Buthus (Buthacus) Birula, 1908: 139, 140; type species by original designation Androctonus (Leiurus) leptochelys Ehrenberg, 1829 [= Buthacus leptochelys (Ehrenberg, 1829)].

Buthacus: Simon, 1910: 74; Werner, 1934: 269; Kästner, 1941: 231; Vachon, 1948: 473–475, figs 229–232; Vachon, 1952: 178–180, figs. 229–232, 578; Bücherl, 1964: 57; Stahnke, 1972: 131; Lamoral and Reynders, 1975: 499; Vachon, 1979: 44; Levy and Amitai, 1980: 75, 76; Francke, 1985: 6, 15; Sissom, 1990: 101; El-Hennawy, 1992: 97, 101, 112–114; Nenilin and Fet, 1992: 15–17; Amr and El-Oran, 1994: 186–187, 189; Kovařík, 1995: 19; 1997a: 49; 1997b: 179; Braunwalder and Fet, 1998: 32–34; Fet et al., 1998: 615, 616; Kabakibi et al., 1999: 82, 88; Fet and Lowe, 2000: 81–86; ICZN, 2000: 7; Lourenço, 2001: 255–268, figs. 23, table 1; Kovařík, 2001: 80; Crucitti and Vignoli, 2002: 438–443, figs. 3–5, 16; Kovařík, 2002: 5; Fet et al., 2003: 1–6, 10, fig. 1, table 1; Kovařík, 2003: 135, 137, 155, table 1; Lourenço, 2003: 877, 880, fig. 10; Soleglad and Fet, 2003a: 5, 26, table 2; 2003b: 7, 13, 88, 149, 151, 152, figs. B-1, B-2, table 9; Kovařík and Whitman, 2004: 106; Lourenço, 2004a: 205–209, figs. 1–6; 2004b: 226–230, figs. 1–16, table 1; Kovařík, 2005: 1–10, figs. 1–8, table 1; Prendini and Wheeler, 2005: 481, table 10; Hendrixson, 2006: 33, 36, 46–56, 59, 91, 99, 100, 109, figs. 4–7, plates 3–5, table 1; Lourenço, 2006: 59–69, figs. 1–43, table 1; Lourenço and Qi, 2006a: 161–164, figs. 1–16, table 1; 2006b: 301–306; Dupré, 2007: 3, 4, 12, 16; Yağmur et al., 2008: 13–17, figs. 2, 4; Navidpour et al., 2008: 1–9, 28, 30, 35, figs. 3, 4, 6, 12, 56–59; Zourgui et al., 2008: 81, 84, 85, 89, fig. 4, plate 5, 7, table 2; Al-Asmari et al., 2009a: 612, 613, 617–619, 621, 622, 624, 626, figs. 8, 9, 12F, G, tables 1,TABLE 22; 2009b: 96, 100, 102, 104, 106, fig. 6K, L, tables 5–7; Lourenço and Leguin, 2009: 103–107, figs. 1–12, table 1; Zarei et al., 2009: 46–49, 51, fig. 2; Zambre and Lourenço, 2010: 115–118, figs. 1–10, table 1; Mirshamsi et al., 2011: 17, 24, 26, fig. 1a, table 1; Shehab et al., 2011: 333–340, figs. 2E, 3–5, tables 1, 2; Lourenço et al., 2012: 307, 309, 313–316, 336, 337, figs. 12–23, 89, 90; Al-Asmari et al., 2013: 2, 5–16, figs. 6–10, 12–13, table 1; Kovařík et al., 2013: 1, 3, 4, 18, figs. 6; Lourenço, 2013: 89–97, figs. 1–14, table 1; Navidpour et al., 2013: 1, 3–5, 21, figs. 2, 4; Amr, 2015: 182–185, 192, 197, 198, figs. 1, 2C, table 3; Caliskan, 2015: 327–332, 336, 337, 343, figs. 1, 3, 5B; Lourenço and Sadine, 2015: 55–59, figs. 1–18; table 1; Navidpour, 2015: 10, 11, table 1; Amr et al., 2015: 30–33, 1C, D; Kovařík et al., 2016: 1–17, figs. 1–63, table 1; Lourenço et al., 2016: 2–10, figs. 1, 4–16; Lourenço et al., 2017a: 31–40, figs. 1, 2, 5–17; 2017b: 18–28, figs. 1, 5–16, tables 1, 2; Saleh et al., 2017: 9, 13, table 2; Sharifinia et al., 2017: 232–236, table 2; Badry et al., 2018: 77, 83; Dehghani and Kassiri, 2018: S881, table 2; Kovařík, 2018: 1–3, 6–10, figs. 10–14, 19–22; Sadine et al., 2018: 51, 53–57, figs. 3C, D, 4, table 1; Al-Khazali and Yağmur, 2019: 85–87, fig. 2C; Alqahtani et al., 2019: 19, 21–25, fig. 2C; Bousmaha et al., 2019: 141–145, figs. 2, 3, table 1; Francke, 2019: 6, 20, 37; Koç et al., 2019: 108–112, figs. 1, 5–7, table 1; Lowe et al., 2019: 1–5, 11, 15, 22, 24, 25, 28, figs. 45, 46, 78–81; Alqahtani and Badry, 2020: 178–183, figs. 1, 3, tables 1, 2; Hussen and Ahmed, 2020: 6711–6713, 6715, 6716, 6718, 6719, 6722, 6723, figs. 1, 7, tables 1, 2; Obuid-Allah et al., 2020: 227, 229–237, figs. 2G, H, 3D, tables 1, 3–6; Alqahtani and Badry, 2021: 3, 4, 7, 12, fig. 4, table 1; Amr et al., 2021: 84–90, 95, 97, fig. 3A, B, tables 1–5, 9–11, 17; Mansouri et al., 2021: 763, 765, 767, table 2; Said et al., 2021: 17, 18, 20, 21, 25, 26, fig. 4, table 1.

Diagnosis: The genus Buthacus is most similar morphologically to other psammophilous buthid genera from the Palearctic deserts, notably Buthiscus, Liobuthus, Pectinibuthus, Plesiobuthus, Trypanothacus, and Vachoniolus, from which it differs as follows. Tibial spurs are present on legs III and IV in Buthacus and Trypanothacus, but may be reduced or absent on leg III; present only on leg IV in Buthiscus; and absent in Liobuthus, Pectinibuthus, Plesiobuthus, and Vachoniolus (Sissom, 1990; Levy et al., 1973; Fet et al., 2001; Lowe, 2010; Kovařík et al., 2016). The pedipalp chela movable finger bears 9–12 subrows of median denticles in Buthacus, compared with 7 in Liobuthus, 7–9 in Trypanothacus, 9 in Vachoniolus, 10 in Pectinibuthus, and 12 in Plesiobuthus (Sissom, 1990; Levy et al., 1973; Fet et al., 2001; Lowe, 2010; Kovařík et al., 2016). The telson vesicle is pyriform, with a long, curved aculeus in Buthacus, whereas the vesicle is more bulbous, with a shorter aculeus in Trypanothacus (Lowe et al., 2019).

Distribution: The species of Buthacus are widespread in the sandy deserts of the Palearctic region, occurring from the Atlantic coast of West Africa across the Sahara and the Sahel, and throughout the Middle East to India. In Africa, the genus has been recorded from the following countries and territories: Algeria, Chad, Egypt, Libya, Mauritania, Morocco, Niger, Nigeria, Senegal, Sudan, Tunisia, and Western Sahara. The presence of Buthacus in Burkina Faso, Eritrea, and Mali, while presently unconfirmed, seems likely. In Asia, the genus has been recorded from the following countries: Afghanistan, Bahrain, India, Iran, Iraq, Israel, Jordan, Kuwait, Lebanon, Oman, Pakistan, Qatar, Saudi Arabia, Syria, Turkey, the United Arab Emirates. Buthacus probably also occurs in Yemen and the Gaza Strip.

Ecology: All species of Buthacus inhabit sandy substrates (fig. 1). However, the stability of the substrate, a function of its hardness, texture, and the density of rock and vegetation vary, from shifting sand dunes, through semiconsolidated dunes, flats, or dry watercourses, to consolidated sand, gravel, and loess plains. Closely related species may inhabit substrata of very different hardness and texture. For example, B. arava, sp. nov., inhabits semiconsolidated to shifting sand dunes, whereas the closely related B. tadmorensis inhabits loose to hard, compacted sandy plains and wadis, often with stones. The suite of ultrapsammophilous to semipsammophilous adaptations these scorpions possess for locomotion and burrowing on sandy substrata include elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, rows of elongated macrosetae (“sand combs”) along the retrolateral margins of the tibae and the pro- and retrolateral margins of the basitarsi of legs I–III, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and elongated ungues, often unequal in length on legs I and II (Prendini, 2001). Additionally, many species exhibit loss or reduction of granulation and carination on the tegument, and pale coloration, often associated with a loss or reduction of infuscation.

Included Species: The genus Buthacus currently includes 30 species and one subspecies: Buthacus agarwali Zambre and Lourenço, 2010; Buthacus ahaggar Lourenço et al., 2017; Buthacus amitaii, sp. nov.; Buthacus arava, sp. nov.; Buthacus arenicola Simon, 1885; Buthacus a. maroccanus Lourenço, 2006; Buthacus armasi Lourenço, 2013, stat. rev.; Buthacus birulai Lourenço, 2006; Buthacus clevai Lourenço, 2001; Buthacus elmenia Lourenço et al., 2017; Buthacus foleyi Vachon, 1948; Buthacus frontalis Werner, 1936; Buthacus fuscata Pallary, 1929, stat. nov. et stat. rev.; Buthacus golovatchi Lourenço et al., 2012; Buthacus leptochelys Ehrenberg, 1829; Buthacus levyi, sp. nov.; Buthacus nigerianus Lourenço and Qi, 2006; Buthacus nigroaculeatus Levy et al., 1973; Buthacus nitzani Levy et al., 1973, stat. nov.; Buthacus occidentalis Vachon, 1953; Buthacus pakistanensis Lourenço and Qi, 2006; Buthacus samiae Lourenço and Sadine, 2015; Buthacus spatzi (Birula, 1911), stat. rev.; Buthacus spinatus Lourenço et al., 2016; Buthacus stockmanni Kovařík et al., 2016; Buthacus striffleri Lourenço, 2004; Buthacus tadmorensis Simon, 1892, stat. rev.; Buthacus villiersi Vachon, 1949; Buthacus williamsi Lourenço and Leguin, 2009; Buthacus yotvatensis Levy et al., 1973, stat. rev.; Buthacus ziegleri Lourenço, 2000.

Remarks: The original description of Buthacus frontalis Werner, 1936, was based on a dry specimen that has never been reexamined According to Kovařík (2003: 152) the “holotype (a female from Asmara, Eritrea) was destroyed during the air raid on Hamburg in 1943… [the] original description does not contain diagnostic characters, and no additional specimens are known [therefore] this name has to be regarded as dubious.” Levy et al. (1973) and Levy and Amitai (1980) also doubted the assignment of this species to the genus Buthacus.

Four species originally described in the genus Buthacus were subsequently transferred to other genera (Vachon, 1979; Kovařík et al., 2013; Kovařík, 2018; Lowe et al., 2019). Buthacus minipectenibus Levy et al., 1973, was transferred by Vachon (1979) to the genus Vachoniolus Levy et al., 1973, becoming Vachoniolus minipectenibus (Levy et al., 1973), and later synonymized with Vachoniolus globimanus Levy et al., 1973, by Hendrixson (2006). Buthacus calviceps, from Somaliland (Pocock, 1900), was transferred to the genus Gint Kovařík et al., 2013, becoming Gint calviceps (Pocock, 1900). Buthacus buettikeri, from Saudi Arabia (Hendrixson, 2006), was transferred to the genus Trypanothacus Lowe et al., 2019, becoming Trypanothacus buettikeri (Hendrixson, 2006). Buthacus maliensis Lourenço and Qi, 2007, from Mali, was synonymized with Androctonus aleksandrplotkini Lourenço and Qi, 2007 by Kovařík (2018).

Additionally, three other species, B. argarwali from India (Zambre and Lourenço, 2010), B. pakistanensis from Pakistan (Lourenço and Qi, 2006a), and B. williamsi from the United Arab Emirates (Lourenço and Leguin, 2009), appear to have been erroneously assigned to Buthacus based on the carapacial carinae, pedipalp shape, and counts of median denticle subrows on the pedipalp chela fingers, among other characters (see, e.g., Kovařík et al., 2016).

Buthacus leptochelys leptochelys: Levy and Amitai, 1980: 76, 77, 79–83, figs. 70–74, map 6; Kabakibi et al., 1999: 82, 88 (misidentification); Fet and Lowe, 2000: 84; Lourenço, 2006: 62.

Buthacus leptochelys: El-Hennawy, 1992: 101, 112, 113 (misidentification); Lourenço, 2006: 59–69, figs. 1, 4, table 1.

Type Material: ISRAEL: Mehoz HaDarom (Southern District): Holotype ♂ (HUJ INVSC 3480 ex AMNH), Mamshit [Kurnub], road to Negev Camel Ranch 31°01′59.9″N 35°04′25″E, 477 m, 28.viii.2011, L. Prendini and T.L. Bird. Paratypes: Mamshit [Kurnub, 31°01′N 35°03′E], 475 m, 14–20.vi.1971, P. Amitai, 3 ♂ (HUJ INVSC 2283–2284, 2294), 2 ♀ (HUJ INVSC 2278, 2292); Mamshit, road to Negev Camel Ranch 31°01′59.9″N 35°04′25″E, 477 m, 28. viii.2011, L. Prendini and T.L. Bird, 65 ♂, 44 ♀(AMNH), 2 juv. ♂, 2 juv. ♀ (AMCC [LP 11176]); Mamshit, Negev Camel Ranch, wadi behind camp, 31°01′46.5″N 35°04′39.2″E, 471 m, 28. viii.2011, L. Prendini and T.L. Bird, 1 ♂, 11 ♀(AMNH), 2 juv. ♂ (AMCC [LP 11173]), 29. viii.2013, L. Prendini, I. Tesler and A. Novikova, 13 ♂, 23 ♀ (AMNH); Mamshit, W of Negev Camel Ranch car park, 31°01′50.7″N 35°04′35.2″E, 475 m, 5.viii.2016, E. Gefen and S. Cain, 3 ♂ (AMCC [LP 15046] ex HUJ INVSC 3202, HUJ INVSC 3203–3204), 2 ♀ (AMNH ex HUJ INVSC 3200, HUJ INVSC 3201); Nahal HaTira, Kholot Tsivoniyim campsite, 30°57′07.9″N 35°01′34.9″E, 304 m, 16.iv.2018, Y. Zvik and S. Cain, 1 ♂ (HUJ INVSC 3463), 3 ♀(HUJ INVSC 3462, 3464 – 3465), 3 juv. ♂ (HUJ INVSC 3466, 3468, 3470), 2 juv. ♀ (HUJ INVSC 3467, 3469); Naal Yamin, SW of Dimona factory, 30°57′01.8″N 35°04′51.7″E, 330 m, 16. iv.2017, Y. Zvik, 2 ♀ (AMCC [LP 15047] ex HUJ INVSC 3207, HUJ INVSC 3205), 1 subad. ♀(HUJ INVSC 3206), 16.iv.2018, Y. Zvik and S. Cain, 1 ♀ (HUJ INVSC 3471), 1 subad. ♀ (HUJ INVSC 3472).

Diagnosis: Buthacus amitaii is most closely related to B. nitzani, also occurring in Israel, and B. leptochelys, occurring in Egypt. Buthacus amitaii differs from B. nitzani as follows. The metasomal segments of the female are shorter and broader in B. amitaii (fig. 28D), with segment IV width:length, 42.8% (40.2%–44.9%, n = 5; table 3), than in B. nitzani (fig. 28C), with segment IV width:length, 38.4% (34%–43.7%, n = 10; table 13). The ventrosubmedian and ventrolateral carinae are more developed on the metasomal segments, especially on segments II and III, in B. amitaii than B. nitzani (figs. 26C, D, 28C, D). The posterior processes of the ventrolateral carinae of metasomal segment V are broad and lobate in B. amitaii, but narrow and spiniform in B. nitzani. The median lateral carinae are distnct on metasomal segments I–III in B. amitaii, but obsolete in B. nitzani. The ventral intercarinal surfaces of metasomal segment V are more coarsely and densely granular in B. amitaii than B. nitzani. The retrolateral accessory denticles on the movable finger of the pedipalp chela are often absent, with variable, usually lower counts (sinistral/dextral: 0–9/2–10; n = 9; table 2; fig. 21A) in B. amitaii, compared to B. nitzani, in which the retrolateral accessory denticles are always present, with constant, usually higher counts (6–9/6–9; n = 16; table 2; fig. 21G).

Buthacus amitaii differs from B. leptochelys as follows. Buthacus amitaii is smaller, measuring 51.7 mm (48–58.8 mm, n = 10; table 3) in total length, than B. leptochelys, measuring 65.8 mm (63.1–68 mm, n = 5; table 4). The pedipalp chela of the adult male is shorter and broader (fig. 32A, B), with chela manus width:chela length, 27.1% (25.1%–28.9%, n = 5; table 3) and chela manus height:length, 69.7% (58.9%–75.5%, n = 5), in B. amitaii than in B. leptochelys (fig. 41A, B), with chela manus width:chela length, 24.2% (table 4), and chela manus height:length, 58.9%. The proximal dentate margins of the pedipalp chela fixed and movable fingers are more deeply emarginate, such that a more pronounced gap is evident proximally when the fingers are closed, in B. amitaii (fig. 32B) than in B. leptochelys (fig. 41B). The pectinal tooth counts are lower in B. amitaii, i.e., 30/31 (28–32/28–33, n = 5; table 2) (♂) and 23/23 (21–26/21–26, n = 9) (♀), than in B. leptochelys, i.e., 35/34 (32–39/32–38, n = 6; table 5) (♂) and 29/29 (27–31/28–31, n = 10) (♀). The telson vesicle dorsal surface is concave and the ventral surface convex and angular in lateral profile, in B. amitaii (fig. 29H), whereas the dorsal surface is flat and the ventral surface convex and rounded in lateral profile, in B. leptochelys (fig. 29D).

Buthacus amitaii differs from B. levyi, another species occurring in Israel with which it may be confused, as follows. The pedipalp chela of the adult male is shorter and broader in B. amitaii (fig. 32A, B), with chela manus length:movable finger length, 81.8% (72.7%–88.3%, n = 5; table 3) and chela manus width:chela length, 27.1% (25.1%–28.9%, n = 5), than in B. levyi (fig. 44A, B), with chela manus length:movable finger length, 62.8% (57.4%–67.3%, n = 8; table 10) and chela manus width:chela length, 19.4% (17.1%–21.3%, n = 8). The proximal dentate margins of the pedipalp chela fixed and movable fingers are emarginate, such that a gap is evident proximally when the fingers are closed, in B. amitaii (fig. 32B), but sublinear, such that no gap is evident proximally when the fingers are closed, in B. levyi (fig. 44B). The counts of retrolateral accessory denticles on the movable finger of the pedipalp chela are usually higher in B. amitaii (sinistral/dextral: 0–9/2–10; n = 9; table 2; fig. 21A) than in B. levyi (0–5/0–3; n = 13; table 5; fig. 21E, F). The posterior processes on the ventrolateral carinae of segment V are larger and more variable in size in B. amitaii (figs. 26D, 28D) than in B. levyi (figs. 26A, 28A). The telson dorsal surface is concave and the ventral surface convex and angular in lateral profile, in B. amitaii (fig. 29H), whereas the dorsal surface is flat and the ventral surface convex in lateral profile, in B. levyi (fig. 29B).

Etymology: The species name is a patronym, honoring Pinchas Amitai, one of the pioneers of arachnology in Israel, whose publications, some coauthored with Gershom Levy (e.g., Levy and Amitai, 1980), constitute a milestone in understanding the scorpions and other arachnids of the Levant.

Description: The following description is based on the type material (see table 2 for counts and table 3 for measurements).

Total length: Medium-sized scorpions, 53.9 mm (48–58.8 mm, n = 5) (♂) or 49.5 mm (48.5–50.8 mm, n = 5) (♀).

Color: Uniformly yellowish olive except as follows. Carapace interocular surface, pedipalp femur, and patella, legs, and metasomal segments orange. Pectines pale yellow. Legs articulation points brown.

Carapace: Carapace shape trapezoidal (fig. 17E, F), anterior width:posterior width, 52.1% (45.8%–54%, n = 10), length:posterior width, 88.7% (83.6%–94.6%, n = 10). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with two or, usually, three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 42.3% (40.6%–43.5%, n = 5) (♂) or 42.7% (40.5%–43.6%, n = 5) (♀). Superciliary and central median carinae distinct, costate-granular, strongly to weakly connected (♂) or weakly connected to disconnected (♀). Anteromedian sulcus distinct, shallow; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulci shallow, wide, curved. Carapace intercarinal surfaces finely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces densely setose. Fixed finger dorsal and ventral surfaces densely setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger dorsal surface smooth and glabrous; ventral surface densely setose; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina absent, represented by fewer than 10 macrosetae; intercarinal surfaces smooth (fig. 31A, B). Patella prolateral median and ventral prolateral carinae discontinuous, each comprising a few spiniform granules; other carinae absent; intercarinal surfaces smooth (fig. 31C–E). Chela short and broad (♂), manus width:length, 62.6% (53.4%–68.3%, n = 5), manus height:length, 69.7% (58.9%–75.5%, n = 5), and manus length:movable finger length, 81.8% (72.7%–88.3%, n = 5) or short and slender (♀), manus width:length, 54.4% (49.4%–58.9%, n = 5), manus height:length, 61.2% (55.1%–64.6%, n = 5), and manus length:movable finger length, 58.1% (54.2%–61.5%, n = 5). Chela manus acarinate; intercarinal surfaces smooth and glabrous (fig. 32). Fixed and movable fingers each with 8–10 (n = 9) oblique median denticle subrows; movable finger with 0–10 (n = 9) retrolateral accessory denticles (fig. 21A); proximal dentate margins of fingers emarginate (fig. 32B), such that gap present proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae obsolete; intercarinal surfaces smooth. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 6:9:11:4; basitarsi, 9:12:15:10; telotarsi, 4:6:5:6 (n = 1). Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Telotarsal ungues long, approximately equal to telotarsus length, unequal on legs I and II, subequal to equal on III and IV (fig. 22H).

