Buthacus Birula, 1908

Cain, Shlomo, Gefen, Eran & Prendini, Lorenzo, 2021, Systematic Revision Of The Sand Scorpions, Genus Buthacus Birula, 1908 (Buthidae C. L. Koch, 1837) Of The Levant, With Redescription Of Buthacus Arenicola (Simon, 1885) From Algeria And Tunisia, Bulletin of the American Museum of Natural History 2021 (450), pp. 1-137 : 21-24

Buthacus Birula, 1908

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, 2003 a: 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, 2; 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 ven- tral 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).