Buthacus leptochelys (Ehrenberg, 1829)

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 : 68-69

Buthacus leptochelys (Ehrenberg, 1829)

Figures 1C , 6 , 15 , 17A, B , 21D , 22D , 24B , 26B , 28B , 29D , 39 , 40 , 41 ; tables 1, 4, 5

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,

Figures

FIGURE 1. Buthacus Birula, 1908, representative habitats. A. Mamshit [Kurnub], Israel, habitat of Buthacus amitaii, sp. nov. B. Naftah to Ong Jmal, Tunisia, habitat of Buthacus arenicola (Simon, 1885). C. Faiyum Oasis, Egypt, habitat of Buthacus leptochelys (Ehrenberg, 1829). D. Ashdod Nizzanim Nature Reserve, Israel, habitat of Buthacus nitzani Levy et al., 1973, stat. nov. E. Yotvata, Israel, habitat of Buthacus arava, sp. nov., and Buthacus yotvatensis Levy et al., 1973, stat. rev. F. Be’er Milka, Israel, Holot Haluza sand dunes, Israel, habitat of Buthacus levyi, sp. nov., and B. nitzani.

FIGURE 6. Map of southwestern Levant, plotting known locality records for Buthacus leptochelys (Ehrenberg, 1829) in Egypt and Israel, based on material examined for the present study and the literature. Pale gray symbol denotes type locality of the junior synonym, Androctonus (Leiurus) thebanus Ehrenberg, 1828 whereas dark gray symbol denotes type locality of the junior synonym, Buthacus granosus Borelli, 1929.

FIGURE 15. Nonmetric multidimensional scaling (NMDS) plots for A, male and B, female Buthacus Birula, 1908, based on 24 morphometric ratios and counts, for 79 adults (each symbol represents an individual): Buthacus amitaii, sp. nov. (white triangles); Buthacus arava, sp. nov. (black triangles); Buthacus arenicola (Simon, 1885) (white inverted triangles); Buthacus leptochelys (Ehrenberg, 1829) (white circles); Buthacus levyi, sp. nov. (white squares); Buthacus nitzani Levy et al., 1973, stat. nov. (white diamonds); Buthacus tadmorensis (Simon, 1892), stat. rev. (black circles); Buthacus yotvatensis Levy et al., 1973, stat. rev. (black squares).

FIGURE 17. Buthacus Birula, 1908, carapace, dorsal aspect. A, B. Buthacus leptochelys (Ehrenberg, 1829), A. ♂ (HUJ INVSC 859), B. ♀ (HUJ INVSC 1807). C, D. Buthacus nitzani Levy et al., 1973, stat. nov., C. ♂ (HUJ INVSC 3227), D. ♀ (HUJ INVSC 3226). E, F. Buthacus amitaii, sp. nov., E. Holotype ♂ (HUJ INVSC 3480), F. Paratype ♀ (HUJ INVSC 3205). Scale bars = 2 mm.

FIGURE 21. Buthacus Birula, 1908, dextral pedipalp chela movable finger, dorsal aspect, illustrating dentition. A. Buthacus amitaii, sp. nov., holotype ♂ (HUJ INVSC 3480). B. Buthacus arava, sp. nov., paratype ♂ (HUJ INVSC 3290). C. Buthacus arenicola (Simon, 1885), ♂ (AMNH). D. Buthacus leptochelys (Ehrenberg, 1829), ♂ (HUJ INVSC 859). E, F. Buthacus levyi, sp. nov., E. Holotype ♂ (HUJ INVSC 2082), F. Paratype ♀ (HUJ INVSC 3252). G. Buthacus nitzani Levy et al., 1973, stat. nov., ♂ (HUJ INVSC 3227). H. Buthacus tadmorensis (Simon, 1892), stat. rev., lectotype ♂ (NHMW 2453) of Buthus pietschmanni Penther, 1912. I. Buthacus yotvatensis Levy et al., 1973, stat. rev., ♂ (HUJ INVSC 3302). Scale bars = 1 mm.