Genital operculum: Genital opercula suboval, completely divided longitudinally, with overlapping, rounded margins (♂) or partially fused longitudinally (♀) (fig. 20C, D). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 8–10 (n = 2) (♂) or 7–9 (n = 3) (♀) median lamellae (fig. 20C, D). Fulcra present. Pectinal teeth along most of length, dentate margin length:pecten length, 98.3% (94.4%–106.7%, n = 5) (♂) or 95% (91.3%–98.3%, n = 5) (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 30/31 (28–32/28–33, n = 5) (♂) or 23/23 (21–26/21–26, n = 9) (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 57.6% (55.1%–61.3%, n = 5) (♂) or 68.9% (55.2%–106.2%, n = 5) (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly finely granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae absent on I, granular, vestigial, restricted to posterior third of II–IV and posterior half of V and VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–IV and posterior third of V and VI. Tergite VII, dorsomedian carina absent to granular, vestigial, restricted to anterior half; dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII smooth and glabrous; III–VI acarinate, VII, ventrolateral carinae vestigial, granular; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 24D, 26D, 28D), segment I shortest, length I:II, 87.5% (84.2%–89.5%, n = 5) (♂) or 86.7% (80.4%–89.8%, n = 5) (♀); segments II–IV similar, length II:III, 96.4% (95.1%–97.1%, n = 5) (♂) or 97.3% (95%–100.2%, n = 5) (♀), length III:IV, 97.4% (93.2%–103.1%, n = 5) (♂) or 99.3% (96.4%–103.9%, n = 5) (♀); segment V longest, length IV:V, 82.2% (80.8%–83.4%, n = 5) (♂) or 77.5% (73.3%–84.8%, n = 5) (♀); width:length segment I, 69.1% (61.8%–72.7%, n = 5) (♂) or 69.6% (64%–77.4%, n = 5) (♀), II, 58.8% (57.8%–61.2%, n = 5) (♂) or 56.6% (52.1%–59.6%, n = 5) (♀), III, 53.8% (52.2%–55.1%, n = 5) (♂) or 52.5% (49.4%–55.7%, n = 5) (♀), IV, 42.8% (37.2%–47.3%, n = 5) (♂) or 42.8% (40.2%–44.9%, n = 5) (♀), V, 35.8% (33.9%–39.5%, n = 5) (♂) or 36.1% (33.2%–38.5%, n = 5) (♀). Dorsosubmedian carinae distinct, granular on segments I and II, obsolete on III, absent on IV and V; dorsolateral carinae distinct, granular on segments I and II, obsolete on III and IV, absent on V; dorsosubmedian and dorsolateral carinae sparsely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 1/0 (0/0–3/1, n = 5):2/3 (2/2–3/4):3/3 (2/3–4/4):3/3 (1/2–4/4):0/0 (0/0–1/0); dorsolateral carinae, 2/2 (0/1–3/3, n = 5):3/3 (2/1–3/4):4/3 (3/1–5/5):3/2 (1/0–5/4):5/6 (2/4–7/8). Median lateral carinae distinct, granular, extending entire length of segment I, restricted to posterior half of II and posterior quarter of III, absent on IV and V. Ventrolateral carinae distinct, costate, becoming granular posteriorly on segment I; costate-granular along entire length, with granules becoming progressively larger and spiniform posteriorly, on II and III; costate-granular along entire length of IV; distinct, serrate, comprising spiniform granules and lobate processes posteriorly, on V. Ventrosubmedian carinae distinct, costate on segment I; costate-granular along entire length, with granules becoming progressively larger and spiniform posteriorly, on II and III; costate-granular along entire length of IV; granular, restricted to anterior three-quarters of V. Ventromedian carina granular, distinct along entire length of segment V. Dorsal and lateral intercarinal surfaces smooth on segments I–V; ventral intercarinal surfaces smooth on I and II, finely granular on III and IV, finely and densely granular across entire surface of V.

Telson: Telson vesicle width:metasomal segment V width, 80.5% (76.6%–82%, n = 5) (♂) or 78.3% (74.3%–80.5%, n = 5) (♀). Vesicle globose, dorsal surface concave, ventral surface convex and angular; vesicle height:length, 69.2% (62.6%–74.5%, n = 5) (♂) or 66.2% (61.5%–69.9%, n = 5) (♀); dorsal and ventral surfaces smooth and glabrous; lateral and ventral surfaces sparsely setose, with 26 (23–29, n = 2) (♂) or 32 (29–39, n = 3) (♀) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 52.1% (49.6%–55.5%, n = 5) (♂) or 55.2% (52.1%–60.5%, n = 5) (♀).

Sexual dimorphism: Adult males and females differ as follows. The pedipalp chela manus of the male is markedly broader and deeper with proportionally shorter fingers than that of the female (fig. 32), as indicated by the higher ratios of chela manus width:length, manus height:length, and manus length:movable finger length, respectively 62.6%, 69.7%, and 81.8% in the male compared with 54.4%, 61.2%, and 58.1% in the female. The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female (fig. 20C, D), and genital papillae are present in the male but absent in the female. The pectinal tooth count is higher in the male (28–33) than in the female (22–26). The spiniform granules of the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and the ventrolateral carinae of segment V are less prominent in the male than in the female.

Distribution: Buthacus amitaii appears to be endemic to the inland sandy soils and loess of the Yamin Plain and Mamshit area in the northern Negev, Israel (fig. 7). The known records are located at elevations ranging from 304 m to 477 m.

Buthacus amitaii is allopatric with the closely related B. leptochelys, distributed from the western Sinai Peninsula across Egypt, and southward along the coast of the Red Sea to Sudan (fig. 6), and B. nitzani, distributed from the Sorek (Rubin) River, throughout the sand dunes of the southern coastal plain of Israel, to the Haluza dunes of the interior and probably the adjacent Sinai Peninsula of Egypt (fig. 7).

Ecology: The types of B. amitaii were collected at night with UV light detection on flat to gently sloping loess plains, comprising compacted sandy loam with soft hummock dunes in places. The substratum was noticeably more compact compared with the habitats in which the closely related B. leptochelys and B. nitzani were collected. Most specimens were found walking, but some were sitting on the ground or on bushes. The habitat and habitus, notably the pale coloration, smooth tegument, obsolescence of some pedipalpal and metasomal carinae, elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins of the tibae and basitarsi of legs I–III, and elongated telotarsal ungues, are consistent with the psammophilous ecomorphotype (Prendini, 2001).

The types of B. amitaii were collected in sympatry with five other buthids, Androctonus bicolor Ehrenberg, 1828, Androctonus amoreuxi (Audouin, 1826), Buthus israelis Shulov and Amitai, 1959, Leiurus quinquestriatus (Ehrenberg, 1828), Orthochirus scrobiculosus (Grube, 1873), and the scorpionid, Scorpio palmatus (Ehrenberg, 1829). Leiurus quinquestriatus usually occurs in rocky habitats, however.

Remarks: According to Levy and Amitai (1980), the population of B. leptochelys inhabiting the inland sandy soils near Mamshit, in the northern Negev, belonged to the nominotypical subspecies, otherwise occurring along the southern coast of Israel and in the Sinai Peninsula. Based on evidence gathered during the present investigation, including multivariate analysis of morphometrics (fig. 15) and multilocus molecular phylogenetics (fig. 14), this population represents a distinct species, described here as Buthacus amitaii, sp. nov.

Figures 1E, 3A, 4, 15, 16A, B, 19A, B, 21B, 22C, 23A, 25A, 27A, 29C, 33, 34, 35; tables 1, 8, 9

Buthacus nigroaculeatus: Amr et al., 2021: 86, fig. 3B (misidentification).

Type Material: Holotype ♂ (HUJ INVSC 3672), ISRAEL: Mehoz HaDarom (Southern District): Lotan, SE of Lotan Plantations, 30°00′10.4″N 35°05′44.1″E, 128 m, 13.ix.2018, S. Cain. Paratypes: ISRAEL: Mehoz HaDarom (Southern District): near Lotan [29°59′N 35°05′E], 123 m, 29.ix.2003, ♂ (HUJ INVSC 2560), U. Shanas; Lotan, SE of Lotan Plantations, 30°00′10.4″N 35°05′44.1″E, 128 m, 15.ix.2017, Y. Olek and S. Cain, 8 ♂ (HUJ INVSC 3282, 3284–3287, 3289–3291), 1 ♂ (AMCC [LP 15042] ex HUJ INVSC 3283) 1 ♂ (AMCC [LP 15043] ex HUJ INVSC 3288), 1 ♀ (HUJ INVSC3381), 27.iii.2018, Y. Olek and S. Cain; Nahal Sha'alav, 4 km S of Kibbutz Yahel, 30°02′40.6″N 35°06′33.7″E, 153 m, 15.ix.2017, Y. Olek and S. Cain, 1 ♂ (HUJ INVSC 3299); Qetura [29°58′N 35°03′E], 100 m, 8.x.1974, M. Israel, 1 ♂ (HUJ INVSC 2329); S of Yahel, sand dunes, 30°02′N 35°06′E, 149 m, 14.x.2004, U. Shanas, 1 ♂ (HUJ INVSC 2587); Yotvata, E side of circle field, 29°53′21.1″N 35°04′34.8″E, 72 m, 14.ix.2017, Y. Olek and S. Cain, 1 ♂ (AMCC [LP 15045] ex HUJ INVSC 3292), 1 ♂ (HUJ INVSC 3293), 29°53′36.1″N 35°04′40.6″E, 72 m, 14.ix.2017, Y. Olek and S. Cain, 3 ♂ (HUJ INVSC 3294–3296), 29°53′45.1″N 35°04′37.5″E, 72 m, 14.ix.2017, Y. Olek and S. Cain, 1 ♂ (AMCC [LP 15044] ex HUJ INVSC 3297), 1 ♂ (HUJ INVSC 3298). JORDAN: Aqaba Governorate: E of Qetura, 29°57′N 35°05′E, 98 m, 11.x.2004, U. Shanas, 2 ♂ (HUJ INVSC 2594–2595); Rahma, 29°56′N 35°06′E, 95 m, 27.x.2003, U. Shanas, 1 ♂ (HUJ INVSC 2569); Wadi Aheimir [Hemer], 30°02′N 35°10′E, 212 m, 16.x.2003, U. Shanas, 2 ♂ (HUJ INVSC 2559, 2562); Wadi Quseib, NE of Yahel, 30°06′N 35°10′E, 189 m, 23.x.2003, U. Shanas, 1 ♂ (HUJ INVSC 2563); E of Yotvata, 29°46′N 35°04′E, 156 m, 24.x.2003, U. Shanas, 1 subad. ♀(HUJ INVSC 2561); NE of Yotvata, sand dunes, 29°54′N 35°04′E, 76 m, 12.x.2004, U. Shanas, 1 ♂ (HUJ INVSC 2597). Karak Governorate: Wadi Khanzeerah (Khanzairh), 30°53′39.9″N 35°25′38.2″E, -294 m, 9.ix.2013, L. Prendini, Z. Amr and L. Al Azam, 1 ♂ (AMNH), 1 ♂ (JUST), 1 juv. ♂, 2 juv. ♀ (AMCC [LP 12198]).

Diagnosis: Buthacus arava differs from the closely related B. tadmorensis and B. yotvatensis, occurring in Israel and neighboring states, as follows. The pedipalp chela of B. arava is shorter and broader, especially in the adult male (fig. 35A, B), with chela manus width:chela length, 29.8% (27.2%–32.8%, n = 7; table 8) (♂), chela manus length:movable finger length, 90.4% (83.5%–95%, n = 7) or 60.8% (♀), and chela manus width:length, 63.3% (57.3%–74.5%, n = 7) or 61.6% (♀), than that of B. tadmorensis (fig. 50A, B), with chela manus width:chela length, 22.3% (♂) (table 4), chela manus length:movable finger length, 64.2% (♂) or 55% (51.9%–59.8%, n = 4) (♀), and chela manus width:length, 58.7% (♂) or 59.1% (53.8%–65.1%, n = 4) (♀), and B. yotvatensis (fig. 53A, B), with chela manus width:chela length, 17.4% (16.5%–18.8%, n = 8; tables 6, 7) (♂), chela manus length:movable finger length, 55.2% (51.7%–58.3%, n = 8) (♂) or 47.1% (45.5%–48%, n = 6) (♀), and chela manus width:length, 50.5% (48.1%–54.8%, n = 8) (♂) or 56.9% (51.3%–61.5%, n = 6) (♀). The ungues of the leg telotarsi are longer (approximately equal to the length of the telotarsus) and unequal on legs I and II in B. arava (fig. 22C) but shorter (approximately two-thirds the length of the telotarsus) and equal on legs I and II in B. tadmorensis and B. yotvatensis (fig. 22E, G). The pectinal tooth counts are lower in B. arava, i.e., 20/20 (18–21/18–22, n = 7; table 9) (♂) and 12/13 (♀), than in B. tadmorensis, i.e., 30/32 (♂) (table 9) and 24/25 (23–26/23–26, n = 3) (♀), and B. yotvatensis, i.e., 34/35 (32–37/32–37, n = 8; table 9) (♂) and 26/26 (25–28/26–28, n = 6) (♀). The metasomal segments and telson are sparsely setose in B. arava (figs. 23A, 25A, 29C), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 0/0 (0–1/0–1, n = 4):2/3 (2–3/2–5):4/3 (2–5/2–6):3/3 (2–3/1–5):0/0 (0–0/0–0), dorsolateral carinae, 1/1 (0–2/0–2):2/2 (1–3/0–3):3/2 (2–4/1–3):3/3 (2–4/2–5):5/6 (3–8/5–7), and telson, 28 (27–29, n = 4), but densely setose in B. tadmorensis (figs. 23B, 25B, 29E), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 10/11 (8–12/10–12, n = 2):19/23 (18–19/21–24):23/23 (23–23/21–25):19/20 (17–20/19–21):9/9 (7–11/7–11), dorsolateral carinae, 6/6 (4–8/5–6):11/13 (8–13/11–15):9/11 (8–9/10–11):9/9 (8–9/6–12):9/9 (8–10/8–10), and telson, 100 (80–120, n = 2), and B. yotvatensis (figs. 23C, 25C, 29G), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 16/15 (14–20/11–21, n = 5):25/26 (20–29/18–30):26/25 (23–31/19–29):30/27 (24–38/23–31):12/11 (6–18/7–15), dorsolateral carinae, 9/8 (7–13/5–13):20/20 (19–23/16–21):22/21 (16–28/18–24):15/14 (11–20/12–17):11/13 (10–13/11–14), and telson, 102 (88–127, n = 5). Additionally, the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and ventrolateral carinae of segment V are less developed in B. arava than in B. tadmorensis (fig. 25A, B), with narrow, spiniform processes posteriorly in B. arava but broad, lobate processes posteriorly in B. tadmorensis.

Etymology: The specific epithet is a noun in apposition, taken from the Arava Valley, to which the species appears to be endemic.

Description: The following description is based on the type material (see table 8 for measurements and table 9 for counts).

Total length: Large scorpions, 70.4 mm (61.7–75 mm, n = 6) (♂) or 62.3 mm (♀).

Color: Uniformly pale yellow to yellowish-olive, except as follows. Mesosomal tergites, sternite VII and metasomal segments I–V, each with narrow brown stripe posteriorly. Pectines pale yellow. Leg articulation points brown.

Carapace: Carapace shape trapezoidal (fig. 16A, B), anterior width:posterior width, 50.6% (44.3%–59.4%, n = 8), length:posterior width, 90.6% (84.6%–96.7%, n = 8). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with two or, usually, three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 43.1% (41%–44.7%, n = 7) (♂) or 45.2% (♀). Superciliary and central median carinae distinct, costate-granular, weakly connected to disconnected (♂) or smooth and obsolete (♀). Anteromedian sulcus absent or shallow; posteromedian sulcus shallow, narrow anteriorly, wide posteriorly; posterolateral sulci shallow, wide, curved. Carapace intercarinal surfaces finely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin finely granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces densely setose. Fixed finger dorsal and ventral surfaces densely setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger dorsal surface smooth and glabrous; ventral surface densely setose; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae vestigial, each comprising a few spiniform granules; retrolateral dorsosubmedian carina absent, represented by fewer than 10 macrosetae; dorsal and ventral intercarinal surfaces finely granular; lateral intercarinal surfaces smooth (fig. 34A, B). Patella dorsal prolateral carina obsolete; prolateral median carina partial, costate-granular; ventral prolateral carinae complete, costate-granular; other carinae absent; dorsal, ventral and retrolateral intercarinal surfaces smooth; prolateral intercarinal surfaces finely granular (fig. 34C–E). Chela globose, short and broad (♂), manus width:length, 63.3% (57.3%–74.5%, n = 7), manus height:length, 69.1% (60%–84%, n = 7), and manus length:movable finger length, 90.4% (83.5%–95%, n = 7) or short and narrow (♀), manus width:length, 61.6%, manus height:length, 69.2%, and manus length:movable finger length, 60.8%. Chela manus acarinate; intercarinal surfaces smooth and glabrous (fig. 35). Fixed and movable fingers respectively with 8 or 9 (n = 8) and 6–10 (n = 8) oblique median denticle subrows; movable finger with 6–9 (n = 8) retrolateral accessory denticles (fig. 21B); proximal dentate margins of fingers sublinear (fig. 35B), such that no gap evident proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae obsolete; intercarinal surfaces smooth. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 8:9:11:4; basitarsi, 10:13:20:7; telotarsi, 4:5:6:6 (n = 1). Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on legs III and IV. Telotarsal ungues long, approximately equal to telotarsus length, unequal on legs I and II, subequal to equal on III and IV (fig. 22C).

Genital operculum: Genital opercula suboval, completely divided longitudinally, with overlapping, rounded margins (♂) or partially fused longitudinally (♀) (fig. 19A, B). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 7–8 (n = 3) (♂) or 6 (♀) median lamellae (fig. 19A, B). Fulcra present. Pectinal teeth along part of length, dentate margin length:pecten length, 88.1% (85.2%–95%, n = 7) (♂) or 67.1% (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 20/20 (18–21/18–22, n = 7) (♂) or 12/13 (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 56.4% (52.6%–59%, n = 7) (♂) or 47.2% (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly finely granular, becoming more coarsely and densely granular posteriorly, VII more coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae absent on I and II, granular, vestigial, restricted to posterior third of III–VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–VI. Tergite VII, dorsomedian carina absent; dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII acarinate, smooth and glabrous; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 23A, 25A, 27A), segment I shortest, length I:II, 88.5% (86.3%–89.9%, n = 7) (♂) or 90.3% (♀); segments II–IV similar, length II:III, 95.8% (93.7%–98.6%, n = 7) (♂) or 100.1% (♀), length III:IV, 95.4% (94.7%–96.7%, n = 7) (♂) or 92.3% (♀); segment V longest, length IV:V, 86.4 (82.4%–92.8%, n = 7) (♂) or 80.5% (♀); width:length segment I, 64.5% (62%–66.4%, n = 7) (♂) or 61% (♀), II, 56.3% (52.3%–66.4%, n = 7) (♂) or 55.4% (♀), III, 50.5% (47.1%–52.7%, n = 7) (♂) or 54.7% (♀), IV, 43.8% (40.6%–46.4%, n = 7) (♂) or 46.9% (♀), V, 35% (31.9%–38.3%, n = 7) (♂) or 37.1% (♀). Dorsosubmedian and dorsolateral carinae distinct, granular on segments I and III, obsolete on IV, absent on V; dorsosubmedian and dorsolateral carinae sparsely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 0/0 (0/0–1/1, n = 4):2/3 (2/2–3/5):4/3 (2/2–5/6):3/3 (2/1–3/5):0/0 (0/0–0/0), dorsolateral carinae, 1/1 (0/0–2/2):2/2 (1/0–3/3):3/2 (2/1–4/3):3/3 (2/2–4/5):5/6 (3/5–8/7). Median lateral carinae distinct, granular, restricted to posterior two-thirds of segment I, posterior half of II and posterior third of III, absent on IV and V. Ventrolateral carinae distinct, costate-granular on segment I; costate-granular, with granules becoming progressively larger and subspiniform posteriorly, on II and III; costate-granular on IV; serrate, comprising spiniform granules of variable size, becoming more prominent posteriorly, on V. Ventrosubmedian carinae distinct, costate-granular on segment I; costate-granular, with granules becoming progressively larger and subspiniform posteriorly, on II and III; costate-granular on IV; granular, restricted to anterior two-thirds of V. Ventromedian carina granular, distinct along entire length of segment V. Dorsal intercarinal surfaces smooth on I–V; lateral intercarinal surfaces smooth on I–III, smooth or very finely granular on segment IV, finely granular on V; ventral intercarinal surfaces finely granular on I–IV, coarsely granular in anterior two-thirds of V.

Telson: Telson vesicle width:metasomal segment V width, 69.4% (67.7%–71.5%, n = 7) (♂) or 69.7% (♀). Vesicle globose, dorsal surface flat, ventral surface convex and rounded; vesicle height:length, 63.2% (56%–68.2%, n = 7) (♂) or 59.9% (♀); dorsal surface smooth, ventral surface granular anteriorly; lateral and ventral surfaces sparsely setose, with 27 (27–29, n = 4) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 50.5% (44.9%–55%, n = 7) (♂) or 53% (♀).

Sexual dimorphism: Adult males and females differ as follows. The carapacial superciliary and central median carinae are distinct and costate-granular in the male, but obsolete and smooth in the female. The carinae of the pedipalp femur and patella are more coarsely granular in the male than in the female. The pedipalp chela manus of the male is markedly broader and deeper with proportionally shorter fingers than that of the female (fig. 35), as indicated by the higher ratios of chela manus width:length and chela manus length:movable finger length, respectively 63.3% and 90.4% in the male compared with 61.6% and 60.8%, in the female. The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female (fig. 19A, B), and genital papillae are present in the male but absent in the female. The pectinal tooth count is higher in the male (18–22) than in the female (12–14). The mesosoma of the male is relatively narrower than that of the female, as indicated by the higher sternite VII length:width ratio in the male (71.1%) compared to the female (67.8%). The spiniform granules of the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and the ventrolateral carinae of segment V are less prominent in the male than in the female.

Distribution: Buthacus arava appears to be endemic to the Arava Valley, straddling the southern border between Israel and Jordan (fig. 4). The species is presently known from only two rather restricted areas in the valley. Most specimens were collected in the central sand dunes in the southern part of the valley, extending from northeast of Yahel in the north to east of Yotvata in the south, on both sides of the border. A few were collected in the northeastern corner of the valley, in Jordan. The known records are located at elevations ranging from -290 m to 212 m.

Buthacus arava is sympatric with the closely related B. yotvatensis, in the Arava Valley (fig. 5), but allopatric with B. tadmorensis, distributed from central Jordan, through Syria, southeastern Turkey and Iraq, to the northern shore of the Persian Gulf, in southern Iran (fig. 10).

Ecology: The types of B. arava were collected from pitfall traps or at night with UV light detection on sparsely vegetated, stable to vegetationless, shifting inland sand dunes. Specimens collected at night were uncommon, sitting still or walking on the surface. The habitat and habitus of B. arava, notably the pale coloration, smooth tegument, loss or obsolescence of pedipalpal and metasomal carinae, reduced pectinal tooth count, elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins of the tibae and the pro- and retrolateral margins of the basitarsi of legs I–III, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and elongated, unequal length telotarsal ungues, are consistent with the ultrapsammophilous ecomorphotype (Prendini, 2001).

The types of B. arava were collected in sympatry with B. yotvatensis. Two other buthids, Androctonus crassicauda (Olivier, 1807) and Orthochirus scrobiculosus, also inhabit the same area, but occur on harder substrata.

Buthus arenicola Simon, 1885: 51, 52; Pocock, 1889: 116; Simon, 1892: 84; Kraepelin, 1895: 82; Birula, 1896: 244; 1905: 138.

Buthacus leptochelys (nec Androctonus leptochelys Ehrenberg, 1829): Pocock, 1895: 299.

Buthus (Buthacus) leptochelys arenicola: Birula, 1914: 636.

