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Figure 98 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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).

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Figure 100 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 96 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 93 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 95 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 91 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 88 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 89 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 87 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 86 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

Figure 86: Leiurus jordanensis Lourenço, Modrý et Amr, 2002, female. A. Carapace and tergites. B. Coxosternal area and sternites. Al-Tawil, Saudi Arabia.

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Figure 84 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

Figure 84: Leiurus abdullahbayrami Yağmur, Koç et Kunt, 2009, male. A. Carapace and tergites. B. Coxosternal area and sternites. Eski Sarkaya Village, Turkey.

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Figure 83 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

Figure 83: Leiurus abdullahbayrami Yağmur, Koç et Kunt, 2009, female. A. Carapace and tergites. B. Coxosternal area and sternites. Çaybasi Village, Turkey.

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Figure 97 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

Figure 97: Carapace and tergites I–III of Cicileiurus monticola Teruel, 2007, holotype female, Atlas Mountains, Morocco. Scale bar: 2 mm.

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Figure 82 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 90 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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.

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Figure 94 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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

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Figure 85 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

Figure 85: Leiurus jordanensis Lourenço, Modrý et Amr, 2002, male. A. Carapace and tergites. B. Coxosternal area and sternites. Al-Tawil, Saudi Arabia

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Figure 80 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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).

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Figure 78 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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

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Figure 81 in A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula

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).

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