17g).

Figure 79: Leiurus quinquestriatus (Ehrenberg, 1828), 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. Kena, Egypt (NHMB 17g).

Figure 80: Leiurus quinquestriatus (Ehrenberg, 1828). Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Kena, Egypt (NHMB 17g).

Figure 81: Leiurus quinquestriatus (Ehrenberg, 1828), 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. Kharge Oasis, Egypt (NHMB 17i).

Figure 82: Leiurus quinquestriatus (Ehrenberg, 1828), 5 of 8 syntypes (ZMHB No. 140). Females: A, #12C01-3; B, #12C04-6. Males: C, #12C07-9; D, #12C10-3. Juvenile: E, #12C14-6. Egypt; Sudan. Scale bars (10 mm) apply to the habitus photos. Body lengths: A, 90 mm; B, 88 mm; C, 63 mm; D, 69 mm; E, 65 mm. Not shown are three additional fragmented juvenile syntypes.

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 96: Carapace and tergites I–III of representatives of six genera of the Buthus group. A. Compsobuthus acutecarinatus (Simon, 1882), male, Jabal Qara, Oman. B. Cicileus exilis (Pallary, 1928), male, Hoggar, Algeria. C. Leiurus quinquestriatus (Ehrenberg, 1828), female, Egypt. D. Sassanidotus gracilis (Birula, 1900), female, Hormozgan Prov., Iran. E. Odontobuthus bidentatus Lourenço & Pézier, 2002, male, Omidiyeh, Iran. F. Apistobuthus susanae Lourenço 1998, male, Albadji, Iran. Scale bars: A, B, D: 1 mm; C, E, F: 2 mm.

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).