Buthacus arenicola: Pallary, 1929: 135, 137, 140, fig. 2C; 1934: 91, 95, 98, fig. 4C; 1938: 280, 281; Vachon, 1949a: 71–79, figs. 247, 253, 256–258, 260, 266; 1950a: 197; 1950b: 400, 407, fig. 586; 1951, 422, 424, 425, 428, 429, 431, 441, 446, 450, 452, 454, 464, figs. 642, 656, 685; 1952: 180, 185, 191, 194–196, 199,201-202 figs. 256, 266; 1958: 184, 185; Bücherl, 1964: 57; Larrouy et al., 1972: 393; Levy et al., 1973: 125, 128; Pérez, 1974: 19; Levy and Amitai, 1980: 89; Kinzelbach, 1985: map III; Vachon and Kinzelbach, 1987: 102; El-Hennawy, 1992: 97, 112 (part); Kovařík, 1998: 105; Fet and Lowe, 2000: 81, 82; Lourenço, 2001: 255; Crucitti and Vignoli, 2002: 440; Kovařík, 2002: 5; Lourenço, 2004a: 205; 2004b: 225; Kovařík and Whitman, 2004: 106 (misidentification); Kovařík, 2005: 1, 2, 6, 9, 10, fig. 1; Lourenço, 2006: 59, 60, 62–69, figs. 21–27; Lourenço and Qi, 2006a: 161; Kaltsas et al., 2008: 213; Zourgui et al., 2008: 81, 84, 85, 88, 89, table 2, plate 7, fig. 4 (misidentification); Lourenço and Leguin, 2009: 103; Lourenço, 2013: 89, 91; Lourenço and Sadine, 2015: 55; Lourenço et al., 2016: 3; 2017a: 33, fig. 1; 2017b: 20, 22, 24, fig. 1, table 1; Sadine et al., 2018: 51, 53–57, figs. 3C, 4, table 1; Bousmaha et al., 2019: 141–145, figs. 2, 3, table 1; Amr et al., 2021: 95, table 17.

Buthacus leptochelys arenicola: Pallary, 1929: 134, fig. 1A; Pallary, 1934: 91, 92, fig. 3A.

Buthacus leptochelys var. arenicola: Pallary, 1929: 135.

Buthacus arenicola arenicola: Vachon, 1949a: 76; 1950a: 197; 1952: 193, 196, figs. 253, 256–258, 260, 266; Bouisset and Larrouy, 1963: 295–297, figs. 1, 2; Pérez, 1974: 19; Kovařík, 1997b: 179; Lourenço, 2006: 65, 67, 69, fig. 42, table 1; 2013: 97, fig. 14; Lourenço and Sadine, 2015: 56, 57, fig. 1; Lourenço et al., 2016: 4, fig. 1.

Type Material: Buthus arenicola: Lectotype ♂ (MNHN RS 1672), TUNISIA: Tozzer [Tozeur, 33°54′N 08°06′E], V. May [examined]; paratype ♀ (MNHN), ALGERIA: Biskra [34°52′N 05°47′E]; paratype (MNHN), TUNISIA: Gabès [Quabis], Séd [33°53′N 10°05′E].

Diagnosis: Buthacus arenicola differs from the closely related species, B. levyi, occurring in Egypt and Israel, as follows. The pedipalp chela movable finger of the adult male is proportionally longer than the manus in B. arenicola (fig. 38A, B), with chela manus length:movable finger length, 54% (52.4%–55.6%, n = 2; table 10), than in B. levyi, with chela manus length:movable finger length, 62.8% (57.4%–67.3%, n = 8; table 10). The metasomal segments of the male are broader in B. arenicola (fig. 27D), with segment width:length, I–V, 72.2% (70.8%–73.6%, n = 2; table 10), 63.8% (60.9%–66.8%), 58.9% (58.9%–59%), 44.8% (43.1%–46.5%), and 34.5% (34%–35.1%), than in B. levyi (fig. 28A), with segment width:length, I–V, 65.1% (60.2%–70.8%, n = 8; table 10), 56.9% (53.9%–64.4%), 50.5% (47.6%–57.8%), 40.1% (35.9%–45%), and 33.8% (30.7%–38.9%). The dorsolateral carinae of metasomal segment IV are well developed in B. arenicola (figs. 23D, 25D), but obsolete to absent in B. levyi (figs. 24A, 26A).

Buthacus arenicola differs from B. amitaii, B. leptochelys and B. nitzani, as follows. The pedipalp chela of the adult male is longer and narrower in B. arenicola (fig. 38A, B), with chela manus length:movable finger length, 54% (52.4%–55.6%, n = 2; table 10) and chela manus width:chela length, 17.6% (17.1%–18.1%, n = 2), than in B. amitaii, B. leptochelys, and B. nitzani (figs. 32A, B, 41A, B, 47A, B), with chela manus length:movable finger length, 81.8% (72.7%–88.3%, n = 5; table 3), 70.7% (table 4), and 84.4% (76.8%–92.4%, n = 8; table 12), and chela manus width:chela length, 27.1% (25.1%–28.9%, n = 5), 24.2%, and 26.3% (22.3%–29.5%, n = 8), respectively. The count of retrolateral accessory denticles on the movable finger of the pedipalp chela is usually lower in B. arenicola (fig. 21C), with 1/1 (table 5), than in B. amitaii, B. leptochelys, and B. nitzani (fig. 21A, D, G), with 0–9/2–10 (n = 9; table 2), 7–9/7–9 (n = 5; table 5) and 6–9/6–9 (n = 16; table 2), respectively.

Description: The following redescription is based on two adult males collected from the vicinity of type locality (see table 5 for counts and table 10 for measurements).

Total length: Medium-sized scorpions, 54.3 mm (54.2–54.4 mm, n = 2) (♂).

Color: Uniformly pale yellow with legs slightly paler. Pectines whitish.

Carapace: Carapace shape trapezoidal (fig. 18C), anterior width:posterior width, 57.1% (55.6%–58.6%, n = 2), length:posterior width, 96.1% (94.3%–98%, n = 2). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them approximately 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 42.8% (42.3%–43.3%, n = 2) (♂). Superciliary and central median carinae distinct, weakly granular, and disconnected (♂). Anteromedian sulcus distinct, shallow; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulci deep, wide, curved. Carapace intercarinal surfaces finely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin and retrodorsal surfaces smooth, prolateral and ventral surfaces setose. Fixed finger dorsal and ventral surfaces densely setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger dorsal surface smooth and glabrous; ventral surface setose; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina comprising discontinuous row of spiniform granules and fewer than 15 macrosetae; dorsal, retrolateral, and intercarinal surfaces smooth; prolateral intercarinal surfaces finely granular (fig. 37A, B). Patella dorsal prolateral carinae obsolete; prolateral median and ventral prolateral carinae each comprising discontinuous row of spiniform granules; other carinae absent; intercarinal surfaces smooth (fig. 37C–E). Chela of male long and slender, manus width:length, 52.6% (50.8%–54.3%, n = 2) (♂), manus height:length, 62% (56.2%–67.7%, n = 2) (♂), and manus length:movable finger length, 54% (52.4%–55.6%, n = 2) (♂). Chela manus acarinate; intercarinal surfaces smooth and setose (fig. 38). Fixed and movable fingers respectively with 10 and 11 oblique median denticle subrows; movable finger with one retrolateral accessory denticle (fig. 21C); proximal dentate margins of fingers sublinear (fig. 37B), such that small gap present proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae absent; intercarinal surfaces smooth. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 7:12:13: 3; basitarsi, 12:18:19:10; telotarsi, 5:6:8:6 (n = 1). Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Telotarsal ungues long, approximately equal to telotarsus length, unequal on legs I and II, subequal to equal on III and IV (fig. 22A).

Genital operculum: Genital opercula suboval, completely divided longitudinally, with overlapping, rounded margins (♂) (fig. 20G) or partially fused longitudinally (♀). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 10–12 (n = 2) (♂) median lamellae (fig. 20G). Fulcra present. Pectinal teeth along most of length, dentate margin length:pecten length, 96.8% (96.1%–97.5%, n = 2) (♂). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 31–32/31–32 (n = 2) (♂).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 56.5% (53.6%–59.4%, n = 2) (♂); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae granular, vestigial, restricted to posterior fifth of I–IV and posterior third of V and VI; dorsosubmedian carinae obsolete to absent on I and II, granular, vestigial, restricted to posterior quarter of III and IV and posterior third of V and VI. Tergite VII, dorsomedian carina obsolete to absent; dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII smooth and glabrous; III–VI acarinate, VII, ventrolateral carinae vestigial, granular; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 23D, 25D, 27D), segment I shortest, length I:II, 89.9% (89.6%–90.2%, n = 2) (♂); segments II and III similar, length II:III, 97.9% (95.1%–100.7%, n = 2) (♂); segment III shorter than IV, length III:IV, 93.1% (90.8%–95.3%, n = 2) (♂); segment V longest, length IV:V, 82.9% (81.3%–84.6%, n = 2) (♂); width:length segment I, 72.2% (70.8%–73.6%, n = 2) (♂), II, 63.8% (60.9%–66.8%, n = 2) (♂), III, 58.9% (58.9%–59%, n = 2) (♂), IV, 44.8% (43.1%–46.5%, n = 2) (♂), V, 34.5% (34%–35.1%, n = 2); dorsosubmedian and dorsolateral carinae distinct, granular on segment I–III, obsolete on IV, absent on V; dorsosubmedian and dorsolateral carinae sparsely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 0/1 (0/1–0/1, n = 2):4/4 (3/3–4/4):4/3 (3/3–4/3):3/4 (3/3–3/4):0/0 (0/0–0/0), dorsolateral carinae, 2/3 (2/2–2/3, n = 2):3/3 (3/2–3/4):3/3 (3/2–3/3):3/2 (2/2–3/2):4/3 (3/2–4/3). Median lateral carinae distinct, granular, extending entire length of segment I, restricted to posterior half of II and posterior quarter of III, absent on IV and V. Ventrolateral carinae distinct, costate, becoming granular posteriorly on segments I–IV; serrate, comprising spiniform granules, becoming more prominent posteriorly, on V. Ventrosubmedian carinae distinct, costate on segment I; costate, becoming costate-granular posteriorly, on II–IV; granular on V. Ventromedian carina obsolete, granular on segment V. Dorsal and lateral intercarinal surfaces smooth on segments I–V; ventral intercarinal surfaces smooth on I, finely and sparsely granular on II–IV, finely and densely granular across entire surface of V.

Telson: Telson vesicle width:metasomal segment V width, 74.3% (72.2%–76.3%, n = 2) (♂). Vesicle globose, dorsal surface flat, ventral surface convex and rounded; vesicle height:length, 59.1% (58.7%–59.5%, n = 2) (♂); dorsal and ventral surfaces smooth and glabrous; lateral and ventral surfaces sparsely setose, with 39 (38–40, n = 2) (♂) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 54.5% (54.3%–54.7%, n = 2) (♂).

Sexual dimorphism: The pedipalp chela manus of the male is slightly broader, with proportionally shorter fingers than that of the female. The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female, and genital papillae are present in the male but absent in the female. Additionally, the pectinal tooth count is higher and the mesosoma relatively narrower in the male than in the female.

Distribution: Buthacus arenicola appears to be endemic to central Tunisia and northeastern Algeria, south of the Tell Atlas mountain range and north of the Chott Melhrir and Chott el Djerid salt lakes (fig. 8), from approximately sea level (5 m) to 126 m elevation (Lourenço et al., 2017b; L.P., personal obs.). It is doubtful whether populations from elsewhere, including those formerly assigned to Buthacus spatzi (Birula, 1911), stat. rev., are conspecific based on DNA sequence data and morphology (L.P., unpublished data). Buthacus arenicola is allopatric with the closely related B. spatzi, occurring south of the Chott Melhrir and Chott el Djerid salt lakes, in southern Tunisia and western Libya, and B. levyi, endemic to Egypt and Israel.

Ecology: Specimens of B. arenicola were collected at night with UV light detection on low, stable vegetated white sand dunes with scattered gypsum outcrops and small shrub-coppice dunes around grass tussocks. Specimens were resting or walking on ground and bushes, mostly on shrub-coppice dunes. The habitat and habitus, notably the pale coloration, smooth tegument, loss or obsolescence of pedipalpal and metasomal carinae, elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, with comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and elongated telotarsal ungues, are consistent with the psammophilous ecomorphotype (Prendini, 2001). Buthacus arenicola was collected in sympatry with another buthid, Androctonus amoreuxi, near Naftah, Tunisia.

Remarks: Simon (1885) described B. arenicola from Gabès (Séd) and Tozzer [Tozeur] in central Tunisia, noting that the species can also be found in Algeria (Biskra, Bou-Sadaa, and Debila) and Egypt (Port-Saïd and Ramleh). Pocock (1895) subsequently synonymized B. arenicola with B. leptochelys, a decision accepted by Kraepelin (1899) and Simon (1910). Birula (1908) only considered specimens cited by Simon (1885) from Egypt conspecific with B. leptochelys, however. Pallary (1938) revalidated B. arenicola and Vachon (1949a) distinguished B. arenicola and B. leptochelys based on the absence or presence of retrolateral accessory denticles on the movable finger of the pedipalp chela. Following Vachon's (1952) suggestion that B. arenicola and B. leptochelys represent species complexes and not single species, Levy and Amitai (1980) recognized the B. arenicola and B. leptochelys species groups. Levy and Amitai (1980) noted that the distributions of B. arenicola and B. leptochelys overlap in parts of Egypt and that the population of B. arenicola in the Sinai Peninsula differs morphologically from the population in Algeria and Tunisia, suggesting it may represent a distinct subspecies, an opinion shared by Lourenço (2006).

During the present investigation, the types of B. arenicola and B. leptochelys were examined, in addition to material from the vicinity of the type locality of B. arenicola, in Tunisia, and from Egypt, including the Sinai Peninsula, and Israel. These comparisons reinforced the distinction between B. arenicola and B. leptochelys, and confirmed previous suggestions that the population of B. arenicola in Egypt and Israel is distinct from the typical population in Algeria and Tunisia (Levy and Amitai, 1980; Lourenço, 2006). Based on evidence gathered during the present investigation, including multivariate analysis of morphometrics (fig. 15) and multilocus molecular phylogenetics (fig. 14), B. arenicola is restricted to central Tunisia and northeastern Algeria and the population occurring in Egypt and Israel represents a distinct species, described below.

Buthacus spatzi, described from the Sahara of southern Tunisia (Birula, 1911), was synonymized with B. arenicola by Lourenço (2006), based on an assessment of the original description and an unpublished opinion of M. Vachon (in litt.). The types of B. spatzi could not be located at the time of Lourenço's (2006) study, nor during the present investigation, and appear to be lost. However, material collected from Jebil and elsewhere in southern Tunisia, matching the descriptions of B. spatzi in Birula (1911) and Lourenço (2006), was found to be morphologically and genetically distinct from material collected near Naftah, in the vicinity of the type locality of B. arenicola (L.P., unpublished data), questioning the decision to synonymize. Buthacus spatzi (Birula, 1911), stat. rev., is therefore revalidated. The limits of B. arenicola and B. spatzi, beyond the scope of the present investigation, will be redefined more precisely, elsewhere. Buthacus birulai Lourenço, 2006, described from El Oued, Algeria, will probably prove conspecific with either B. arenicola or B. spatzi.

Buthacus arenicola fuscata Pallary, 1929, described from In Ameri [23°17′N 05°30′E] in the Hoggar Mountains of southern Algeria, and synonymized with B. arenicola by El-Hennawy (1992), is unlikely to be conspecific with the latter, and is hereby revalidated and elevated to the rank of species. Buthacus fuscata Pallary, 1929, stat. nov. et stat. rev., is probably a senior synonym of one or more of the other species of Buthacus described from the vicinity of the Hoggar and Tassili n'Ajjer mountain ranges, e.g., Buthacus ahaggar Lourenço, 2017, Buthacus armasi Lourenço, 2013, or Buthacus foleyi Vachon, 1948, none of which were compared with it in the original descriptions. Which of these species are junior synonyms requires further investigation.

Material Examined: TUNISIA: Tozeur Governorate: Entrée Nefta, 24.v.1994, S. Nouira, dunes de sable, barkhanes, chasse nocturne, UV, 1 ♀, 1 juv. ♂, 3 juv. ♀ (UTM 153); Naftah, ca. 15 km W on P3 to Hazoua, 33°51.209′N 07°44.011′E, 6 m, 24.x.2017, L. Prendini, J. Babay, M. Khammasi and R. Ouni, 1 subad. ♂, 1 juv. ♀ (AMCC [LP 14859]); Naftah, ca. 7 km N on road to Ong Jmal, 33°56.366′N 07°50.321′E, 17 m, 23.x.2017, L. Prendini and J. Babay, 1 ♂, 11 ♀, 5 subad. ♂, 12 subad. ♀, 2 juv. ♂, 4 juv. ♀ (AMNH), 7 juv. ♂, 4 juv. ♀(AMCC [LP 14858]); Naftah, ca. 8 km N on road to Ong Jmal, 33°56.588′N 07°50.228′E, 5 m, 23.x.2017, L. Prendini, J. Babay and R. Ouni, 1 juv. ♂ (AMCC [LP 14857]).

Androctonus (Leiurus) leptochelys Ehrenberg in Hemprich and Ehrenberg, 1829: 355.

Androctonus (Leiurus) thebanus Ehrenberg in Hemprich and Ehrenberg, 1828: pl. 1, fig. 4; Ehrenberg in Hemprich and Ehrenberg, 1829: 355, 356; Braunwalder and Fet, 1998: 30, 32 fig. 2.

Androctonus (Leiurus) macrocentrus Ehrenberg in Hemprich and Ehrenberg, 1828: pl. 1, fig. 6 (synonymized by Kraepelin, 1891: 60); Ehrenberg in Hemprich and Ehrenberg, 1829: 355; Vachon, 1952: 199, 201; Braunwalder and Fet, 1998: 30, 32 fig. 2; syn. nov.

Androctonus (Liurus) leptochelys: Hemprich and Ehrenberg, 1831; Moritz and Fischer, 1980: 317.

Androctonus (Liurus) macrocentrus: Hemprich and Ehrenberg, 1831; Moritz and Fischer, 1980: 317.

Androctonus (Liurus) thebanus: Hemprich and Ehrenberg, 1831; Moritz and Fischer, 1980: 324.

Scorpio (Androctonus) leptochelis: Gervais, 1844: 44.

Scorpio (Androctonus) macrocentrus: Gervais, 1844: 44.

Scorpio (Androctonus) thebanus: Gervais, 1844: 44.

Androctonus leptochelys: C.L. Koch, 1845: 7–9, pl. CCCXCIX, fig. 964.

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

Buthus leptochelis: Simon, 1872: 250, 251; Lucas, 1873: 172; Simon, 1879: 99; 1880: 29; 1892: 83; Birula, 1905: 138.

Buthus macrocentrus: Simon, 1879: 99.

Leiurus macrocentrus: Karsch, 1881a: 8.

Leiurus thebanus: Karsch, 1881a: 8.

Buthus leptochelys: Karsch, 1881a: 8; 1881b: 89; Kraepelin, 1891: 60–62, pl. I, fig. 8; Pocock, 1895: 299, 300; Kraepelin, 1899: 17; 1901: 266; King, 1925: 81; Gough and Hirst, 1927: 3, fig. 3; Schenkel, 1932: 379.

Buthus (Buthacus) leptochelys: Birula, 1908: 140; 1910: 155–157, 170; 1917: 21, 23, 214, 224, 229; Giltay, 1929: 196; Werner, 1929: 30; Whittick, 1947: 122; 1955: [unpaginated] (part).

Buthacus leptochelys: Simon, 1910: 75, 76, fig. 11; Borelli, 1914: 158, 159; 1927: 352; Pallary, 1929: 133–135; Caporiacco, 1932: 396; Borelli, 1934: 172; Pallary, 1934: 90–92, 99 (misidentification); Werner, 1934: 269, fig. 332; Caporiacco, 1936: 94, 98; Werner, 1936a: 176; Bodenheimer, 1937: 235; Caporiacco, 1937: 347; Sergent, 1941: 355, fig. 2.9, pl. 35, fig. 9; Vachon, 1949a: 79–83, figs. 262–266; 1950a: 197; 1950b: 406 (part); 1951: 432–434, 436, 437, 441, 446, 447, 450, 453, 456, 458, 460, 462, 465, figs. 661, 677, 697 (misidentification); 1952: 180, 185, 191, 194, 198–203, figs. 262–266 (part); 1954: 188 (misidentification); 1958: 184; Pringle, 1960: 76, fig. 2 (misidentification); L. Khalaf, 1962: 1, 2 (misidentification); K.I. Khalaf, 1963: 59 (misidentification); Vachon, 1966: 210; Habibi, 1971: 43 (misidentification); Larrouy et al., 1972: 393; Levy et al., 1973: 126, 128, 134; Pérez, 1974: 19; Lamoral and Reynders, 1975: 499; Vachon, 1979: 31, 36–39, 41, 47, 49, 63, 65, figs. 7, 8, 26, 29, 61–63 (part); Levy and Amitai, 1980: 77, 78; Kettel, 1982: 6; Kinzelbach, 1984: 99; 1985: map III; Vachon and Kinzelbach, 1987: 101; Amr et al., 1988: 374; El-Hennawy, 1987: 17; Farzanpay, 1988: 36 (misidentification); El-Hennawy, 1991: 86, 87; 1992: 97, 101, 112, 113 (part); Amr and El-Oran, 1994: 187, 188 (misidentification); Kovařík, 1998: 105; Braunwalder and Fet, 1998: 32–34; Fet and Lowe, 2000: 82–84; ICZN, 2000: 7; Kovařík, 2001: 80; Lourenço, 2001: 255; Stathi and Mylonas, 2001: 288; Crucitti and Vignoli, 2002: 440, 441; Kovařík, 2002: 5; 2003: 135, 155, table 1 (misidentification); Lourenço, 2003: 877, 880, fig. 10 (misidentification); 2004a: 205; 2004b: 225, 226, 229; Kovařík, 2005: 1–3, 6, 8, 9, 10, figs. 2, 5, 7; Lourenço, 2006: 59–69, figs. 1, 4, table 1; Lourenço and Qi, 2006a: 161; Kaltsas et al., 2008: 214; Kamenz and Prendini, 2008: 8, 40, table 2 (misidentification); Navidpour et al., 2008: 8; Zourgui et al., 2008: 81, 84, 85, 88, 89, table 2, plate 5, fig. 4 (misidentification); Al-Asmari et al., 2009a: 612, 613, 617–619, 621, 622, 624, 626, figs. 9, 12G, tables 1, 2 (misidentification); 2009b: 96, 100, 102, 104, 106, fig. 6K, L, tables 5–7 (misidentification); Lourenço and Leguin, 2009: 103; Zarei et al., 2009: 46–49, 51, fig. 2, table 1 (misidentification); Lourenço et al., 2012: 315; Al-Asmari et al., 2013: 2, 5–7, 10–12, 15, figs. 6, 7, 9,10, 12A, table 1 (misidentification); Kovařík et al., 2013: 3; Lourenço, 2013: 89, 91, 93; Amr, 2015: 183–185, 192, 197, figs. 1, 2, table 3 (misidentification); Amr et al., 2015: 32; Lourenço and Sadine, 2015: 55; Kovařík et al., 2016: 1, 2; Lourenço et al., 2016: 3; Badry et al., 2018: 77, 83; Kovařík, 2018: 1, 9, 10; Saleh et al., 2017: 7–9, fig. 3, table 2; Francke, 2019: 20; Lowe et al., 2019: 22; Alqahtani and Badry, 2020: 178–183, figs. 1–3, tables 1, 2; Hussen and Ahmed, 2020: 6722; Obuid-Allah et al., 2020: 227, 229–237, figs. 2G, H, 3D, tables 1, 3–6; Amr et al., 2021: 85, 88–90, tables 4, 5, 9, 10 (misidentification); Said et al., 2021: 17, 18, 20, 21, 25, 26, fig. 4, table 1.