FIGURE 22. Buthacus Birula, 1908, dextral legs I–IV, basitarsus and telotarsus, ventral aspect. A. Buthacus arenicola (Simon, 1885), ♂ (AMNH). B. Buthacus levyi, sp. nov., holotype ♂ (HUJ INVSC 2082). C. Buthacus arava, sp. nov., paratype♂ (HUJ INVSC 3295). D. Buthacus leptochelys (Ehrenberg, 1829), ♀ (HUJ INVSC 1807). E. Buthacus tadmorensis (Simon, 1892), stat. rev., lectotype ♂ (NHMW 2453) of Buthus pietschmanni Penther, 1912. F. Buthacus nitzani Levy et al., 1973, stat. nov., ♀ (HUJ INVSC 3226). G. Buthacus yotvatensis Levy et al., 1973, stat. rev., ♂ (HUJ INVSC 3302). H. Buthacus amitaii, sp. nov., holotype ♂ (HUJ INVSC 3480). Scale bars = 1 mm.

FIGURE 24. Buthacus Birula, 1908, metasomal segments I–V and telson, dorsal aspect. A. Buthacus levyi, sp. nov., paratype ♀ (HUJ INVSC 3274). B. Buthacus leptochelys (Ehrenberg, 1829), ♀ (HUJ INVSC 1807). C. Buthacus nitzani Levy et al., 1973, stat. nov., ♀ (HUJ INVSC 3226). D. Buthacus amitaii, sp. nov., paratype ♀ (HUJ INVSC 3205). Scale bars = 5 mm.

FIGURE 26. Buthacus Birula, 1908, metasomal segments I–V and telson, lateral aspect. A. Buthacus levyi, sp. nov., paratype ♀ (HUJ INVSC 3274). B. Buthacus leptochelys (Ehrenberg, 1829), ♀ (HUJ INVSC 1807). C. Buthacus nitzani Levy et al., 1973, stat. nov., ♀ (HUJ INVSC 3226). D. Buthacus amitaii, sp. nov., paratype ♀ (HUJ INVSC 3205). Scale bars = 5 mm.

FIGURE 28. Buthacus Birula, 1908, metasomal segments I–V and telson, ventral aspect. A. Buthacus levyi, sp. nov., paratype ♀ (HUJ INVSC 3274). B. Buthacus leptochelys (Ehrenberg, 1829), ♀ (HUJ INVSC 1807). C. Buthacus nitzani Levy et al., 1973, stat. nov., ♀ (HUJ INVSC 3226). D. Buthacus amitaii, sp. nov., paratype ♀ (HUJ INVSC 3205). Scale bars = 5 mm.

FIGURE 29. Buthacus Birula, 1908, telson, lateral aspect. A. Buthacus arenicola (Simon, 1885), ♂ (AMNH). B. Buthacus levyi, sp. nov., holotype ♂ (HUJ INVSC 2082). C. Buthacus arava, sp. nov., paratype ♂ (HUJ INVSC 2559). D. Buthacus leptochelys (Ehrenberg, 1829), ♂ (HUJ INVSC 859). E. Buthacus tadmorensis (Simon, 1892), stat. rev., lectotype ♂ (NHMW 2453) of Buthus pietschmanni Penther, 1912. F. Buthacus nitzani Levy et al., 1973, stat. nov., ♂ (HUJ INVSC 3227). G. Buthacus yotvatensis Levy et al., 1973, stat. rev., ♂ (HUJ INVSC 3302). H. Buthacus amitaii, sp. nov., holotype ♂ (HUJ INVSC 3480). Scale bars = 1 mm.

FIGURE 39. Buthacus leptochelys (Ehrenberg, 1829), habitus, dorsal (A, C), and ventral (B, D) aspects. A, B. ♂ (HUJ INVSC 859). C, D. ♀ (HUJ INVSC 1807). Scale bars = 1 cm.

FIGURE 40. Buthacus leptochelys (Ehrenberg, 1829), ♂ (HUJ INVSC 859), dextral pedipalp femur (A, B) and patella (C, E), prolateral (A, C), dorsal (B, D), and retrolateral (E) aspects. Scale bar = 2 mm.

FIGURE 41. Buthacus leptochelys (Ehrenberg, 1829), dextral pedipalp chela, dorsal (A, C) and retrolateral (B, D) aspects. A, B. ♂ (HUJ INVSC 859). C, D. ♀ (HUJ INVSC 1807). Scale bar = 2 mm.