Buthacus granosus Borelli, 1929: 297–299, pl. VI, figs. 1–3; Probst, 1973: 329; Lamoral and Reynders, 1975: 499; syn. nov.

Buthus granosus: Vachon, 1949b: 162; 1952: 248.

Buthacus leptochelys: Le Corroller, 1967: 63.

Buthacus macrocentrus: Kovařík, 2005: 1, 7, 9, 10, fig. 8 (part); Lourenço, 2006: 63, 64, figs. 15, 16; Lourenço and Qi, 2006a: 161; Kovařík et al., 2016: 1; Kovařík, 2018: 10.

Buthacus thebanus: Vachon, 1952: 180; Pérez, 1974: 19; Lourenço, 2006: 62.

Buthacus leptochelys leptochelys: Levy et al., 1973: 125; Levy and Amitai, 1980: 76, 77, 79–83, figs. 70–74, map 6; Kinzelbach, 1984: 99, 101 (misidentification); Kovařík, 1995: 19 (misidentification); Kovařík, 1997a: 49 (misidentification); Kabakibi et al., 1999: 82, 88 (misidentification); Fet and Lowe, 2000: 84; Hendrixson, 2006: 50; Lourenço, 2006: 62, 69, fig. 42.

Buthacus leptochelys granosus: Levy et al., 1973: 125; Vachon, 1979: 38; Levy and Amitai, 1980: 76; El-Hennawy, 1992: 97, 113; Kovařík, 1998: 105; Fet and Lowe, 2000: 84.

Type Material: Androctonus (Leiurus) leptochelys: Lectotype ♀, paralectotype ♀ (No. 1) (ZMB 152), Sinai [examined]. Androctonus (Leiurus) macrocentrus: Lectotype ♀, paralectotype ♀ (ZMB 153), EGYPT: Sinai [examined]. Androctonus (Leiurus) thebanus: Lectotype ♂, paralectotype ♂ (ZMB 154), EGYPT: Luxor Governorate: Thebae [Thebes, Luxor, 25°43′N 32°37′E]. Buthacus granosus: Holotype ♂(MIZT), SUDAN: Red Sea State: Port Sudan [19°33′N 37°11′E], 24.iv.1926, H.B. Johnston.

Diagnosis: Buthacus leptochelys differs from the closely related species B. amitaii and B. nitzani, occurring in Israel, as follows. Buthacus leptochelys is larger, measuring 65.8 mm (63.1–68 mm, n = 5; table 4) in total length, than B. amitaii, measuring 51.7 mm (48–58.8 mm, n = 10; table 3) and B. nitzani, measuring 50.6 mm (44.4–58.5 mm, n = 18; tables 12, 13). The pedipalp chela of the adult male is longer and narrower (fig. 41A, B), with chela manus width:chela length, 24.2% (table 4) and chela manus height:length, 58.9%, in B. leptochelys than in B. amitaii (fig. 32A, B), with chela manus width:chela length, 27.1% (25.1%–28.9%, n = 5; table 3) and chela manus height:length, 69.7% (58.9%–75.5%, n = 5), and B. nitzani (fig. 47A, B), with chela manus width:chela length, 26.3% (22.3%–29.5%, n = 8; table 12) and chela manus height:length, 62.5% (57.7%–66.6%, n = 8). The proximal dentate margins of the pedipalp chela fixed and movable fingers are shallowly emarginate to sublinear, such that a less pronounced gap is evident proximally when the fingers are closed in B. leptochelys (fig. 41B) than in B. amitaii and B. nitzani (figs. 32B, 47B). The pectinal tooth counts are higher in B. leptochelys, i.e., 35/34 (32–39/32–38, n = 6; table 5) (♂) and 29/29 (27–31/28–31, n = 10) (♀), than in B. amitaii, i.e., 30/31 (28–32/28–33, n = 5; table 2) (♂) and 23/23 (21–26/21–26, n = 9) (♀), and B. nitzani, i.e., 31/31 (26–34/27–35, n = 6; table 2) (♂) and 23/23 (20–27/21–27, n = 17) (♀). The metasomal segments of the female are longer and narrower in B. leptochelys (fig. 28B), with the segment IV width:length, 42.1% (40.3%–44.6%, n = 4; table 4), than in B. nitzani (fig. 28C), 38.4% (34%–43.7%, n = 10; table 13). The median lateral carinae of metasomal segments I–III are distinct in B. leptochelys, but obsolete in B. nitzani. The telson dorsal surface is flat and the ventral surface convex and rounded in lateral profile in B. leptochelys (fig. 29D), whereas the dorsal surface is concave and the ventral surface convex and angular in lateral profile in B. amitaii and B. nitzani (fig. 29F, H).

Buthacus leptochelys differs from B. levyi, occurring in sympatry in Egypt, including the Sinai Peninsula, as follows. The pedipalp chela of the adult male is broader and deeper in B. leptochelys (fig. 41A, B), with chela manus width:length, 59.1% and chela manus height:length, 58.9% (table 4), than in B. levyi (fig. 44A, B), with chela manus width:chela length, 19.4% (17.1%–21.3%, n = 8; table 10) and chela manus height:length, 59.7% (55.8%–64%, n = 8). The proximal dentate margins of the pedipalp chela fixed and movable fingers are shallowly emarginate, such that a small gap is evident proximally when the fingers are closed, in B. leptochelys (fig. 41B), but sublinear, such that no gap is evident proximally when the fingers are closed, in B. levyi (fig. 44B). The retrolateral accessory denticles on the movable finger of the pedipalp chela are present, with higher counts, in B. leptochelys (sinistral/dextral: 7–9/7–9; n = 5; table 5; fig. 21D), than in B. levyi, in which the denticles are often absent (0–5/0–3; n = 13; table 5; fig. 21E, F).

Description: The following description is based on specimens collected in Egypt, including Sinai (see table 4 for measurements and table 5 for counts).

Total length: Large scorpions, 63.7 mm (♂) or 66.3 mm (63.1–68 mm, n = 4) (♀).

Color: Uniformly yellow. Legs paler with articulation points brown. Pectines whitish.

Carapace: Shape trapezoidal (fig. 17A, B), anterior width:posterior width, 49.7% (43.7%–56.5%, n = 5), length:posterior width, 84.8% (77.4%–88.7%, n = 5). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 40.1% (♂) or 44% (41%–45.9%, n = 4) (♀). Superciliary and central median carinae distinct, costate-granular, strongly to weakly connected. Anteromedian sulcus distinct; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulcus deep, wide, curved. Carapace intercarinal surfaces coarsely granular.

Chelicerae: Cheliceral manus prodorsal margin granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces densely setose. Fixed finger dorsal and ventral surfaces densely setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger dorsal surface smooth and glabrous; ventral surface densely setose; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina comprising discontinuous row of spiniform granules and fewer than 10 macrosetae; intercarinal surfaces smooth (fig. 40A, B). Patella prolateral median and ventral prolateral carinae each comprising discontinuous row of spiniform granules; dorsal prolateral carinae obsolete; other carinae absent; intercarinal surfaces smooth (fig. 40C–E). Chela long and broad (♂), manus width:length, 59.1%, manus height:length, 58.9%, and manus length:movable finger length, 70.7% or long and slender (♀), manus width:length, 51.7% (50.4%–52.4%, n = 4), manus height:length, 60.5% (58.4%–63.2%, n = 4), and manus length:movable finger length, 51.9% (48.2%–54.1%, n = 4). Chela manus acarinate; intercarinal surfaces smooth and glabrous (fig. 41). Fixed and movable fingers respectively with 7–11 (n = 5) and 10–11 (n = 5) oblique median denticle subrows; movable finger with 7–9 (n = 5) retrolateral accessory denticles (fig. 21D); proximal dentate margins of fingers sublinear (fig. 41B), such that small gap weakly present when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae obsolete; intercarinal surfaces smooth. Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 10:11:11:3; basitarsi, 9:13:21:9; telotarsi, 6:8:9:7 (n = 1). Telotarsal ungues long, approximately equal to telotarsus length, unequal on legs I and II, subequal to equal on III and IV (fig. 22D).

Genital operculum: Genital opercula suboval, completely divided longitudinally, with overlapping, rounded margins (♂) or partially fused longitudinally (♀). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 11–13 (n = 3) (♂) or 11–14 (n = 2) (♀) median lamellae. Fulcra present. Pectinal teeth along most of length, dentate margin length:pecten length, 90.8% (♂) or 91.8% (90.7%–92.9%, n = 4) (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 35/34 (32–39/32–38, n = 6) (♂) or 29/29 (27–31/28–31, n = 10) (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 58.4% (♂) or 57.2% (55%–59.5%, n = 4) (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae granular, vestigial, restricted to posterior fifth of I–III and posterior two-thirds of IV–VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–III, and posterior two-thirds of IV–VI. Tergite VII, dorsomedian carina granular, vestigial, restricted to anterior half; dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII acarinate; III sparsely granular, IV–VII smooth and glabrous; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 24B, 26B, 28B), segment I shortest, length I:II, 87.5% (♂) or 87.5% (86.6%–89%, n = 4) (♀); segments II–IV similar, length II:III, 95.1% (♂) or 97.7% (95.7%–99.7%, n = 4) (♀), length III:IV, 94.6% (♂) or 98.7% (95.4%–101.7%, n = 4) (♀); segment V longest, length IV:V, 87% (♂) or 84.5% (78.8%–79.7%, n = 4) (♀); width:length segment I, 66.9% (♂) or 68.4% (66.2%–70%, n = 4) (♀), II, 52.8% (♂) or 55.7% (53.3%–57.6%, n = 4) (♀), III, 49.4% (♂) or 51.3% (48.4%–52.6%, n = 4) (♀), IV, 37.9% (♂) or 42.1% (40.3%–44.6%, n = 4) (♀), V, 33.4% (♂) or 37.5% (35.2%–39.3%, n = 4) (♀); dorsosubmedian and dorsolateral carinae distinct, granular on segments I–III, dorsosubmedian carinae obsolete, dorsolateral carinae granular on IV; dorsosubmedian and dorsolateral carinae absent on V; dorsosubmedian and dorsolateral carinae sparsely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 1/0 (0/0–1/0, n = 2):2/3 (2/3–2/3):3/3 (3/3–3/3):2/2 (1/1–2/2):0/0 (0/0–0/0), dorsolateral carinae, 2/3 (2/2–2/3, n = 2):2/3 (1/2–3/3):4/3 (3/2–4/3):5/4 (4/3–6/4):4/5 (4/5–4/5). Median lateral carinae distinct, granular, extending entire length of segment I, restricted to posterior half of II and posterior third of III, absent on IV and V. Ventrolateral carinae distinct, costate-granular on segments I–IV; serrate, comprising spiniform granules of variable size, becoming more prominent posteriorly on V. Ventrosubmedian carinae distinct, costate-granular on segment I; granular, with granules becoming progressively larger and subspiniform posteriorly on II and III; costate-granular on IV; granular, restricted to anterior three-quarters of V. Ventromedian carina granular, distinct along entire length of segment V. Dorsal and lateral intercarinal surfaces smooth to finely and sparsely granular on segment I, smooth on II–V; ventral intercarinal surfaces smooth on I and II, finely and sparsely granular on III, finely granular across entire surface on IV and V.

Telson: Telson vesicle width:metasomal segment V width, 81.3% (♂) or 77.2% (76.2%–77.8%, n = 4) (♀). Vesicle globose, dorsal surface flat, ventral surface convex and rounded; vesicle height:length, 58.4% (♂) or 63.3% (57.2%–67.8%, n = 4) (♀); dorsal and ventral surfaces smooth and glabrous; lateral and ventral surfaces sparsely setose, with 20 (♂) or 25 (♀) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 49.4% (♂) or 56.7% (56%–58.4%, n = 4) (♀).

Sexual dimorphism: Adult males and females differ as follows. The pedipalp chela manus of the male is slightly broader than that of the female (fig. 41), as indicated by the higher chela manus width:length ratio in the male (59.1%) compared with the female (58.9%). The pedipalp chela manus of the male is markedly broader and deeper with proportionally shorter fingers than that of the female (fig. 41), as indicated by the higher ratios of chela manus width:length and chela manus length:movable finger length, respectively 59.1% and 70.7% in the male compared with 58.9% and 51.9% in the female. The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female, and genital papillae are present in the male but absent in the female. The pectinal tooth count is higher in the male (32–33) than in the female (27–30). The mesosoma of the male is relatively narrower than that of the female, as indicated by the higher sternite VII length:width ratio in the male (66.5%) compared with the female (57.5%). The spiniform granules of the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and the ventrolateral carinae of segment V are less prominent in the male than in the female.

Distribution: Buthacus leptochelys appears to be distributed from the western Sinai Peninsula across Egypt, and southward along the coast of the Red Sea to Sudan (fig. 6). It probably also occurs in Libya. The known records range from -28 m to 1155 m in elevation.

Buthacus leptochelys is allopatric with the closely related B. amitaii, endemic to the inland sandy soils and loess of the Yamin Plain and Mamshit area in the northern Negev, Israel (fig. 7), and B. nitzani, distributed from the Sorek (Rubin) River, throughout the sand dunes of the southern coastal plain of Israel, to the Haluza dunes of the interior and probably the adjacent Sinai Peninsula of Egypt (fig. 7).

Ecology: Specimens of B. leptochelys were collected at night with UV light detection in arid to hyperarid sandy habitats, including gravel plains and soft sandy plains, inland and coastal sand dunes, with variable vegetation cover (Badry et al., 2018; L.P., personal obs.). The habitat and habitus, notably the pale coloration, smooth tegument, loss or obsolescence of pedipalpal and metasomal carinae, elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins of the tibae and the pro- and retrolateral margins of the basitarsi of legs I–III, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and elongated, unequal-length telotarsal ungues, are consistent with the ultrapsammophilous ecomorphotype (Prendini, 2001).

Buthacus leptochelys has been collected in sympatry with the following buthids: Androctonus australis (Linnaeus, 1758), Buthus intumescens (Ehrenberg, 1829), Compsobuthus egyptiensis Lourenço et al., 2009, Leiurus quinquestriatus, Orthochirus aristidis (Simon, 1882), Orthochirus innesi Simon, 1910, and Parabuthus abyssinicus Pocock, 1910 (Badry et al., 2018; L.P., personal obs.).

Remarks: Ehrenberg (1829) described B. leptochelys and B. macrocentrus from the same region, the Sinai Peninsula, one year after describing B. thebanus from Thebes, near Luxor, in southern Egypt. Kraepelin (1891) regarded the three species from Egypt and the Sinai Peninsula as a single species, B. leptochelys. Pocock (1895) subsequently synonymized B. arenicola with B. leptochelys, an opinion accepted by Kraepelin (1899), who also synonymized B. tadmorensis with B. leptochelys. Birula (1908) revalidated B. arenicola and B. tadmorensis, but the limits and validity of B. arenicola, B. macrocentrus, and B. tadmorensis continued to be debated thereafter (Levy et al., 1973; Levy and Amitai, 1980; Kinzelbach, 1985; Fet and Lowe, 2000; Kovařík, 2005, 2018; Lourenço, 2006; Lourenço and Qi, 2006a; Lourenço and Leguin, 2009; Kovařík et al., 2016).

Levy et al. (1973) downgraded Buthacus granosus Borelli, 1929, described from Port Sudan on the coast of the Red Sea, to a subspecies of B. leptochelys, without justification. Comparison of Borelli's (1929) description of B. granosus with the redescription of B. leptochelys presented here revealed few differences that might merit their recognition as different taxa, justifying the following synonymy: Buthacus granosus Borelli, 1929 = Buthacus leptochelys (Ehrenberg, 1829), syn. nov.

Kovařík (2018) recently synonymized Buthacus armasi Lourenço, 2013, from the Tassili n'Ajjer of southern Algeria, with B. leptochelys. Setting aside the vast geographical distance between the type locality of B. armasi and the known locality records of B. leptochelys, most of which occur in Egypt, comparison of Lourenço's (2013) description and illustrations with specimens of B. leptochelys from Egypt including the Sinai Peninsula examined during the present investigation casts doubt on Kovařík's (2018) decision. For example, differences are evident in the dentition of the pedipalp chela fingers of B. armasi and B. leptochelys, specifically the curvature of the median denticle subrows and the deviations of the retrolateral accessory denticles from the subrows, cf. Lourenço's (2013: 94, 96) description and figure 9 with figure 21D of the present contribution. The two species also differ in pectinal tooth counts, with 27–28 (♂) or 21–23 (♀) in B. armasi and 35/34 (32–39/32–38, n = 6) (♂) or 29/29 (27–31/28–31, n = 10) (♀) in B. leptochelys. The ventral carinae of the metasoma of B. armasi were described as “vestigial on segment I, weak on II, moderate on III–IV” (Lourenço, 2013: 96), whereas in B. leptochelys these carinae are distinct on segments I–IV. Unpublished DNA sequence data from samples originating in the vicinity of the type locality of B. armasi are also inconsistent with the hypothesis that the two taxa are conspecific. Insofar as the evidence refutes the hypothesis that it is conspecific with B. leptochelys, Buthacus armasi Lourenço, 2013, stat. rev., is hereby revalidated, although it may prove to be a junior synonym of B. fuscata and/or B. foleyi, two other species described from the vicinity of the Hoggar and Tassili n'Ajjer mountain ranges, neither of which were compared with it in the original description.

During the present investigation, the types of B. arenicola, B. leptochelys, and B. macrocentrus were examined in addition to material from the vicinity of the type locality of B. arenicola in Tunisia and from Egypt and Israel. These comparisons, including multivariate analysis of morphometrics (fig. 15) and multilocus molecular phylogenetics (fig. 14), confirmed the validity of B. arenicola and B. leptochelys. Kraepelin's (1891) synonymy of A. (L.) thebanus with B. leptochelys was also confirmed, the lectotype male of A. (L.) thebanus closely matching a male B. leptochelys from Wadi Sidri (HUJ INVSC 859).

As concluded by Kovařík (2005), the type series of B. leptochelys comprises more than one species: paralectotype 2 is heterospecific with the lectotype and paralectotype 1, instead representing a species of the B. arenicola group. However, Kovařík (2005) misidentified paralectotype 2 as the Tunisian species, B. spatzi; it is conspecific with B. levyi, the distribution of which overlaps that of B. leptochelys in the Sinai Peninsula (fig. 9).

Also contrary to Kovařík (2005) and others (e.g., Lourenço, 2006), the types B. macrocentrus were found to be conspecific with one another and with the types of B. leptochelys, as originally deduced by Kraepelin (1891) and confirmed by Vachon (1952) and Levy and Amitai (1980). No evidence was found to support Kovařík's (2005) contention that the types of B. macrocentrus are heterospecific, with only the paralectotype conspecific with B. leptochelys. The lectotype of B. macrocentrus closely resembles the paralectotype of B. macrocentrus, the lectotype and paralectotype 1 of B. leptochelys, and other material of B. leptochelys from southwestern Sinai (e.g., HUJ INVSC 1807, 2233, 2278) in the armature of the leg tarsi and the shape and ventral carination of the metasomal segments, notably II and III, and differs markedly in these characters from B. tadmorensis and B. yotvatensis, two species erroneously synonymized with B. macrocentrus by Kovařík (2005) and revalidated below. Similarly, Kovařík's (2005: 9) assertion that the “Type locality ‘Sinai’ (Ehrenberg in Hemprich and Ehrenberg, 1829: 355 and label) must be regarded as erroneous” is unfounded. The type locality of B. macrocentrus is consistent with the type locality of B. leptochelys as well as the known distribution of that species (fig. 6). In view of the evidence, B. macrocentrus is returned to synonymy: Androctonus (Leiurus) macrocentrus Ehrenberg, 1829 = Buthacus leptochelys (Ehrenberg, 1829), syn. nov.

Material Examined: EGYPT: 1994, ex. M. Scharmach, 1 ♀ (AMNH [LP 324]). Sinai, 1890, Ehrenberg, 1 ♀ (ZMH ex Mus. Berol.). Caïro Governorate: Caïro, vi.1913, 1 ♂ (ZMH ex Mus. Basel). Faiyum Governorate: Faiyum Oasis, N side, 29°32′11″N 30°48′08.8″E, -28 m, 24.ii.2019, L. Prendini, E. Attia, T. Saïd, F. Sayed, and N. Abdelrazik, 1 ♀, 4 subad. ♂, 6 subad. ♀, 23 juv. ♂, 23 juv. ♀ (AMNH), 5 juv. ♂, 2 juv. ♀(AMCC [LP 16600]). North Sinai Governorate: Gebel Iqme [28°38′N 33°18′E], 1 ♀ (SMNH TAU AR 50805); Mitla Pass [Ma'avar Ha Mithleh], 30°00′N 32°55′E, 414 m, xi.1970, J. Gershoni, 1 ♀ (HUJ INVSC 1807), 15.xii.1970, A. Bara'am, 1 ♀ (HUJ INVSC 2233). Red Sea Governorate: Zafarana, ca. 22 km SW on al Kurimat–al Zafrana Road, 23.ii.2019, L. Prendini, E. Attia, M. Omran, and T. Saïd, 29°04′36.3″N 32°27′07.9″E, 109 m, 4 juv. ♀ (AMNH), 1 juv. ♂, 2 juv. ♀ (AMCC [LP 16598]), 29°04′22″N 32°26′48.9″E, 129 m, 2 subad. ♂, 1 juv. ♂, 2 juv. ♀ (AMNH), 1 juv. ♂, 3 juv. ♀ (AMCC [LP 16599]). South Sinai Governorate: Ram-let Himayir (Sinai), sands SE of Abu Zanima, 29°04′N 33°27′E, 509 m, 25.xi.1969, A. Haim, 1 juv. ♀ (HUJ INVSC 1545); on way to Sarabitel-Hadem, 29°03′N 33°25′E, 485 m, 4.iii.1970, A. Haim, 1 juv. ♂ (HUJ INVSC 1750); Shechel Ka'rai, 29°05′N 33°20′E, 523 m, 8.iv.1968, Oranim Seminar, 1 subad. ♀ (HUJ INVSC 857); Wadi Sidri, Abu Rudies area, 28°51′N 33°15′E, 113 m, 3.vii.1968, N. Tademor and A. Haim, 1 ♂ (HUJ INVSC 859); Wadi Zalka, 28°52′N 34°00′E, 1155 m, 1.iii.1970, A. Haim, 1 juv. ♂ (HUJ INVSC 1749), 1 juv. ♀ (HUJ INVSC 1748). Suez Governorate: E of Suez City, 29°55′N 32°37′E, 16 m, v.1971, D. Simon, 1 ♂ (HUJ INVSC 2274), 1 juv. ♀ (HUJ INVSC 2275).

Buthus arenicola: Simon, 1885: 50–52.

Buthus leptochelys: Pocock, 1895: 299, 300.

Buthacus leptochelys: Birula, 1908: 140 (misidentification).

Buthacus arenicola: Vachon, 1949a: 71; Levy et al., 1973: 125, 128; Levy and Amitai, 1980: 76, 77, 82, 86–89, figs. 79–81, map 6; El-Hennawy, 1987: 17 (misidentification); El-Hennawy, 1992: 97, 112 (part); Fet and Lowe, 2000: 81, 82; Kovařík and Whitman, 2004: 106 (part); Lourenço, 2006: 59, 65, 62–69; Kovařík et al., 2013: 3, fig. 5; Kaltsas et al., 2008: 213; Kamenz and Prendini, 2008: 8, 40, table 2; Badry et al., 2018: 77, 83; Saleh et al., 2017: 7, 8, fig. 3, table 2; Alqahtani and Badry, 2020: 178–183, figs. 1–3, tables 1, 2.

Buthacus spatzi: misidentification: Androctonus (Leiurus) leptochelys paralectotype ♀ (No. 2): Kovařík, 2005: 3, 6, 7, fig. 6; Kovařík et al., 2016: 1.

Type Material: Holotype ♂ (HUJ INVSC 2082), EGYPT: North Sinai Governorate: Quseima, 7.5 km SW, 30°37′N 34°18′E, 236 m, 12.ix.1970, I. Israeli. Paratypes: EGYPT: North Sinai Governorate: Quseima, 7.5 km SW, 30°37′N 34°18′E, 236 m, 12.ix.1970, I. Israeli, 4 ♂ (HUJ INVSC 2077–2079, 2083), 3 ♀ (HUJ INVSC 2076, 2080, 2081), 2 juv. ♂ (HUJ INVSC 2085, 2086), 2 juv. ♀ (HUJ INVSC 2084). Giza Governorate: Abou Rawash, Imbaba, 30°02′32.3″N 31°05′10.2″E, 34 m, 16.xii.2018, T. Adly, 1 ♀(AMCC [LP 16587]). Ismailia Governorate: Qantara, 10 km E, 30°55′N 32°26′E, 4 m, 5.iv.1975, A. Dahlman, 1 ♀ (HUJ INVSC 2371); Timsach Lake [30°35′N 32°18′E], 10 m, 1.iii.1971, Yehezkel, 1 subad. ♂ (HUJ INVSC 2265), 1 juv. ♂(HUJ INVSC 2264), 1 juv. ♀ (HUJ INVSC 2266). Matrouh Governorate: Qesm Ad Dabaah, N of El Dabaa, 31°02′34.7″N 28°26′37.2″E, 23 m, 7. ii.2019, M. Mehana, 3 subad. ♂ (AMCC [LP 16586]); Qesm Ad Dabaah, SW of El Dabaa, 30°58′40.1″N 28°28′33.6″E, 43 m, 25.ii.2019, L. Prendini, E. Attia, T. Saïd and F. Sayed, 2 juv. ♂(AMCC [LP 16601]), 1 juv. ♂, 2 juv. ♀ (AMNH). North Sinai Governorate: Bir Gifgafa, 30°26′N 33°07′E, 313 m, 1968, A. Shulov, 1 juv. ♀ (HUJ INVSC 856), 8.iv.1968, A. Shulov, 2 ♀ (HUJ INVSC 1245, 1251), 15.ii.1973, D. Simon, 1 ♂(HUJ INVSC 2529); Gebel Libri, near road, 30°40′N 33°44′E, 129 m, 15.xi.1967, P. Amitai, 1 juv. ♂ (HUJ INVSC 661); sands between Gebel Ya'laq and Gebel Me'rara [30°33′N 33°26′E], 268 m, 2.ii.1970, J. Gershoni, 1 ♂ (HUJ INVSC 1772), 1 ♀ (HUJ INVSC 1771), 1 juv. ♂ (HUJ INVSC 1773); Qesm Remanah, Om Okba, 30°59′39.1″N 32°46′17.9″E, 17 m, 5.xii.2018, T. Ragab, 1 ♀ (AMCC [LP 16585]); Wadi Masagid, 30°45′N 33°27′E, 274 m, 2.iv.1968, O. Seminar, 1 ♀ (HUJ INVSC 858), 2 juv. ♀ (HUJ INVSC 854, 855). South Sinai Governorate: Wadi Tal, 29°08′N 33°04′E, 131 m, 14.i.1970, M. Broza, 1 juv. ♀(HUJ INVSC 1684); Suez Governorate: E of Suez City, 29°55′N 32°37′E, 16 m, v.1971, D. Simon, 1 ♂ (HUJ INVSC 2273), 1 ♀ (HUJ INVSC 2276), 1 juv. ♂ (HUJ INVSC 2277). ISRAEL: Mehoz HaDarom (Southern District): Holot Haluza sand dunes, E of Be'er Milka, 30°55′42.57″N 34°24′41.95″E, 213 m, 19.ix 2016, B. Shacham, 1 ♀ (HUJ INVSC 3260), 28.ix.2016, S. Talal and S. Cain, 1 ♂ (HUJ INVSC 3253), 2 ♀ (AMCC [LP 15063, 15064] ex HUJ INVSC 3251, 3252), 6 juv. ♂ (AMCC [LP 15065–15067] ex HUJ INVSC 3257–3259, HUJ INVSC 3254–3256), 23.iv.2017, S. Talal and Oranim students, 1 ♀ (AMNH ex HUJ INVSC 3261), 20.iv.2017, Y. Zvik, 1 ♀(AMNH ex HUJ INVSC 3112), 2 subad. ♂(AMNH ex HUJ INVSC 3110, 3111), 25.v.2017, Y. Zvik and S. Cain, 2 subad. ♂ (AMNH ex HUJ INVSC 3268, 3270), 2 juv. ♂ (AMNH ex HUJ INVSC 3265, 3266), 5 juv. ♀ (AMNH ex HUJ INVSC 3263, 3267, 3271, HUJ INVSC 3264, 3269), 31.vii.2017, S. Cain, 2 ♀ (HUJ INVSC 3272, 3274), 4 subad. ♂ (AMNH ex HUJ INVSC 3276, 3278, HUJ INVSC 3275, 3279), 2 subad. ♀(HUJ INVSC 3273, 3281), 1 juv. ♂ (HUJ INVSC 3277), 15.iv.2018, Y. Zvik and S. Cain, 1 ♂ (HUJ INVSC 3382), 2 ♀ (HUJ INVSC 3383, 3386), 2 subad. ♂ (HUJ INVSC 3384, 3385), 1 juv. ♀(HUJ INVSC 3387), 14.vi.2019, D. Kotter, R. Agmon, A. Rinot and S. Cain, 1 ♂ (HUJ INVSC 3669), 1 ♀ (HUJ INVSC 3670), 1 subad. ♂ (HUJ INVSC 3671).

Diagnosis: Buthacus levyi differs from the closely related species, B. arenicola, occurring in Algeria and Tunisia, as follows. The pedipalp chela movable finger of the adult male is proportionally shorter than the manus in B. levyi (fig. 44A, B), with chela manus length:movable finger length, 62.8% (57.4%–67.3%, n = 8; table 10), than in B. arenicola (fig. 38A, B), with chela manus length:movable finger length, 54% (52.4%–55.6%, n = 2; table 10). The metasomal segments of the male are narrower in B. levyi (fig. 28A), with segment width:length, I–V, 65.1% (60.2%–70.8%, n = 8; table 10), 56.9% (53.9%–64.4%), 50.5% (47.6%–57.8%), 40.1% (35.9%–45%), and 33.8% (30.7%–38.9%), than in B. arenicola (fig. 27D), with segment width:length, I–V, 72.2% (70.8%–73.6%, n = 2; table 10), 63.8% (60.9%–66.8%), 58.9% (58.9%–59%), 44.8% (43.1%–46.5%), and 34.5% (34%–35.1%). The dorsolateral carinae of metasomal segment IV are obsolete to absent in B. levyi (figs. 24A, 26A), but well developed in B. arenicola (figs. 23D, 25D).

Buthacus levyi differs from B. amitaii, B. leptochelys and B. nitzani, occurring in Egypt and Israel, as follows. The pedipalp chela of the adult male is longer and narrower in B. levyi (fig. 44A, B), with chela manus length:movable finger length, 62.8% (57.4%–67.3%, n = 8; table 10) and chela manus width:chela length, 19.4% (17.1%–21.3%, n = 8), than in B. amitaii, B. leptochelys, and B. nitzani (figs. 32A, B, 41A, B, 47A, B), with chela manus length:movable finger length, 81.8% (72.7%–88.3%, n = 5; table 3), 70.7% (table 4), and 84.4% (76.8%–92.4%, n = 8; table 10), and chela manus width:chela length, 27.1% (25.1%–28.9%, n = 5), 24.2%, and 26.3% (22.3%–29.5%, n = 8), respectively. The count of retrolateral accessory denticles on the movable finger of the pedipalp chela is usually lower in B. levyi (fig. 21E), with 0–5/0–3 (n = 13; table 5), than in B. amitaii, B. leptochelys, and B. nitzani (fig. 21A, D, G), with 0–9/2–10 (n = 9; table 2), 7–9/7–9 (n = 5; table 5) and 6–9/6–9 (n = 16; table 2), respectively.

Buthacus levyi differs further from B. amitaii and B. nitzani, as follows. The proximal dentate margins of the pedipalp chela fixed and movable fingers are sublinear, such that no gap is evident proximally when the fingers are closed, in B. levyi (fig. 44B), but emarginate, such that a gap is evident proximally when the fingers are closed, in the other species (figs. 32B, 47B). The posterior processes on the ventrolateral carinae of segment V are smaller and more uniform in size in B. levyi (figs. 26A, 28A) than in the other species (figs. 26C, D, 28C, D). The telson dorsal surface is flat and the ventral surface convex and rounded in lateral profile in B. levyi (fig. 29B), whereas the dorsal surface is concave and the ventral surface convex and angular in lateral profile in the other species (fig. 29F, H).

Etymology: The species name is a patronym, honoring Gershom Levy, former Curator of the Arachnida Collection at the National Natural History Collections, Hebrew University of Jerusalem, Israel. Levy promoted the study of arachnids in Israel in particular, and in the Middle East in general; his publications (e.g., Levy and Amitai, 1980) constitute a milestone in understanding the scorpions and other arachnids of the region.

Description: The following description is based on the type material (see table 5 for counts and tables 10 and 11 for measurements).

Total length: Medium-sized scorpions, 49.2 mm (45.4–52.5 mm, n = 8) (♂) or 57.2 mm (53.6–62 mm, n = 7; 1 ♀ with tip of aculeus broken, 67.2 mm) (♀).

Color: Uniformly yellowish olive, except as follows. Carapace interocular surface orange. Mesosomal tergite VII and metasomal segments I–V, each with narrow brown stripe posteriorly. Pectines whitish. Leg articulation points brown. Immatures more orange in color than adults.

Carapace: Carapace shape trapezoidal (fig. 18A, B), anterior width:posterior width, 51.5% (43.9%–60.5%, n = 17), length:posterior width, 90.1% (80.2%–99.8%, n = 17). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with two or, usually, three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 42.5% (40.2%–45.5%, n = 8) (♂) or 43.2% (41.3%–45.3%, n = 9) (♀). Superciliary and central median carinae distinct, costate-granular, and weakly connected to disconnected (♂) or smooth and obsolete (♀). Anteromedian sulcus distinct, shallow; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulci shallow, wide, curved. Carapace intercarinal surfaces finely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin finely granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces setose. Fixed finger dorsal and ventral surfaces densely setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger dorsal surface smooth and glabrous; ventral surface setose; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina comprising discontinuous row of spiniform granules and fewer than 10 macrosetae; dorsal, ventral, and retrolateral intercarinal surfaces smooth; prolateral intercarinal surfaces finely granular (fig. 43A, B). Patella prolateral median and ventral prolateral carinae each comprising discontinuous row of spiniform granules; other carinae absent; intercarinal surfaces smooth (fig. 43C–E). Chela long and slender in both sexes, manus width:length, 52.3% (47.2%–55.9%, n = 8) (♂) or 56% (52.3%–60.3%, n = 9) (♀), manus height:length, 59.7% (55.8%–64%, n = 8) (♂) or 63% (59.5%–66.5%, n = 9) (♀), and manus length:movable finger length, 62.8% (57.4%–67.3%, n = 8) (♂) or 46.6% (43%–52%, n = 9) (♀). Chela manus acarinate; intercarinal surfaces smooth and setose (fig. 44). Fixed and movable fingers respectively with 8–10 (n = 13) and 7–11 (n = 13) (♀) oblique median denticle subrows; movable finger with 0–5 (n = 13) retrolateral accessory denticles (fig. 21E, F); proximal dentate margins of fingers sublinear (fig. 44B), such that small gap present proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae absent; intercarinal surfaces smooth. Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 8:10:12:3; basitarsi, 12:16:16:10; telotarsi, 6:7:8:8 (n = 1). Telotarsal ungues long, approximately equal to telotarsus length, unequal on legs I and II, subequal to equal on III and IV (fig. 22B).

Genital operculum: Genital opercula suboval, completely divided longitudinally, with overlapping, rounded margins (♂) or partially fused longitudinally (♀) (fig. 20E, F). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 11–13 (n = 3) (♂) or 10–13 (n = 3) (♀) median lamellae (fig. 20E, F). Fulcra present. Pectinal teeth along most of length, dentate margin length:pecten length, 97.4% (90.5%–101.2%, n = 8) (♂) or 91.8% (86.4%–96.3%, n = 9) (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 33/32 (31–35/31–34, n = 7) (♂) or 27/27 (25–29/25–28, n = 9) (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 53.3% (44.6%–60.3%, n = 8) (♂) or 53.8% (47.1%–57.5%, n = 9) (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae granular, vestigial, restricted to posterior fifth of I–IV and posterior third of V and VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–III and posterior third of IV–VI. Tergite VII, dorsomedian carina granular, vestigial, restricted to anterior half; dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII smooth and glabrous; III–VI acarinate, VII, ventrolateral carinae vestigial, granular; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 24A, 26A, 28A), segment I shortest, length I:II, 90.8% (85.9%–95%, n = 8) (♂) or 90.4% (82.8%–101%, n = 9) (♀); segments II–IV similar, length II:III, 96.8% (95.3%–100.5%, n = 8) (♂) or 95.7% (86.6%–98.9%, n = 9) (♀), length III:IV, 97.8% (91.2%–108.7%, n = 8) (♂) or 97.8% (95%–100.6%, n = 9) (♀); segment V longest, length IV:V, 83.6% (73.1%–90.8%, n = 8) (♂) or 84.5% (78.2%–91.8%, n = 9) (♀); width:length segment I, 65.1% (60.2%–70.8%, n = 8) (♂) or 66.9% (62.5%–71%, n = 9) (♀), II, 56.9% (53.9%–64.4%, n = 8) (♂) or 56.4% (51.1%–61.6%, n = 9) (♀), III, 50.5% (47.6%–57.8%, n = 8) (♂) or 51.3% (46.2%–56.4%, n = 9) (♀), IV, 40.1% (35.9%–45%, n = 8) (♂) or 41.9% (39.9%–44.1%, n = 9) (♀), V, 33.8% (30.7%–38.9%, n = 8) (♂) or 35.4% (31.8%–40.5%, n = 9) (♀); dorsosubmedian carinae distinct, granular on segment I, obsolete on II and III, absent on IV and V; dorsolateral carinae distinct, granular on segments I and II, obsolete on III and IV, absent on V; dorsosubmedian and dorsolateral carinae sparsely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 1/1 (0/0–2/1, n = 5):3/2 (2/2–4/3):3/2 (2/2–3/3):2/2 (0/1–3/2):0/0 (0/0–0/0), dorsolateral carinae, 2/2 (2/1–2/3, n = 5):3/3 (2/2–3/3):2/3 (1/2–2/3):3/2 (1/1–3/3):3/4 (1/2–4/5). Median lateral carinae distinct, granular, restricted to posterior half of segment I, posterior third of II, vestigial, reduced to posterior margin on III, absent on IV and V. Ventrolateral carinae distinct, costate, becoming granular posteriorly on segments I–IV; serrate, comprising spiniform granules, becoming more prominent posteriorly, on V. Ventrosubmedian carinae distinct, costate on segment I; costate, becoming costate-granular posteriorly, on II and III; costate on IV; obsolete, granular on V. Ventromedian carina obsolete, granular on segment V. Dorsal and lateral intercarinal surfaces smooth on segments I–V; ventral intercarinal surfaces smooth on segments I–IV, finely and sparsely granular on V.

Telson: Telson vesicle width:metasomal segment V width, 72.8% (65.8%–81.9%, n = 8) (♂) or 74.8% (71.7%–80.1%, n = 9) (♀). Vesicle globose, dorsal surface flat, ventral surface convex and rounded; vesicle height:length, 56% (51.6%–59.7%, n = 8) (♂) or 59.3% (55.4%–65%, n = 9) (♀); dorsal and ventral surfaces smooth and glabrous; lateral and ventral surfaces sparsely setose, with 22 (♂: 18–26, n = 2; ♀: 22–23, n = 3) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 52.5% (49.4%–57.8%, n = 8) (♂) or 54.9% (53.1%–58.8%, n = 7) (♀).

Sexual dimorphism: Adult males and females differ as follows. Males are slightly smaller, on average 49.2 mm in total length, than females, on average 57.2 mm. The pedipalp chela of the male has proportionally shorter fingers than that of the female (fig. 44), as indicated by the higher chela manus length:movable finger length ratio in the male (62.8%) compared with the female (46.6%). The mesosoma of the male is relatively narrower than that of the female, as indicated by the higher sternite VII length:width ratio in the male (67.9%) compared with the female (59%). The pectinal tooth count is higher in the male (31–35) than in the female (25–29). The genital opercula are completely divided longitudinally, with rounded margins in the male but partially fused longitudinally in the female (fig. 20E, F), and genital papillae are present in the male but absent in the female.

Distribution: The known locality records suggest B. levyi may be endemic to Egypt and Israel, extending from the coast of northern Egypt, across the sand dunes of the northern and western Sinai Peninsula, to the southern part of the Haluza sand dunes, near the Israeli border with Egypt (fig. 9). Records of B. arenicola from northeastern Libya (Kovařík and Whitman, 2004) may be referable to B. levyi, but this requires confirmation. The known records range from 4 m to 313 m in elevation. Buthacus levyi is allopatric with the closely related B. arenicola, endemic to central Tunisia and northeastern Algeria.

Ecology: The types of B. levyi were collected at night with UV light detection on soft, sparsely vegetated to vegetationless inland sand dunes. The habitat and habitus, notably the pale coloration, smooth tegument, loss or obsolescence of pedipalpal and metasomal carinae, elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins of the tibae and the pro- and retrolateral margins of the basitarsi of legs I–III, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and elongated, unequal length telotarsal ungues, are consistent with the ultrapsammophilous ecomorphotype (Prendini, 2001). Buthacus levyi was collected in sympatry with three other buthids, Androctonus amoreuxi, Buthus israelis, and Orthochirus scrobiculosus, at Be'er Milka, Israel.

Remarks: As noted above, the types of B. arenicola and B. leptochelys were examined during the present investigation, along with material from the vicinity of the type locality of B. arenicola, in Tunisia, and from Egypt, including the Sinai Peninsula, and Israel. These comparisons, multivariate analysis of morphometrics (fig. 15), and multilocus molecular phylogenetics (fig. 14), confirmed previous suggestions (Levy and Amitai, 1980; Lourenço, 2006) that the population of B. arenicola occurring in Egypt and Israel represents a distinct species, described here as Buthacus levyi, sp. nov.

Additionally, as noted above, Kovařík's (2005) conclusion that the type series of B. leptochelys comprises more than one species was also confirmed: paralectotype 2 is heterospecific with the lectotype and paralectotype 1, instead representing a species of the B. arenicola group. However, Kovařík (2005) misidentified paralectotype 2 as the Tunisian species, B. spatzi; it is conspecific with B. levyi, the distribution of which overlaps that of B. leptochelys in the Sinai Peninsula (fig. 6).

Material Examined: EGYPT: Sinai, Androctonus (Leiurus) leptochelys paralectotype ♀ (No. 2) (ZMB 152).

Buthacus leptochelys nitzani Levy et al., 1973: 126; Vachon, 1979: 38, 39; Levy and Amitai, 1980: 83–86, figs. 70–74, map 6; Kinzelbach, 1984: 101 (misidentification); Fet et al., 1998: 616; Kovařík, 1998: 105; Fet and Lowe, 2000: 84; Hendrixson, 2006: 50; Lourenço, 2006: 64; Al-Asmari et al., 2013: 5, 8, 9, fig. 8, table 1 (misidentification).

Buthacus leptochelys: Vachon, 1950b: 406 (part); 1952: 180, 185, 191, 194, 198–203, figs. 262–266 (part); Kinzelbach, 1984: 101; Vachon and Kinzelbach, 1987: 101; El-Hennawy, 1992: 101, 112, 113 (misidentification); Kovařík, 2005: 2, 3; Kaltsas et al., 2008: 214; Lowe et al., 2019: 5.

Type Material: Holotype 1 ♀ (HUJ INVSC 280), ISRAEL: Mehoz HaDarom (Southern District): Haluza sand dunes, Ze'elim [31°12′N 34°32′E], 20.vi.1961, M. Nizan [examined].

Diagnosis: Buthacus nitzani is most closely related to B. amitaii, also occurring in Israel, and B. leptochelys, occurring in Sinai and Egypt. Buthacus nitzani differs from B. amitaii as follows. The metasomal segments of the female are longer and narrower in B. nitzani (fig. 28C), with segment IV width:length, 38.4% (34%–43.7%, n = 10; table 13), than in B. amitaii (fig. 28D), with segment IV width:length, 42.8% (40.2%–44.9%, n = 5; table 3). The ventrosubmedian and ventrolateral carinae are less developed on the metasomal segments, especially on segments II and III, in B. nitzani than in B. amitaii (figs. 26C, D, 28C, D). The posterior processes of the ventrolateral carinae of metasomal segment V are narrow and conical in B. nitzani, but broad and lobate in B. amitaii. The median lateral carinae are obsolete on metasomal segments I–III in B. nitzani, but distinct in B. amitaii. The ventral intercarinal surfaces of metasomal segment V are more finely and sparsely granular in B. nitzani than in B. amitaii. The retrolateral accessory denticles on the movable finger of the pedipalp chela are always present, with constant, usually higher counts (sinistral/dextral: 6–9/6–9; n = 16; table 2; fig. 21G) in B. nitzani, compared to B. amitaii, in which the retrolateral accessory denticles are often absent, with variable, usually lower counts (0–9/2–10; n = 9; table 2; fig. 21A).

Buthacus nitzani differs B. leptochelys as follows. Buthacus nitzani is smaller, measuring 50.6 mm (44.4–58.5 mm, n = 18; tables 12, 13) in total length, than B. leptochelys, measuring 65.8 mm (63.1–68 mm, n = 5; table 4). The pedipalp chela of the adult male is shorter and broader (fig. 47A, B), with chela manus length:movable finger length, 84.4% (76.8%–92.4%, n = 8; table 12) and chela manus width:chela length, 26.3% (22.3%–29.5%, n = 8), in B. nitzani than in B. leptochelys (fig. 41A, B), with chela manus length:movable finger length, 70.7% (table 4) and chela manus width:chela length, 24.2%. The proximal dentate margins of the pedipalp chela fixed and movable fingers are more deeply emarginate, such that a more pronounced gap is evident proximally when the fingers are closed, in B. nitzani (fig. 47B), than in B. leptochelys (fig. 41B). The pectinal tooth counts are lower in B. nitzani, i.e., 31/31 (26–34/27–35, n = 6; table 2) (♂) and 23/23 (20–27/21–27, n = 17) (♀), than in B. leptochelys, i.e., 35/34 (32–39/32–38, n = 6; table 5) (♂) and 29/29 (27–31/28–31, n = 10) (♀). The metasomal segments of the female are shorter and broader in B. nitzani (fig. 28C), with segment IV width:length, 38.4% (34%–43.7%, n = 10; table 13), than in B. leptochelys (fig. 28B), with segment IV width:length, 42.1% (40.3%–44.6%, n = 4; table 4). The median lateral carinae of metasomal segments I–III are obsolete in B. nitzani, but distinct in B. leptochelys. The telson dorsal surface is concave and the ventral surface convex and angular in lateral profile, in B. nitzani (fig. 29F), whereas the dorsal surface is flat and the ventral surface convex and rounded in lateral profile, in B. leptochelys (fig. 29D).

Buthacus nitzani differs from B. levyi, with which it is sympatric in Israel and the Sinai Peninsula, as follows. The pedipalp chela of the adult male is shorter and broader in B. nitzani (fig. 47A, B), with chela manus length:movable finger length, 84.4% (76.8%–92.4%, n = 8; table 12) and chela manus width:chela length, 26.3% (22.3%–29.5%, n = 8), than in B. levyi (fig. 44A, B), with chela manus length:movable finger length, 62.8% (57.4%–67.3%, n = 8; table 10) and chela manus width:chela length, 19.4% (17.1%–21.3%, n = 8). The proximal dentate margins of the pedipalp chela fixed and movable fingers are emarginate, such that a gap is evident proximally when the fingers are closed, in B. nitzani (fig. 47B), but sublinear, such that no gap is evident proximally when the fingers are closed, in B. levyi (fig. 44B). The retrolateral accessory denticles on the movable finger of the pedipalp chela are present, with higher counts, in B. nitzani (sinistral/dextral: 6–9/6–9; n = 16; table 2; fig. 21G), than in B. levyi, in which the denticles are often absent (0–5/0–3; n = 13; table 5; fig. 21E, F). The posterior processes on the ventrolateral carinae of segment V are larger and more variable in size in B. nitzani (figs. 26C, 28C) than in B. levyi (figs. 26A, 28A). The telson dorsal surface is concave and the ventral surface convex and angular in lateral profile, in B. nitzani (fig. 29F), whereas the dorsal surface is flat and the ventral surface convex and rounded in lateral profile, in B. levyi (fig. 29B).

Description: The following description is based on the type material and specimens from across the distribution of the species in Israel (see table 2 for counts and tables 12 and 13 for measurements).

Total length: Medium-sized scorpions, 53 mm (47.4–58.5 mm, n = 8) (♂) or 48.6 mm (44.4–54.5 mm, n = 10) (♀).

Color: Uniformly yellowish olive except as follows. Carapace interocular surface, pedipalp femur and patella, legs, anterior part of metasomal segments I–IV, and entire segment V orange. Pectines pale yellow. Leg articulation points brown.

Carapace: Carapace shape trapezoidal (fig. 17C, D), anterior width:posterior width, 52.8% (47.7%–57.4%, n = 18), length:posterior width, 88.3% (75.2%–98.3%, n = 18). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with two or, usually, three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 39.7% (35.7%–43.1%, n = 8) (♂) or 41.3% (39.2%–44.6%, n = 10) (♀). Superciliary and central median carinae distinct, costate-granular, strongly to weakly connected (♂) or weakly connected to disconnected (♀). Anteromedian sulcus distinct, shallow; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulci shallow, wide, curved. Carapace intercarinal surfaces finely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin finely granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces setose. Fixed finger dorsal and ventral surfaces densely setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger dorsal surface smooth and glabrous; ventral surface setose; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina absent, represented by fewer than 10 macrosetae; intercarinal surfaces smooth (fig. 46A, B). Patella prolateral median and ventral prolateral carinae discontinuous, each comprising a few spiniform granules; other carinae absent; intercarinal surfaces smooth (fig. 46C–E). Chela short and broad (♂), manus width:length, 59% (48.6%–67%, n = 8), manus height:length, 62.5% (57.7%–66.6%, n = 8), and manus length:movable finger length, 84.4% (76.8%–92.4%, n = 8) or short and slender (♀), manus width:length, 54.1% (51.2%–63.1%, n = 10), manus height:length, 61.5% (57.3%–68.5%, n = 10), and manus length:movable finger length, 63.8% (56.4%–83%, n = 10). Chela manus acarinate; intercarinal surfaces smooth and glabrous (fig. 47). Fixed and movable fingers each with 7–10 (n = 16) oblique median denticle subrows; movable finger with 6–9 (n = 16) retrolateral accessory denticles (fig. 21G); proximal dentate margins of fingers emarginate (fig. 47B), such that gap present proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae obsolete; intercarinal surfaces smooth. Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 7:9:11:4; basitarsi, 9:14:17:7; telotarsi, 4:6:7:7 (n = 1). Telotarsal ungues long, approximately equal to telotarsus length, unequal on legs I and II, subequal to equal on III and IV (fig. 22F).

Genital operculum: Genital opercula suboval, completely divided longitudinally, with overlapping, rounded margins (♂) or partially fused longitudinally (♀) (fig. 20A, B). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 7–9 (♂) or 8–10 (n = 3) (♀) median lamellae (fig. 20A, B). Fulcra present. Pectinal teeth along most of length, dentate margin length:pecten length, 96.9% (93.9%–101.6%, n = 8) (♂) or 94.5% (88.7%–99%, n = 10) (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 31/31 (26–34/27–35, n = 6) (♂) or 23/23 (20–27/21–27, n = 17) (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 55.9% (51.9%–61.9%, n = 8) (♂) or 59.7% (52.2%–70.8%, n = 10) (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae granular, vestigial, restricted to posterior fifth of I–V and posterior third of VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–VI. Tergite VII, dorsomedian carina granular, vestigial, restricted to anterior half; dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII smooth and glabrous; III–VI acarinate, VII, ventrolateral carinae vestigial, granular; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 24C, 26C, 28C), segment I shortest, length I:II, 87.7% (83.1%–91.8%, n = 8) (♂) or 89.2% (84.4%–97.8%, n = 10) (♀); segments II–IV similar, length II:III, 98.2% (95.7%–100%, n = 8) (♂) or 98.3% (91.1%–103.1%, n = 10) (♀), length III:IV, 95.4% (93.2%–98.6%, n = 8) (♂) or 95% (91.6%–104.8%, n = 10) (♀); segment V longest, length IV:V, 83.4% (77.6%–88.5%, n = 8) (♂) or 81.7% (77.9%–88.9%, n = 10) (♀); width:length segment I, 64.2% (60.4%–68.8%, n = 8) (♂) or 64.6% (59.1%–67.8%, n = 10) (♀), II, 55.1% (53.3%–58%, n = 8) (♂) or 54.3% (47.9%–61%, n = 10) (♀), III, 51% (49%–54.4%, n = 8) (♂) or 50.8% (46%–56%, n = 10) (♀), IV, 42% (37.9%–50.7%, n = 8) (♂) or 38.4% (33.9%–43.7%, n = 10) (♀), V, 34.1% (30.8%–36.1%, n = 8) (♂) or 34% (31.7%–38.7%, n = 10) (♀); dorsosubmedian carinae distinct, granular on segments I and II, obsolete on III, absent on IV and V; dorsolateral carinae distinct, granular on segment I, granular on II, obsolete on III and IV, absent on V; dorsosubmedian and dorsolateral carinae sparsely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 1/1 (0/0–1/2, n = 4):2/2 (1/1–4/3):2/3 (2/3–3/3):2/1 (1/0–2/3):0/0 (0/0–0/0), dorsolateral carinae, 2/2 (1/1–3/3, n = 4):3/2 (2/2–4/3):3/3 (1/2–4/3):4/3 (4/1–4/4):4/4 (3/3–6/5). Median lateral carinae obsolete, granular along entire length of segment I, restricted to posterior third of II and posterior quarter of III, absent on IV and V. Ventrolateral carinae distinct, costate-granular, restricted to posterior two-thirds of segment I; costate-granular, extending entire length of II–IV; serrate, comprising spiniform granules of variable size, becoming more prominent posteriorly, on V. Ventrosubmedian carinae distinct, costate on segment I; granular, with granules becoming progressively larger and subspiniform posteriorly on II and III; costate-granular along entire length of IV; granular, restricted to anterior three-quarters of V. Ventromedian carina granular, distinct along entire length of segment V. Dorsal and lateral intercarinal surfaces smooth to finely and sparsely granular on segment I, smooth on II–V; ventral intercarinal surfaces smooth on I and III, finely granular on IV, finely and densely granular across entire surface of V.

Telson: Telson vesicle width:metasomal segment V width, 77.2% (71.9%–79.9%, n = 8) (♂) or 79.5% (73.7%–85.6%, n = 10) (♀). Vesicle globose, dorsal surface concave, ventral surface convex and angular; vesicle height:length, 58.5% (44.7%–63.5%, n = 8) (♂) or 65.6% (56.4%–76.8%, n = 10) (♀); dorsal and ventral surfaces smooth and glabrous; lateral and ventral surfaces sparsely setose, with 19 (♂) or 24 (n = 3) (♀) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 50.1% (43.1%–53.6%, n = 8) (♂) or 55% (51%–60.4%, n = 10) (♀).

Sexual dimorphism: Adult males and females differ as follows. The carinae of the pedipalp femur and patella are more coarsely granular in the male than in the female. The pedipalp chela manus of the male is markedly broader and deeper with proportionally shorter fingers, than that of the female (fig. 47), as indicated by the higher ratios of chela manus width:length and manus length:movable finger length, respectively 59% and 84.4% in the male compared with 54.1% and 54.1% in the female. The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female (fig. 20A, B) and genital papillae are present in the male but absent in the female. The pectinal tooth count is higher in the male (26–35) than in the female (22–27). The spiniform granules of the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and the ventrolateral carinae of segment V are less prominent in the male than in the female.

Distribution: The known distribution of B. nitzani extends from the Sorek (Rubin) River, which appears to be the northern limit of the distribution of Buthacus on the Mediterranean coast, throughout the sand dunes of the southern coastal plain of Israel, to the Haluza dunes of the interior (fig. 7). The species probably also occurs in the Gaza Strip and adjacent Sinai Peninsula of Egypt. The known records range from 17 m to 340 m in elevation.

Buthacus nitzani is allopatric with the closely related B. amitaii, endemic to the inland sandy soils and loess of the Yamin Plain and Mamshit area in the northern Negev, Israel (fig. 7), and B. leptochelys, distributed from the western Sinai Peninsula across Egypt, and southward along the coast of the Red Sea to Sudan (fig. 6).

Ecology: Specimens of B. nitzani were collected at night with UV light detection on semiconsolidated to stable yellowish-white, partially vegetated to vegetationless coastal and inland sand dunes. Specimens were usually fairly common, mostly sitting still in open areas, some walking on the ground or resting on bushes. The habitat and habitus, notably the pale coloration, smooth tegument, loss or obsolescence of pedipalpal and metasomal carinae, elongation of the legs, especially legs III and IV, dorsoventral compression of the basitarsi of legs I–III, comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins of the tibae and the pro- and retrolateral margins of the basitarsi of legs I–III, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and elongated, unequal length telotarsal ungues, are consistent with the ultrapsammophilous ecomorphotype (Prendini, 2001).

Buthacus nitzani was collected in sympatry with B. levyi and three other buthids, Androctonus amoreuxi, Buthus israelis, and Orthochirus scrobiculosus at Be'er Milka. In addition to these species, it is sympatric with Androctonus bicolor and Androctonus crassicauda in the Haluza sand dunes and along the coastal plain.

Remarks: According to Levy et al. (1973) and Levy and Amitai (1980), the subspecies B. leptochelys nitzani is restricted to the sand dunes of the central Negev in Israel, and no connection was found to the nominotypical population of B. leptochelys, extending from the Egyptian mainland, across the Sinai Peninsula to the Gaza Strip and southern coastline of Israel. Several authors have suggested that B. l. nitzani is synonymous with other species, however. Kovařík (2005) synonymized B. l. nitzani with the nominotypical form of B. leptochelys, whereas Lourenço (2006) suggested it could be synonymous with B. tadmorensis.

During the present investigation, examination of the types of B. leptochelys and B. l. nitzani, as well as material from Egypt, including the Sinai Peninsula, and Israel, demonstrated that B. nitzani is a valid species, distinct from both B. leptochelys and B. tadmorensis, the latter revalidated in accordance with previous authors, e.g., Birula (1908), Kinzelbach (1985), and Lourenço (2006) (see below). This finding is also strongly supported, for both sexes, by multivariate analysis of morphometrics (fig. 14). Buthacus nitzani Levy et al., 1973, stat. nov., is therefore elevated to the rank of species.

Material Examined: ISRAEL: Mehoz HaDarom (Southern District): Ashdod Nizzanim Nature Reserve, near Nizzan, 31°44′09″N 34°37′11.4″E, 17 m, 29.viii.2011, L. Prendini and T.L. Bird, 5 ♂, 5 ♀(AMNH), 3 juv. ♂, 3 juv. ♀ (AMCC [LP 11174]); Ashdod Nizzanim Nature Reserve, 750 m W of turnoff to Nizzanim Military Base, 31°44′06.7″N 34°37′02.4″E, 17 m, 6.viii.2016, E. Gefen and S. Cain, 1 ♀ (HUJ INVSC3214), 1 subad. ♀(AMCC [LP 15049] ex HUJ INVSC 3213); Ashdod Nizzanim Nature Reserve, dunes N of Nizzan, 31°45′16.0″N 34°38′06.8″E, 39 m, 9.viii.2016, A. Allon, M. Lugassi, S. Livne, and S. Cain, 3 ♂(AMCC [LP 15051, 15052] ex HUJ INVSC 3219, 3220, HUJ INVSC 3215), 3 ♀ (AMCC [LP 15050] ex HUJ INVSC 3218, HUJ INVSC 3216, 3217), 1.vi.2017, S. Cain, 1 ♀ (HUJ INVSC 3208); Ashdod, the Great Dune, N part of Ashdod Nizzanim Nature Reserve, 31°46′18.3″N 34°39′09.7″E, 43 m, 22.viii.2017, I. Ofer and S. Cain, 2 ♂ (AMNH ex HUJ INVSC 3210, 3212), 2 ♀ (AMCC [LP 15048] ex HUJ INVSC 3211, HUJ INVSC 3209); Ashdod, sand dune at corner of Ben Gurion Ave. and Sderot Herzl, 31°48′53.5″N 34°39′11.3″E, 18 m, 22.viii.2017, I. Ofer and S. Cain, 2 ♂ (AMCC [LP 15054] ex HUJ INVSC 3234, HUJ INVSC 3232), 3 ♀ (AMNH ex HUJ INVSC 3231, HUJ INVSC 3230, 3233); Ashqelon, S of Carlsberg Brewery, E of Ha-Tokhen St., 31°37′50.7″N 34°32′46.3″E, 31 m, 23.viii.2017, S. Cain, 3 ♂ (HUJ INVSC 3227–3229), 2 ♀ (AMCC [LP 15056] ex HUJ INVSC 3225, HUJ INVSC 3226); Holot Haluza sand dunes, Gevulot [31°06′N 34°48′E], 329 m, 20.iii.1969, I. Israeli, 1 ♀ (HUJ INVSC 1330); Holot Haluza sand dunes, Revivim, 31°05′N 34°41′E, 252 m, 27.vii.1970, G. Levy, P. Amitai, and A. Shulov, 2 ♂ (HUJ INVSC 1989, 1991), 1 ♀ (HUJ INVSC 1990); Holot Haluza sand dunes, 3.1 km N of turnoff to Retamim on road Revivim–Ze'elim, 31°04′32″N 34°40′33.1″E, 248 m, 27.viii.2011, L. Prendini and T.L. Bird, 15 ♂, 5 ♀ (AMNH), 1 juv. ♂ (AMCC [LP 11175]); Holot Haluza sand dunes, 2.2 km N of turnoff to Retamim on road Revivim–Ze'elim, 31°04′16.5″N 34°41′55.1″E, 259 m, 5.viii.2016, E. Gefen and S. Cain, 5 ♂ (AMCC [LP 15057, 15058] ex HUJ INVSC 3239, 3240, HUJ INVSC 3236–3238), 31.vii.2017, S. Cain, 6 ♂ (AMCC [LP 15059, 15060] ex HUJ INVSC 3241, 3246, AMNH ex HUJ INVSC 3243, HUJ INVSC 3242, 3244, 3245); Holot Haluza sand dunes, 12.5 km N of turnoff to Retamim on road Revivim–Ze'elim, 31°08′10.1″N 34°37′16.0″E, 205 m, 5.viii.2016, E. Gefen and S. Cain, 1 ♂ (AMCC [LP 15053] ex HUJ INVSC 3235); Holot Haluza sand dunes, E of Be'er Milka, 30°55′42.57″N 34°24′41.95″E, 213 m, 23.iv.2017, S. Talal and Oranim students, 2 ♀ (AMCC [LP 15061] ex HUJ INVSC 3247, AMNH ex HUJ INVSC 3262), 25.v.2017, Y. Zvik and S. Cain, 3 ♀(AMCC [LP 15062] ex HUJ INVSC 3250, HUJ INVSC 3248, 3249), 31.vii.2017, S. Cain, 1 ♀ (HUJ INVSC 3280), 15.iv.2018, Y. Zvik and S. Cain, 6 ♀(HUJ INVSC 3388–3392, 3395), 4 subad. ♀ (HUJ INVSC 3393, 3394, 3397, 3398), 2 juv. ♀ (HUJ INVSC 3396, 3399); Holot Haluza sand dunes, 4.5 km S of Ramat Hovav factory, W of road 40, 31°05′32.7″N 34°49′01.6″E, 340 m, 16.iv.2018, Y. Zvik and S. Cain, 7 ♀ (HUJ INVSC 3450–3456), 2 subad. ♀ (HUJ INVSC 3457, 3458), 1 juv. ♂(HUJ INVSC 3461), 2 juv. ♀ (HUJ INVSC 3459, 3460); Mashabbim sand dunes, between Mash'abei Sade and Revivim on road 222, 31°05′00.8″N 34°41′11.0″E, 246 m, 27.viii.2013, L. Prendini, Y. Lubin, E. Gavish-Regev, E. Gefenl, and Israel Taxonomy Initiative scorpion course participants, 24 ♂, 1 ♀ (AMNH), 1 juv. ♀ (AMCC [LP 12200]); Palmahim sand dunes, E of Palmahim air base, 31°54′39.4″N 34°43′31.8″E, 19 m, 22.viii.2017, I. Ofer, N. Braun and S. Cain, 2 ♂ (AMCC [LP 15055] ex HUJ INVSC 3222, HUJ INVSC 3224), 2 ♀ (AMNH ex HUJ INVSC 3221, HUJ INVSC 3223).

Figures 10, 15, 16C, D, 19C, D, 21H, 22E, 23B, 25B, 27B, 29E, 48, 49, 50; tables 1, 4, 9

Buthus tadmorensis Simon, 1892: 84; Kraepelin, 1895: 82; Birula, 1905: 136–138; Werner, 1936b: 204.

Buthacus leptochelys: Kraepelin, 1899: 17; Pringle, 1960: 76, fig. 2; L. Khalaf, 1962: 1, 2; K.I. Khalaf, 1963: 59; Levy et al., 1973: 126; Levy and Amitai, 1980: 77, 82; Fet and Lowe, 2000: 82; Dehghani and Kassiri, 2018: S881.

Buthus (Buthacus) tadmorensis: Birula, 1908: 140; 1910: 172, 173; 1917: 23, 214, 229.

Buthus pietschmanni Penther, 1912: 112–114, fig.; Vachon, 1966: 211; Kachel et al., 2021: 2, 3; syn. nov.

Buthus (Buthacus) leptochelys: Whittick, 1947: 122; 1955: [unpaginated] (part).

Buthacus tadmorensis: Vachon, 1948: 475; 1966: 210; Habibi, 1971: 43; Pérez, 1974: 19; Kinzelbach, 1984: 101; 1985: map III; El-Hennawy, 1992: 101, 114 (part); Kovařík, 1997a: 49; 1998: 105; Kovařík and Whitman, 2004: 106 (misidentification, part); Lourenço, 2006: 63, 64, 69, figs. 17, 18, 42; Lourenço and Qi, 2006a: 161, 162; Lourenço and Leguin, 2009: 103, 104; Shehab et al., 2011: 333–340, fig. 2E, tables 1, 2; Dehghani and Kassiri, 2018: S881; Lowe et al., 2019: 5.

Buthacus yotvatensis: Levy et al., 1973: 134 (misidentification).

Buthacus tadmorensis tadmorensis: Kinzelbach, 1985: map III; Kovařík, 2002: 5.

Mesobuthus pietschmanni: El-Hennawy, 1992: 101, 128.

Buthacus tadmorensis yotvatensis: El-Hennawy, 1992: 101, 114 (misidentification); Kabakibi et al., 1999: 82, 88 (misidentification).

Buthacus macrocentrus: Kovařík, 2005: 1, 7; Lourenço and Qi, 2006a: 161; Kaltsas et al., 2008: 214; Navidpour et al., 2008: 1, 3–8, 28, 30, 35, figs. 3, 6, 12, 56–59; Yağmur et al., 2008: 13–17, figs. 2, 4; Mirshamsi et al., 2011: 17, 24, 26, fig. 1A, table 1; Navidpour et al., 2013: 1, 3–5, 21, figs. 2, 4; Loria and Prendini, 2014: 9, 10, fig. 2C; Caliskan, 2015: 327–332, 336, 337, 343, figs. 1, 3, 5B; Navidpour, 2015: 10, 11; Kovařík et al., 2016: 1; Sharifinia et al., 2017: 232, 233, 235, 236, table 2; Dehghani and Kassiri, 2018: S881; Al-Khazali and Yağmur, 2019: 85–87, fig. 2C (misidentification); Koç et al., 2019: 108–112, figs. 1, 5–7, table 1; Hussen and Ahmed, 2020: 6711–6713, 6715, 6716, 6718, 6719, 6722, 6723, figs. 1, 7, tables 1, 2; Alqahtani and Badry, 2021: 10; Amr et al., 2021: 85, 87, 89, 90, 97, tables 3, 10; Mansouri et al., 2021: 763, 765, 767, table 2; Kachel et al., 2021: 2, 3, 5, 7, table 1.

Buthacus yotvatensis: Crucitti and Vignoli, 2002: 438–440, fig. 3 (misidentification).

Type Material: Buthus pietschmanni: IRAQ: Saladin Governorate: Lectotype ♂, 1 ♀, 1 juv. ♂, 2 juv. ♀ paralectotypes (NHMW 2453), Mesopotamien, Kal'at Shergat [Qal'at Sherqat, 35°27′N 43°16′E], 11.v.1910, Pietschmann [examined]; 3 ♀, 1 subad. ♀, 3 juv. ♀ paralectotypes (NHMW 2452), Mesopotamien, Assur [Aššur/Ashur, 35°27′N 43°16′E], 19.v.1910, Pietschmann [examined]. Buthus tadmorensis: 1 ♂, 1 ♀ syn-types (MNHN), SYRIA: Homs Governorate: Palmira [Palmyra/Tadmur/Tadmor, 34°34′N 38°17′E].

Diagnosis: Buthacus tadmorensis differs from the closely related B. arava and B. yotvatensis, occurring in Israel and Jordan, as follows. The ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and ventrolateral carinae of segment V are well developed, with broad, lobate processes posteriorly, in B. tadmorensis, but weakly developed, with narrow, spiniform processes posteriorly, in the other species (figs. 25A, B, C, 27A, B, C).

Buthacus tadmorensis differs further from B. arava, as follows. The pedipalp chela of B. tadmorensis is longer and narrower, especially in the adult male (fig. 50A, B), with chela manus width:chela length, 22.3% (♂) (table 4), chela manus length:movable finger length, 64.2% (♂) or 55% (51.9%–59.8%, n = 4) (♀), and chela manus width:length, 58.7% (♂) or 59.1% (53.8%–65.1%, n = 4) (♀), than that of B. arava (fig. 35A, B), with chela manus width:chela length, 29.8% (27.2%–32.8%, n = 7; table 8) (♂), chela manus length:movable finger length, 90.4% (83.5%–95%, n = 7) or 60.8% (♀), and chela manus width:length, 63.3% (57.3%–74.5%, n = 7) or 61.6% (♀). Additionally, the ungues of the leg telotarsi are shorter (approximately two-thirds the length of the telotarsus) and equal on legs I and II in B. tadmorensis (fig. 22E) but longer (approximately equal to the length of the telotarsus) and unequal on legs I and II in B. arava (fig. 22C). The pectinal tooth counts are higher in B. tadmorensis, i.e., 30/32 (♂) (table 9) and 24/25 (23–26/23–26, n = 3) (♀), than in B. arava, i.e., 20/20 (18–21/18–22, n = 7; table 9) (♂) and 12/13 (♀). The metasomal segments and telson are densely setose in B. tadmorensis (figs. 23B, 25B, 29E), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 10/11 (8–12/10–12, n = 2):19/23 (18–19/21–24):23/23 (23–23/21–25):19/20 (17–20/19–21):9/9 (7–11/7–11), dorsolateral carinae, 6/6 (4–8/5–6):11/13 (8–13/11–15):9/11 (8–9/10–11):9/9 (8–9/6–12):9/9 (8–10/8–10), and telson, 100 (80–120, n = 2), but sparsely setose in B. arava (figs. 23A, 25A, 29C), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 0/0 (0–1/0–1, n = 4):2/3 (2–3/2–5):4/3 (2–5/2–6):3/3 (2–3/1–5):0/0 (0–0/0–0), dorsolateral carinae, 1/1 (0–2/0–2):2/2 (1–3/0–3):3/2 (2–4/1–3):3/3 (2–4/2–5):5/6 (3–8/5–7), and telson, 28 (27–29, n = 4).

Buthacus tadmorensis differs further from B. yotvatensus, as follows. The pedipalp chela of B. tadmorensis is shorter and broader (fig. 50A, B), with chela manus width:chela length, 22.3% (♂) (table 4), chela manus length:movable finger length, 64.2% (♂) or 55% (51.9%–59.8%, n = 4) (♀), and chela manus width:length, 58.7% (♂) or 59.1% (53.8%–65.1%, n = 4) (♀), than that of B. yotvatensis (fig. 53A, B), with chela manus width:chela length, 17.4% (16.5%–18.8%, n = 8; table 6) (♂), chela manus length:movable finger length, 55.2% (51.7%–58.3%, n = 8) (♂) or 47.1% (45.5%–48%, n = 6; table 7) (♀), and chela manus width:length, 50.5% (48.1%–54.8%, n = 8) (♂) or 56.9% (51.3%–61.5%, n = 6) (♀). The telson ventral surface is densely granular anteriorly in B. tadmorensis but sparsely granular anteriorly in B. yotvatensis (fig. 29E, G).

Description: The following description is based on the type material of B. pietschmanni and additional material examined (see table 4 for measurements and table 9 for counts).

Total length: Large scorpions, 66.8 mm (♂) or 70.2 mm (66.3–75.2 mm) (♀).

Color: Uniformly yellowish olive with pale yellow legs and whitish pectines.

Carapace: Carapace shape trapezoidal (fig. 16C, D), anterior width:posterior width, 47.8% (44%–53.1%, n = 5), length:posterior width, 86.7% (82.2%–92.8%, n = 5). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 48% (♂) or 47.1% (45.2%–49.4%, n = 4) (♀). Superciliary and central median carinae distinct, costate-granular and disconnected to weakly connected. Anteromedian sulcus distinct, deep; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulcus deep, wide, curved. Carapace intercarinal surfaces coarsely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin finely granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces setose. Fixed finger dorsal surface setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger smooth and glabrous; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina obsolete, comprising discontinuous row of spiniform granules and more than 15 macrosetae; dorsal, prolateral, and ventral intercarinal surfaces finely granular; retrolateral intercarinal surfaces smooth (fig. 49A, B). Patella prolateral median and ventral prolateral carinae complete, costate-granular to finely serrate; dorsal prolateral carinae complete, costate-granular; other carinae obsolete; intercarinal surfaces smooth (fig. 49C–E). Chela long and slender in both sexes, manus width:length, 58.7% (♂) or 59.1% (53.8%–65.1%, n = 4) (♀), manus height:length, 67.4% (♂) or 67.7% (59.2%–70.8%, n = 4) (♀), and manus length:movable finger length, 64.2% (♂) or 55% (51.9%–59.8%, n = 4) (♀). Chela manus acarinate; intercarinal surfaces smooth and setose (fig. 50). Fixed and movable fingers respectively with 8–10 (n = 5) and 9–11 (n = 5) oblique median denticle subrows; movable finger with 8–10 (n = 5) retrolateral accessory denticles (fig. 21H); proximal dentate margins of fingers sublinear (fig. 50B), such that no gap evident proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae obsolete; intercarinal surfaces smooth. Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 9:13:12:5; basitarsi, 13:18:21:8; telotarsi, 7:7:6:5 (n = 1). Telotarsi ungues short, approximately equal to three-quarters of telotarsus length, equal on legs I–IV (fig. 22E).

Genital operculum: Genital opercula suboval, completely divided longitudinally, without overlapping, rounded margins (♂) or partially fused longitudinally (♀) (fig. 19C, D). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 12 (♂) or 9–12 (n = 4) (♀) median lamellae (fig. 19C, D). Fulcra present. Pectinal teeth along part of length, dentate margin length:pecten length, 89.7% (♂) or 84.4% (81.5%–87.7%, n = 4) (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 30/32 (♂) or 24/25 (23–26/23–26, n = 3) (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 56.5% (♂) or 60.7% (55.3%–65.1%, n = 4) (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly finely granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae smooth, vestigial, restricted to posterior half of I, granular, complete on II–VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–V and posterior half of VI. Tergite VII, dorsomedian carina obsolete, granular, dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII smooth and glabrous; III–VI acarinate, VII, ventrosubmedian and ventrolateral carinae obsolete, smooth; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (fig. 23B, 25B, 27B), segment I shortest, length I:II, 90.5% (♂) or 92.2% (89.9%–95.3%, n = 4) (♀); segments II and III similar, length II:III, 100.5% (♂) or 97.6% (97.1%–98%, n = 4) (♀); segments IV longer, length III:IV, 92.4% (♂) or 94.4% (92.7%–96.7%, n = 4) (♀); segment V longest, length IV:V, 94.7% (♂) or 88.9% (86.7%–90.2%, n = 4) (♀); width:length segment I, 70.5% (♂) or 71.4% (67.8%–76.2%, n = 4) (♀), II, 60.1% (♂) or 62.1% (60.1%–64.7%, n = 4) (♀), III, 57.9% (♂) or 58.6% (55.2%–63.1%, n = 4) (♀), IV, 48.3% (♂) or 49.2% (48.9%–49.4%, n = 4) (♀), V, 42.1% (♂) or 42.7% (40.9%–43.9%, n = 4) (♀); dorsosubmedian and dorsolateral carinae distinct, granular on segments I–III, obsolete on IV, absent on V; dorsosubmedian and dorsolateral carinae densely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 10/11 (8/10–12/12, n = 2):19/23 (18/21–19/24):23/23 (23/21–23/25):19/20 (17/19–20/21):9/9 (7/7–11/11), dorsolateral carinae, 6/6 (4/5–8/6, n = 2):11/13 (8/11–13/15):9/11 (8/10–9/11):9/9 (8/6–9/12):9/9 (8/8–10/10). Median lateral carinae distinct, granular in posterior two-thirds of segment I, posterior half of II and posterior third of III, absent on IV and V. Ventrolateral carinae distinct, weakly granular on segment I; costate-granular to serrate, with granules becoming progressively larger and spiniform posteriorly, on II and III; costate-granular on IV; serrate, comprising spiniform granules of variable size and lobate processes posteriorly, on V. Ventrosubmedian carinae distinct, weakly granular on segment I, costate-granular to serrate, with granules becoming progressively larger and spiniform to lobate posteriorly, on II and III; costate-granular on IV, granular, restricted to anterior three-quarters of V. Ventromedian carina granular, distinct along entire length of segment V. Dorsal and lateral intercarinal surfaces matte and punctate on segments I–V; ventral intercarinal surfaces smooth on I and II, finely granular across entire surface on III–V.

Telson: Telson vesicle width:metasomal segment V width, 76.1% (♂) or 86.2% (83.4%–88%, n = 4) (♀). Vesicle globose, dorsal surface flat, ventral surface convex and rounded; vesicle height:length, 67.1% (♂) or 72.9% (65.4%–79.2%, n = 4) (♀); dorsal surface smooth; entire ventral surface finely granular; lateral and ventral surfaces densely setose, with 80 (♂) or 120 (♀) (n = 2) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 47.2% (♂) or 51.7% (50.2%–53.7%, n = 4) (♀).

Sexual dimorphism: Adult males and females differ as follows. The carinae of the pedipalp femur and patella are more coarsely granular in the male than in the female. The pedipalp chela of the male has proportionally shorter fingers than that of the female (fig. 50), as indicated by the higher chela manus length:movable finger length ratio in the male (64.2%) compared with the female (55%). The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female (fig. 19C, D) and genital papillae are present in the male but absent in the female. The pectinal tooth count is higher in the male (30–32) than in the female (23–26). The spiniform granules of the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and the ventrolateral carinae of segment V are less prominent in the male than in the female.

Distribution: Buthacus tadmorensis is widely distributed across the Fertile Crescent of the Middle East (fig. 10) from central Jordan, through Syria, southeastern Turkey and Iraq, to the northern shore of the Persian Gulf, in southern Iran (Simon, 1892; Penther, 1912; El-Hennawy, 1992; Kabakibi et al., 1999; Crucitti and Vignoli, 2002; Kovařík and Whitman, 2004; Kovařík, 2005; Yağmur et al., 2008; Navidpour et al., 2008; Shehab et al., 2011; Sharifinia et al., 2017). The known distribution covers most of the drainage of the Tigris and Euphrates rivers. Records are located at elevations ranging from 13 m to 924 m (Navidpour et al., 2008; Shehab et al., 2011; Sharifinia et al., 2017).

Buthacus tadmorensis is allopatric with the closely related B. arava and B. yotvatensis, both endemic to the Arava Valley, straddling the southern border between Israel and Jordan (figs. 4, 5), and, in the case of B. yotvatensis, extending west into the Negev.

Ecology: Buthacus tadmorensis inhabits loose to hard, compacted sandy habitats (Shehab et al., 2011). Specimens have been collected under stones or in pitfall traps on sparsely vegetated plains and wadis with hard soil and few to many stones (Shehab et al., 2011). The habitat and habitus, e.g., the pale coloration, lobate granules and processes posteriorly on the ventrosubmedian and ventrolateral carinae of metasomal segments II, III and V, dorsoventral compression of the basitarsi of legs I–III, with weakly developed comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins, are consistent with the semipsammophilous ecomorphotype (Prendini, 2001).

Other scorpion species recorded in sympatry with B. tadmorensis include the buthids Androctonus crassicauda, Apistobuthus susanae Lourenço, 1998, Compsobuthus jakesi Kovařík, 2003, Hottentotta saulcyi (Simon, 1880), Leiurus hebraeus (Birula, 1908), Mesobuthus phillipsii (Pocock, 1889), Odontobuthus bidentatus Lourenço and Pézier, 2002, Orthochirus iranus Kovařík, 2004, Orthochirus scrobiculosus, Razianus zarudnyi (Birula, 1903); the hemiscorpiid Hemiscorpius lepturus Peters 1861; and the scorpionids Scorpio kruglovi Birula, 1910 and Scorpio townsendi (Pocock, 1900) (Navidpour et al., 2008; Shehab et al., 2011; Sharifinia et al., 2017).

Remarks: Buthacus tadmorensis, originally described from Palmyra, Syria (Simon, 1892), was synonymized with B. leptochelys by Kraepelin (1899), but incorrectly attributed to Levy et al. (1973) by Fet and Lowe (2000). Others (e.g., Birula, 1905, 1908, 1910, 1917; Werner, 1936; Vachon, 1948, 1966; Habibi, 1971) continued to recognize B. tadmorensis, however. Pérez (1974) subsequently synonymized Buthus pietschmanni, described from Qal'at Sherqat [Aššur, Iraq], with B. tadmorensis. Kinzelbach (1985) relegated B. yotvatensis and B. yotvatensis nigroaculeatus Levy et al., 1973, to subspecies of B. tadmorensis, a decision accepted by some (e.g., Vachon and Kinzelbach, 1987; Amr et al., 1988; El-Hennawy, 1992) and rejected by others (e.g., Fet and Lowe, 2000; Lourenço, 2004a).

Kovařík (2005) subsequently revalidated B. macrocentrus, originally synonymized with B. leptochelys by Kraepelin (1891), and synonymized B. tadmorensis, B. pietschmanni, and B. yotvatensis with it. This decision was accepted uncritically by most subsequent workers (e.g., Kaltsas et al., 2008; Navidpour et al., 2008; 2013; Navidpour, 2015; Sharifinia et al., 2017: Dehghani and Kassiri, 2018; Al-Khazali and Yağmur, 2019; Koç et al., 2019; Hussen and Ahmed, 2020; Amr et al., 2021; Kachel et al., 2021).

After comparing two specimens from Palmyra, the type locality of B. tadmorensis, presumed to be part of the original type series, with a paratype of B. yotvatensis, deposited at the MNHN, Lourenço (2006) suggested that B. tadmorensis is a valid species whereas B. yotvatensis could be a junior synonym of B. macrocentrus. Lourenço (2006: 64) noted “the very developed ventral carinae of metasomal segments II and III” in B. tadmorensis, a character originally described by Simon (1892: 84) as “segmentis caudae 2° et 3° carinis inferioribus apicem versus sensim validioribus et dente apicali reliquis majoribus,” and which is absent in the types of B. macrocentrus and B. yotvatensis. Nevertheless, Kovařík et al. (2016) and others (Kaltsas et al., 2008; Navidpour et al., 2008; 2013; Navidpour, 2015; Kachel et al., 2021) continued to regard B. tadmorensis, B. pietschmanni, and B. yotvatensis as junior synonyms of B. macrocentrus. Moreover, Lowe et al. (2019) recently rejected Lourenço's (2006) assessment of the validity of B. tadmorensis, along with the suggestion that Buthacus buettikeri could be a junior synonym of the latter. According to Lowe et al. (2019: 5):

Examination of the types of B. leptochelys and B. macrocentrus for the present investigation demonstrated that they are conspecific, as originally concluded by Kraepelin (1891), requiring B. macrocentrus to be returned to synonymy with B. leptochelys (see above). In addition, examination of the types of B. pietschmanni, as well as conspecific material from Jordan, Turkey, Iraq, and Iran, closely matching the descriptions of the type specimens from Palmyra by Simon (1892) and the probable type specimens, also from Palmyra, described by Lourenço (2006), demonstrated that B. tadmorensis is a valid species and B. pietschmanni a junior synonym thereof. This finding is supported by multivariate analysis of morphometrics (fig. 15) as well as multilocus phylogenetic analysis (L.P., unpublished data). Buthacus tadmorensis (Simon, 1892), stat. rev., is revalidated accordingly and B. pietschmanni returned to synonymy with it: Buthus pietschmanni Penther, 1912 = Buthacus tadmorensis (Simon, 1892), syn. nov. On the contrary, examination of the holotype of B. yotvatensis as well as material from across the distributions of both species, multivariate analysis of morphometrics (fig. 15), and multilocus phylogenetic analysis (fig. 14), confirmed that B. yotvatensis is a valid species, as originally indicated by Levy et al. (1973). Furthermore, it would appear, based on the evidence available, that B. buettikeri is a valid species, regardless of its generic assignment. Material conspecific with B. buettikeri, from al-Mudawwarah, Jordan, slightly north of the border with Saudi Arabia, was collected by the last author and deposited in the AMNH, confirming the validity of the record doubted by Lowe et al. (2019).

Material Examined: IRAN: Bushehr Province: Delvar, 28°42′59″N 51°04′52″E, 4 m, xi.2007, Masihipour and Hayader, 1 ♂ (AMCC [LP 11060]); Omidiyeh to Genaveh road, 30°13′42″N 50°12′01″E, 128 m, vi.2007, S. Navidpour and B. Masihipour, 1 ♂, 1 juv. ♀ (AMCC [LP 9720]). Ilam Province: Dashte Abbas, Ein Saleh village, 32°25′24″N 47°43′86″E, 182 m, x.2007, S. Navidpour, 2 juv. ♂ (AMCC [LP 9721, 11076]). Khoozestan Province: Ahvaz–Omidiyeh road (40 km to Omidiyeh), 30°37′49″N 49°31′47″E, 79 m, v.2007, B. Masihipour and H. Bahrani, 1 ♂ (AMCC [LP 9719]); Shush (Apadana Palace), 32°10′55″N 48°15′39″E, 75 m, x.2007, S. Navidpour, B. Masihipour and H. Bahrani, 1 ♀, 1 subad. ♂ (AMCC [LP 9722]). IRAQ: Erbil Governorate: Bardber, 36°02′50.8″N 44°19′20.5″E, 14.vii.2019, F.S. Hussen, 1 ♀(AMCC [LP 16876]). JORDAN: Dieban District: Wadi Deb'em [31°31′N 35°49′E], SE of Amman, 709 m, vii.1938, O. Theodor, 2 juv. ♀ (HUJ INVSC 2491). TURKEY: Şanlıurfa (Urfa) Province: Birecik, ca. 20 km E, 37°00′38.4″N 38°11′41.1″E, 686 m, 23.v.2007, A.V. Gromov and E.A. Yağmur, 2 ♀, 2 subad. ♂, 1 subad. ♀(AMNH), 2 juv. (AMCC [LP 7355]); Harran ruins, 36°51′50.5″N 39°01′42.5″E, 358 m, 17.v.2007, A.V. Gromov, H. Koç and E.A. Yağmur, 56 ♀, 52 subad. ♂, 46 subad. ♀, 17 juv. ♂, 79 juv. ♀ (AMNH), 2 juv. (AMCC [LP 7357]).

Buthacus yotvatensis Levy et al., 1973: 130–133, 134 (misidentification, part), figs. 32–37; Levy and Amitai, 1980: 90–93, figs. 82–85, map 6; Kinzelbach, 1984: 99, 101 (misidentification); Fet et al., 1998: 615, 616; Fet and Lowe, 2000: 85; Soleglad and Fet, 2003a: 5 (misidentification); 2003b: 7, 149, 151, 152, 156, fig. B1-2 (misidentification); Fet et al., 2003: 3, 4, table 1 (misidentification); Hendrixson, 2006: 47, 52, 54, 56; Amr et al., 2015: 30–33, fig. 1D; Alqahtani et al., 2019: 19, 22, 24, 25, fig. 2C (misidentification); Alqahtani and Badry, 2021: 4, 10, table 1; Amr et al., 2021: 86, 88, fig. 3A, table 4; Kachel et al., 2021: 2, 3.

Buthacus yotvatensis yotvatensis: Vachon, 1979: 36; Fet and Lowe, 2000: 85; Lourenço, 2004a: 206; Hendrixson, 2006: 55.

Buthacus tadmorensis yotvatensis: Kinzelbach, 1985: map III; Vachon and Kinzelbach, 1987: 101; Amr et al., 1988: 374; El-Hennawy, 1992: 101, 114 (part).

Buthacus tadmorensis: Vachon, 1952: 180 (misidentification); El-Hennawy, 1992: 101, 104 (misidentification); Kovařík, 2001: 80.

Buthacus macrocentrus: Kovařík, 2005: 1, 8; Lourenço, 2006: 64; Kaltsas et al., 2008: 214.

Type Material: Holotype 1 ♂ (SMNH TAU AR 5223 old NS 5223), ISRAEL: Mehoz HaDarom (Southern District): Arava Valley, dunes E of Yotvata [29°53′N 35°04′E], 2.v.1965, H. Zinner [examined].

Diagnosis: Buthacus yotvatensis differs from the closely related B. arava, also occurring in Israel and Jordan, as follows. The pedipalp chela of B. yotvatensis is longer and narrower, especially in the adult male (fig. 53A, B), with chela manus width:chela length, 17.4% (16.5%–18.8%, n = 8; table 6) (♂), chela manus length:movable finger length, 55.2% (51.7%–58.3%, n = 8) (♂) or 47.1% (45.5%–48%, n = 6, table 7) (♀), and chela manus width:length, 50.5% (48.1%–54.8%, n = 8) (♂) or 56.9% (51.3%–61.5%, n = 6) (♀), than that of B. arava (fig. 35A, B), with chela manus width:chela length, 29.8% (27.2%–32.8%, n = 7; table 8) (♂), chela manus length:movable finger length, 90.4% (83.5%–95%, n = 7) or 60.8% (♀), and chela manus width:length, 63.3% (57.3%–74.5%, n = 7) or 61.6% (♀). The ungues of the leg telotarsi are shorter (approximately two-thirds the length of the telotarsus) and equal on legs I and II in B. yotvatensis (fig. 22G) but longer (approximately equal to the length of the telotarsus) and unequal on legs I and II in B. arava (fig. 22C). The pectinal tooth counts are higher in B. yotvatensis, i.e., 34/35 (32–37/32–37, n = 8; table 9) (♂) and 26/26 (25–28/26–28, n = 6) (♀), than in B. arava, i.e., 20/20 (18–21/18–22, n = 7; table 9) (♂) and 12/13 (♀). The metasomal segments and telson are densely setose in B. yotvatensis (figs. 23C, 25C, 29G), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 16/15 (14–20/11–21, n = 5):25/26 (20–29/18–30):26/25 (23–31/19–29):30/27 (24–38/23–31):12/11 (6–18/7–15), dorsolateral carinae, 9/8 (7–13/5–13):20/20 (19–23/16–21):22/21 (16–28/18–24):15/14 (11–20/12–17):11/13 (10–13/11–14), and telson, 102 (88–127, n = 5), but sparsely setose in B. arava (figs. 23A, 25A, 29C), with macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 0/0 (0–1/0–1, n = 4):2/3 (2–3/2–5):4/3 (2–5/2–6):3/3 (2–3/1–5):0/0 (0–0/0–0), dorsolateral carinae, 1/1 (0–2/0–2):2/2 (1–3/0–3):3/2 (2–4/1–3):3/3 (2–4/2–5):5/6 (3–8/5–7), and telson, 28 (27–29, n = 4).

Buthacus yotvatensis differs from the closely related B. tadmorensis, occurring in neighboring countries, as follows. The pedipalp chela of B. yotvatensis is longer and narrower (fig. 53A, B), with chela manus width:chela length, 17.4% (16.5%–18.8%, n = 8; table 6) (♂), chela manus length:movable finger length, 55.2% (51.7%–58.3%, n = 8) (♂) or 47.1% (45.5%–48%, n = 6; table 7) (♀), and chela manus width:length, 50.5% (48.1%–54.8%, n = 8) (♂) or 56.9% (51.3%–61.5%, n = 6) (♀), than that of B. tadmorensis (fig. 50A, B), with chela manus width:chela length, 22.3% (♂) (table 4), chela manus length:movable finger length, 64.2% (♂) or 55% (51.9%–59.8%, n = 4) (♀), and chela manus width:length, 58.7% (♂) or 59.1% (53.8%–65.1%, n = 4) (♀). The ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and ventrolateral carinae of segment V are less developed in B. yotvatensis than in B. tadmorensis (fig. 25B, C), with narrow, spiniform processes posteriorly in B. yotvatensis but broad, lobate processes posteriorly in B. tadmorensis. The telson ventral surface is sparsely granular anteriorly in B. yotvatensis but densely granular anteriorly in B. tadmorensis (fig. 29E, G).

Description: The following description is based on specimens from across the distribution of the species in Israel (see tables 6 and 7 for measurements, and table 9 for counts).

Total length: Large scorpions, 67.4 mm (61.9–71.9 mm, n = 8) (♂) or 76.6 mm (71.6–85 mm, n = 6) (♀).

Color: Uniformly pale yellow to yellowish olive, except as follows. Mesosomal tergites, sternite VII and metasomal segments I–V, each with narrow brown stripe posteriorly. Pectines pale yellow. Leg articulation points brown.

Carapace: Carapace shape trapezoidal (fig. 16C, D), anterior width:posterior width, 53.3% (46.3%–59.2%, n = 14), length:posterior width, 89.4% (81.7%–98.4%, n = 14). Five, rarely four, pairs of lateral ocelli; each lateral ocular tubercle with three major ocelli (ALMa, MLMa, PLMa), similar in size, situated anterolaterally, and one or, usually, two minor ocelli (ADMi, PDMi; PDMi may be absent) situated posterodorsal to posterior major ocellus. Median ocelli larger than lateral ocelli, distance between them more than 2× ocellus width. Median ocular tubercle situated anteromedially, distance from anterior carapace margin:carapace length, 45.4% (42.6%–47%, n = 8) (♂) or 45.5% (43.9%–46.8%, n = 6) (♀). Superciliary and central median carinae distinct, costate-granular and weakly connected to disconnected. Anteromedian sulcus distinct; posteromedian sulcus deep, narrow anteriorly, wide posteriorly; posterolateral sulcus deep, wide, curved. Carapace intercarinal surfaces coarsely and densely granular.

Chelicerae: Cheliceral manus prodorsal margin finely granular; retrodorsal surfaces smooth or finely granular; prolateral and ventral surfaces setose. Fixed finger dorsal surface setose; dorsal margin with subdistal, medial, and proximal denticles; ventral margin with proximal and medial denticles. Movable finger smooth and glabrous; dorsal margin with retrodistal, subdistal, medial, and pair of proximal denticles; ventral margin with prodistal, medial, and proximal denticles.

Pedipalps: Femur dorsal prolateral, dorsal retrolateral and ventral prolateral carinae complete, costate-granular; prolateral ventral and prolateral ventrosubmedian carinae each comprising discontinuous row of spiniform granules; retrolateral dorsosubmedian carina obsolete, comprising discontinuous row of spiniform granules and more than 15 macrosetae; dorsal, prolateral and ventral intercarinal surfaces finely granular; retrolateral intercarinal surfaces smooth (fig. 52A, B). Patella prolateral median and ventral prolateral carinae complete, costate-granular to finely serrate; dorsal prolateral carinae complete, costate-granular; other carinae obsolete; intercarinal surfaces smooth (fig. 52C–E). Chela long and slender in both sexes, manus width:length, 50.5% (48.1%–54.8%, n = 8) (♂) or 56.9% (51.3%–61.5%, n = 6) (♀), manus height:length, 56.9% (53.3%–60.7%, n = 8) (♂) or 63.5% (60.1%–67.2%, n = 6) (♀), and manus length:movable finger length, 55.2% (51.7%–58.3%, n = 8) (♂) or 47.1% (45.5%–48%, n = 6) (♀). Chela manus acarinate; intercarinal surfaces smooth and setose (fig. 53). Fixed and movable fingers each with 9–12 (n = 12) oblique median denticle subrows; movable finger with 6–11 (n = 12) retrolateral accessory denticles (fig. 21I); proximal dentate margins of fingers straight (fig. 53B), such that no gap evident proximally when fingers closed.

Legs: Legs I–IV, femoral ventral carinae granular; patellar ventral carinae obsolete; intercarinal surfaces smooth. Legs I–IV, tibial spurs absent on I and II, present on III and IV; pro- and retroventral basitarsal (pedal) spurs present, more developed on III and IV. Legs I–IV, macrosetal counts on retrolateral margins of tibiae, 11:14:12:9; basitarsi, 13:20:23:10; telotarsi, 7:8:7:6 (n = 1). Telotarsi ungues slightly elongated, more than three-quarters of telotarsus length, equal on legs I–IV (fig. 22G).

Genital operculum: Genital opercula suboval, completely divided longitudinally, without overlapping, rounded margins (♂) or partially fused longitudinally (♀) (fig. 19E, F). Genital papillae present (♂) or absent (♀).

Pectines: Three marginal lamellae; 11–12 (n = 3) (♂) or 9–12 (n = 2) (♀) median lamellae (fig. 19E, F). Fulcra present. Pectinal teeth along most (♂) or part (♀) of length, dentate margin length:pecten length, 94.1% (90.4%–99.6%, n = 8) (♂) or 86.3% (81.2%–90.2%, n = 6) (♀). Pectinal teeth curved, similar in size; tooth count (sinistral/dextral), 34/35 (32–37/32–37, n = 8) (♂) or 26/27 (25–28/26–28, n = 6) (♀).

Mesosoma: Tergites I–VII progressively increasing in length posteriorly, tergite VI length:tergite VII length, 53.8% (47.7%–58.3%, n = 8) (♂) or 55.9% (51.8%–60.5%, n = 6) (♀); increasing in width posteriorly from I–IV, decreasing in width posteriorly from V–VII. Pretergites smooth; posttergites I–VI, intercarinal surfaces uniformly finely granular, becoming more coarsely and densely granular posteriorly, VII, finely to coarsely and sparsely granular. Tergites I–VI, dorsomedian carinae granular, vestigial, restricted to posterior fifth of I–III and posterior third of IV–VI; dorsosubmedian carinae granular, vestigial, restricted to posterior fifth of I–III and posterior third of IV–VI. Tergite VII, dorsomedian carina granular, vestigial, restricted to anterior half, dorsosubmedian and dorsolateral carinae distinct, granular. Sternites III–VII acarinate, smooth, and glabrous; IV–VI, respiratory spiracles (stigmata) width approximately 3× length.

Metasoma: Metasomal segments I–V becoming longer and narrower posteriorly (figs. 23C, 25C, 27C), segment I shortest, length I:II, 88% (84.3%–89.7%, n = 8) (♂) or 89.5% (87.1%–91%, n = 6) (♀); segments II–IV similar, length II:III, 96.3% (93.4%–98.1%, n = 8) (♂) or 96.7% (92.8%–100.7%, n = 6) (♀), length III:IV, 96.3% (94.1%–98.2%, n = 8) (♂) or 96.3% (93.8%–99.1%, n = 6) (♀); segment V longest, length IV:V, 86.7% (83.6%–91.8%, n = 8) (♂) or 82.3% (79.1%–85.9%, n = 6) (♀); width:length segment I, 61.1% (57.8%–64.4%, n = 8) (♂) or 62.5% (60.7%–65.2%, n = 6) (♀), II, 51.3% (47.2%–53.7%, n = 8) (♂) or 53.5% (51.1%–56.1%, n = 6) (♀), III, 47.7% (43.2%–51%, n = 8) (♂) or 50% (48.4%–53%, n = 6) (♀), IV, 40.1% (36.3%–41.8%, n = 8) (♂) or 41.7% (39.9%–45.1%, n = 6) (♀), V, 33% (31.2%–36.2%, n = 8) (♂) or 33.8% (32.7%–35.5%, n = 6) (♀). Dorsosubmedian and dorsolateral carinae distinct, granular on segments I–III, obsolete on IV, absent on V; dorsosubmedian and dorsolateral carinae densely setose, macrosetal counts on segments I–V (sinistral/dextral), dorsosubmedian carinae, 16/15 (14/11–20/21, n = 5):25/26 (20/18–29/30):26/25 (23/19–31/29):30/27 (24/23–38/31):12/11 (6/7–18/15), dorsolateral carinae, 9/8 (7/5–13/13, n = 5):20/20 (19/16–23/21):22/21 (16/18–28/24):15/14 (11/12–20/17):11/13 (10/11–13/14). Median lateral carinae distinct, granular in posterior three-quarters of segment I, posterior half of II and posterior third of III, absent on IV and V. Ventrolateral carinae distinct, granular, restricted to posterior edge of segment I; granular, with granules becoming progressively larger and subspiniform posteriorly, on II and III; costate-granular on IV; serrate, comprising spiniform granules of variable size, becoming more prominent posteriorly, on V. Ventrosubmedian carinae distinct, costate on segment I; granular, with granules becoming progressively larger and subspiniform posteriorly, on II and III; costate-granular on IV; granular, restricted to anterior three-quarters of V. Ventromedian carina granular, distinct along entire length of V. Dorsal and lateral intercarinal surfaces smooth to finely and sparsely granular on segment I, smooth on II–V; ventral intercarinal surfaces smooth on I and III, finely granular across entire surface on IV and V.

Telson: Telson vesicle width:metasomal segment V width, 80.7% (74.8%–84.6%, n = 8) (♂) or 83% (77.6%–88.5%, n = 6) (♀). Vesicle globose, dorsal surface flat, ventral surface convex and rounded; height:length, 61.2% (51.6%–69.4%, n = 8) (♂) or 62.9% (59.5%–67.6%, n = 6) (♀); dorsal surface smooth; ventral surface finely granular anteriorly, smooth posteriorly; lateral and ventral surfaces densely setose, with 98 (91–102, n = 3) (♂) or 107 (88–127, n = 2) (♀) macrosetae. Aculeus long, gently curved; aculeus length:telson length, 49.6% (43.3%–55.5%, n = 8) (♂) or 51.8% (47.8%–55.5%, n = 6) (♀).

Sexual dimorphism: Adult males and females differ as follows. Males are slightly smaller, on average 67.4 mm in total length, than females, on average 76.6 mm. The carapace intercarinal surfaces are more densely granular and the carinae of the pedipalp femur and patella more coarsely granular in the male than in the female. The pedipalp chela of the male has proportionally shorter fingers than that of the female (fig. 53), as indicated by the higher chela manus length:movable finger length ratio in the male (55.2%) compared with the female (47.1%). The genital opercula are completely divided longitudinally, with overlapping, rounded margins in the male but partially fused longitudinally in the female (fig. 19E, F) and genital papillae are present in the male but absent in the female. The pectinal tooth count is higher in the male (32–37) than in the female (25–28). The spiniform granules of the ventrosubmedian and ventrolateral carinae of metasomal segments II and III, and the ventrolateral carinae of segment V are less prominent in the male than in the female.

Distribution: Buthacus yotvatensis appears to be endemic to Israel and Jordan. Most of the known records occur in the northern and southern parts of the Arava Valley, straddling the southern border between Israel and Jordan, but the species has also been recorded at one locality in the Negev (fig. 5). Whereas records from the Arava Valley range from -294 m to 127 m in elevation, the Negev locality, Ein Zik, is situated at 345 m.

Buthacus yotvatensis is sympatric with the closely related B. arava, in the Arava Valley (fig. 4), but allopatric with B. tadmorensis, distributed from central Jordan, through Syria, southeastern Turkey and Iraq, to the northern shore of the Persian Gulf, in southern Iran (fig. 10).

Ecology: Specimens of B. yotvatensis were collected at night with UV light detection on gravel flats and sparsely vegetated, stable to vegetationless, shifting inland sand dunes. Specimens were fairly common at night, mostly sitting still or walking on the ground surface, some sitting on bushes. The habitat and habitus, notably the pale coloration, smooth tegument, obsolescence of some pedipalpal and metasomal carinae, elongation of the legs, especially legs III and IV, slight dorsoventral compression of the basitarsi of legs I–III, with comblike rows of elongated macrosetae (“sand combs”) along the retrolateral margins, elongated macrosetae on the lateral and ventral surfaces of the telotarsi, and slightly elongated telotarsal ungues, are consistent with the psammophilous ecomorphotype (Prendini, 2001).

Specimens of B. yotvatensis were collected in sympatry with B. arava. Two other buthids, Androctonus crassicauda and Orthochirus scrobiculosus, also inhabit the same area, but occur on harder substrata.

Remarks: In the original description, Levy et al. (1973: 133), stated: “B. yotvatensis is completely different from all scorpions so far described [and] for some time it was doubted whether it should not be placed in a genus of its own.” Kinzelbach (1985) relegated B. yotvatensis to a subspecies of B. tadmorensis, a decision accepted by some authors (e.g., Vachon and Kinzelbach, 1987; Amr et al., 1988; El-Hennawy, 1992) and rejected by others (e.g., Fet and Lowe, 2000; Lourenço, 2004a). Kovařík (2005) subsequently revalidated B. macrocentrus, originally synonymized with B. leptochelys by Kraepelin (1891), and synonymized B. tadmorensis and B. yotvatensis with it. After comparing a paratype of B. yotvatensis, deposited at the MNHN, with two specimens from Palmyra, the type locality of B. tadmorensis, presumed to be part of the original type series, Lourenço (2006) suggested that B. tadmorensis is a valid species and B. yotvatensis could be a junior synonym of B. macrocentrus. Nevertheless, Kovařík et al. (2016) and others (Kaltsas et al., 2008; Navidpour et al., 2008, 2013; Navidpour, 2015) continued to regard B. tadmorensis, B. pietschmanni, and B. yotvatensis as junior synonyms of B. macrocentrus. Lowe et al. (2019) recently rejected Lourenço's (2006) assessment of the validity of B. tadmorensis.

During the present investigation, examination of the types of B. leptochelys and B. macrocentrus, as well as additional material from Sinai and Israel, demonstrated that they are conspecific, as originally concluded by Kraepelin (1891), requiring B. macrocentrus to be returned to synonymy with B. leptochelys (see above). On the contrary, examination of the holotype of B. yotvatensis as well as material from across the distributions of both species, multivariate analysis of morphometrics (fig. 15), and multilocus phylogenetic analysis (fig. 14), confirmed that B. yotvatensis is a valid species, as originally indicated by Levy et al. (1973). Buthacus yotvatensis Levy et al., 1973, stat. rev., is revalidated accordingly.

Material Examined: ISRAEL: Mehoz HaDarom (Southern District): Arava Valley, Avrona Nature Reserve, 29°41′12.5″N 34°59′46.2″E, 48 m, 14–18.ix.2017, E. Gavish-Regev, N. Segev and S. Cain, 13 ♂ (HUJ INVSC 3360, 3362–3367, 3370, 3371, 3373, 3374, 3376, AMNH ex HUJ INVSC 3379), 1 ♀ (HUJ INVSC 3378), 2 subad. ♂ (AMNH ex HUJ INVSC 3377, HUJ INVSC 3372), 1 subad. ♀ (HUJ INVSC 3368), 3 juv. ♂(HUJ INVSC 3369, 3375, 3380), 1 juv. ♀ (HUJ INVSC 3361); Ein-Zik campsite, 30°48′22.4″N 34°51′03.1″E, 345 m, 11.ix.2018, Y. Zvik, 2 ♂(HUJ INVSC 3483, 3485), ♀ (HUJ INVSC 3484), juv. ♀ (HUJ INVSC 3481, 3482); Hai-Bar Yotvata Nature Reserve, 29°50′51.0″N 35°01′42.5″E, 68 m, 26.viii.2011, L. Prendini and T.L. Bird, 32 ♂, 2 ♀, 1 subad. ♀ (AMNH), 1 juv. ♂, 3 juv. ♀(AMCC [LP 11172]), 4.viii.2016, E. Gefen and S. Cain, 3 ♂ (AMCC [LP 15075] ex HUJ INVSC 3331, HUJ INVSC 3333, 3335), 1 ♀ (AMCC [LP 15076] ex HUJ INVSC 3332), 2 subad. ♀ (HUJ INVSC 3329, 3334), 1 juv. ♂ (HUJ INVSC 3330); Hazeva [30°42′N 35°15′E], -129 m, 7.vii.1983, A. Bouskila, 1 ♀ (HUJ INVSC 2532); Hazeva, Nahal Gidron, 30°46′52.2″N 35°14′33.5″E, -136 m, 16. ix.2017, Y. Olek and S. Cain, 1 ♂ (HUJ INVSC 3352); Hazeva, Nahal Mashaq, 30°47′26.1″N 35°15′26.5″E, -155 m, 16.ix.2017, Y. Olek and S. Cain, 2 ♂ (AMNH ex HUJ INVSC 3354, HUJ INVSC 3355), 3 ♀ (AMNH ex HUJ INVSC 3353, 3357, HUJ INVSC 3356); Hazeva, Nahal Shezaf, 30°44′36.0″N 35°16′03.2″E, -134 m, 16.ix.2017, Y. Olek and S. Cain, 1 ♂ (AMCC [LP 15080] ex HUJ INVSC 3359), 1 subad. ♀ (HUJ INVSC 3358); Iddan, N of (Hamada), 30°50′N 35°16′E, -174 m, 15.ix.1988, B. Shalmon, 1 subad. ♂ (HUJ INVSC 2611); Lotan, SE of Lotan Plantations, 30°00′10.4″N 35°05′44.1″E, 127 m, 15.ix.2017, Y. Olek and S. Cain, 5 ♂ (AMCC [LP 15079] ex HUJ INVSC 3350, HUJ INVSC 3346, 3347, 3349, 3351), 1 ♀ (AMCC [LP 15078] ex HUJ INVSC 3348); Nahal Sha'alav, 4 km S of Kibbutz Yahel, 30°02′40.6″N 35°06′33.7″E, 153 m, 15.ix.2017, Y. Olek and S. Cain, 3 ♂ (AMCC [LP 15077] ex HUJ INVSC 3342, AMNH ex HUJ INVSC 3343, HUJ INVSC 3344), 1 ♀ (HUJ INVSC 3345); Qetura [29°58′N 35°03′E], 100 m, M. Israel, 8.x.1974, 1 ♂ (HUJ INVSC 2330), 20.x.1974,1 ♀(HUJ INVSC 2332), 17.iv.1975, 2 ♀ (HUJ INVSC 2336, 2337); Samar [29°50′N 35°01′E], 103 m, 25.ii.1985, U. Mendel, 1 subad. ♀ (HUJ INVSC 2612); Samar sand dunes, E of Eliphaz date plantations, 29°48′06.6″N 35°02′19.2″E, 64 m, 4. viii.2016, E. Gefen and S. Cain, 4 ♂ (AMCC [LP 15073, 15074] ex HUJ INVSC 3311, 3312, HUJ INVSC 3313, 3314), 30.vii.2017, S. Cain, 4 ♂(AMNH ex HUJ INVSC 3319, 3320, HUJ INVSC 3316, 3317), 1 ♀ (HUJ INVSC 3318), 24.v.2017, S. Cain, 1 subad. ♀ (HUJ INVSC 3315); Samar sand dunes, old quarry, 29°48′25.9″N 35°02′06.6″E, 70 m, 13.ix.2017, Y. Olek and S. Cain, 3 ♂ (AMNH ex HUJ INVSC 3308, HUJ INVSC 3305, 3307), 1 subad. ♀ (HUJ INVSC 3306); Samar sand dunes, E side of Samar date plantations, 29°48′44.2″N 35°02′27.9″E, 67 m, 4.viii.2016, E. Gefen and S. Cain, 1 ♂ (HUJ INVSC 3321), 24.v.2017, S. Cain, 4 ♂ (AMNH ex HUJ INVSC 3324, 3327, HUJ INVSC 3322, 3326), 1 ♀ (AMNH ex HUJ INVSC 3323), 1 subad. ♂ (AMNH ex HUJ INVSC 3328), 1 juv. ♂(HUJ INVSC 3325); Samar sand dunes, N part of Samar Nature Reserve, 29°49′36.8″N 35°02′35.8″E, 65 m, 13.ix.2017, Y. Olek and S. Cain, 5 ♂ (AMNH ex HUJ INVSC 3303, 3304, HUJ INVSC 3300–3302); Samar sand dunes, Samar Nature Reserve, 29°49′06.9″N 35°02′40.2″E, 64 m, 13.ix.2017, Y. Olek and S. Cain, 2 ♂ (AMNH ex HUJ INVSC 3309, HUJ INVSC 3310); Timna, sand dunes E of, [29°45′N 35°00′E], 109 m, 24.iii.1972, Y. Ayal, 1 ♂ (HUJ INVSC 2303), 1 juv. ♀ (HUJ INVSC 2304), 22.ii.1972, M. Broza, 1 juv. ♀ (HUJ INVSC 2305); Wadi Fukra [Zin Tichon, 30°50′N 34°49′E], 384 m, 20.iii.1944, 1 ♀ (HUJ INVSC 2492); Yotvata, [29°53′N 35°03′E], 64 m, 20.x.1974, M. Israel, 1 ♀ (HUJ INVSC 2333); Yotvata, E side of circle field, 29°53′28.7″N 35°04′40.5″E, 72 m, 14. ix.2017, Y. Olek and S. Cain, 3 ♂ (AMNH ex HUJ INVSC 3340, HUJ INVSC 3339, 3341); Yotvata, NE side of circle field, 29°54′02.6″N 35°04′42.1″E, 77 m, 30.vii.2017, S. Cain, 2 ♂(AMNH ex HUJ INVSC 3337, HUJ INVSC 3338); Yotvata, SE side of circle field, 29°53′21.8″N 35°04′40.9″E, 71 m, 30.vii.2017, S. Cain, 1 ♂(AMNH ex HUJ INVSC 3336). JORDAN: Karak Governorate: Wadi Khanzeerah (Khanzairh), 30°53′39.9″N 35°25′38.3″E, -294 m, 9.ix.2013, L. Prendini, Z. Amr and L. Al Azam, 1 ♀ (AMNH). Tafilah Governorate: Wadi Al Ghwaibeh (Ghwaqibeh), 30°47′35.6″N 35°23′39.7″E, -119 m, 9. ix.2013, L. Prendini, Z. Amr and L. Al Azam, 2 ♂, 1 ♀ (AMNH), [leg] (AMCC [LP 13561]); Wadi Al Ghwaibeh (Ghwaqibeh), 30°48′13.3″N 35°24′21.6″E, -106 m, 9.ix.2013, L. Prendini, Z. Amr, and L. Al Azam, 2 ♂, 1 ♀ (AMNH).