Odontobuthus, VACHON, 1950

Barahoei, Hossein, Prendini, Lorenzo, Navidpour, Shahrokh, Tahir, H. Muhammad, Aliabadian, Mansour, Siahsarvie, Roohollah & Mirshamsi, Omid, 2022, Integrative systematics of the tooth-tailed scorpions, Odontobuthus (Buthidae), with descriptions of three new species from the Iranian Plateau, Zoological Journal of the Linnean Society 195, pp. 355-398 : 361-364

GENUS ODONTOBUTHUS VACHON, 1950

( FIGS 1–22 , TABLES 1 –5)

Odontobuthus Vachon, 1950: 153 , 154.

Type species: Buthus doriae Thorell, 1876 [= Odontobuthus doriae ( Thorell, 1876) ]; Vachon, 1952: 325, 326; Pringle, 1960: 83, fig.8; Khalaf, 1963: 59; Vachon, 1966: 213; Habibi, 1971: 44; Stahnke, 1972: 132; Vachon, 1974: 906; Tikader & Bastawade, 1983: 247–253, figs 714–726; Francke, 1985: 10, 16; Kinzelbach, 1985: map I; Farzanpay, 1987: 154–156; Vachon & Kinzelbach, 1987: 92; Farzanpay, 1988: 39; Farzanpay, 1990: 7; Sissom, 1990: 92, 98, 101, 102, fig. 3.17J; Nenilin & Fet, 1992: 17; El-Hennawy, 1992: 102, 105, 128; Akbari et al., 1997: 112; Kovarik, 1997: 47, 49, figs 11–13; Dehghani et al., 1998: 126; Kovarik, 1998: 115; Pooladgar, 1999: 176; Fet & Lowe, 2000: 187; Kovařík, 2002: 9; Lourenço & Pézier, 2002: 115–124, figs 1–27; Soleglad & Fet, 2003: 26; Fet et al. 2005: 3, 12, 22; Prendini & Wheeler, 2005: 481; Vignoli & Crucitti, 2005: 6–7, fig. 2d; Dehghani et al., 2008: 176; Navidpour et al., 2008a: 3, 5, 13–14, 28, figs 18, 22, 52–55; 2008b: 1, 3, 15, 21, figs 33–36; 2008c: 3, 11–12, 17, figs 21–24; 2008d: 3, 7–8, 14, figs 27–30; Shahi et al., 2008: 207, 210–212; Kovařík, 2009: 24; Lourenço & Duhem, 2009: 38, 44, 45, 48; Navidpour & Lowe, 2009: 49, 58; Pirali-Kheirabadi et al., 2009: 1, 3, 9–10, 13, figs 10, 24–24; Vatani & Khoobdel, 2009: 7–8, 10; Lowe, 2010: 1–22, figs 1–65; Ramezani et al., 2010: 55–56, 58–59, figs 2, 3, 5; Navidpour et al., 2011: 3, 13, 22, figs 5, 8, 16, 21–24; Fekri et al., 2012: 137, 138, 140; Karataş et al., 2012: 116; Navidpour et al., 2012: 3, 14, 23, figs 2, 4, 5, 11, 25, 26; Mirshamsi et al., 2013: 153–170, figs 4–41; Mozaffari et al., 2013: 5414, fig. 7; Navidpour et al., 2013: 3, 11–14, 21, figs 3, 7, 10, 13–23, 38, 39; Pirali-Kheirabadi et al., 2013: 48; Azghadi et al., 2014: 169–179, figs 2, 6; Moradi et al., 2014: 8; Navidpour & Mirshamsi, 2014: 112; Nejati et al., 2014: 219–220; Ghassemi et al., 2015: 264, 265, fig. 3; Kassiri et al., 2015: 101–103, fig. 1; Moradi et al., 2015: 13; Nazari et al., 2015: 257, 259; Ahmadi et al., 2016:1568; Ahmadimarzale et al., 2017: 337–339; Dehghani & Kassiri, 2017: 433–438, figs 2, 3; Dehghani et al., 2017:113–116, figs 1, 3; Sharifinia et al., 2017: 243, 245, 246; Nejati et al., 2018: 7141, 7142; Navidpour et al., 2019: 2, 5, figs 45, 46, 49, 57; Barahoei et al., 2020: 399–402, figs 18–20; Motevalli Haghi et al., 2020: 69–72.

Odonbuthus: Pérez Minnocci, 1974: 28 . Diagnosis: The triangular sternum and absence of trichobothria on the ventral part of the pedipalp patella place Odontobuthus in the family Buthidae . The presence of prominent tooth-like processes (spiniform granules) on the ventrosubmedian carinae of metasomal segments II and III separate Odontobuthus from most other Iranian Buthidae , except Buthacus tadmorensis (Simon, 1892) and Kraepelinia palpator Vachon, 1974 . Species of Odontobuthus are easily distinguished from B.tadmorensis by the lyriform configuration of the carapacial carinae, and from Kraepelinia by their larger size (˃ 35 mm) and relatively longer fingers of the pedipalp chela. Kraepelinia is also easily distinguished from Odontobuthus by the extremely thickened base of the aculeus, the height of which is about half the height of the telson vesicle in lateral aspect, much greater than in Odontobuthus .

Odontobuthus may be separated from other genera of Buthidae by the following combination of characters ( Stahnke, 1972; Farzanpay, 1987; Lourenço & Pézier, 2002; Lowe, 2010; Mirshamsi et al., 2013; Navidpour et al., 2013): cheliceral movable finger with two ventral denticles; carapace dorsal surface entirely horizontal in lateral aspect; carapacial carinae distinct, with lyriform configuration; pedipalp trichobothriotaxy type A-β; trichobothrium eb situated at distal end of chela manus or on fixed finger; patella trichobothrium d 3 situated prolateral to dorsomedian carina; proximal dentate margins of chela fixed and movable fingers not sexually dimorphic in adult male, i.e. without notch and lobe, respectively, in lateral aspect; median denticle row of chela fixed and movable fingers each comprising 10–14 oblique, non-imbricate subrows of denticles, each flanked by single prolateral and retrolateral accessory denticles; tibial spurs present on legs III and IV; pectines with fulcra; ventrosubmedian carinae of metasomal segments II and III comprising prominent tooth-like processes (spiniform granules); anteroventral carina of metasomal segment IV comprising prominent granules; ventrolateral carinae of metasomal segment V with prominent lobate processes; anal arch with prominent lobes laterally and reduced lobes posteroventrally; telson globose, without subaculear tubercle; hemispermatophore capsule with hook-like basal lobe and sperm hemiduct of three lobes.

Figures

Figure 1. Geographical distribution of the buthid scorpion genus Odontobuthus plotted on topography of the Middle East. A, known locality records obtained from fieldwork and the literature. Circles with black or white dots indicate type localites of each species. B, localities of material used in molecular phylogenetic analysis (see Table 1 for details).

Figure 2. Boxplots of sex-corrected total length for species of Odontobuthus occurring in Iran.

Figure 3. Lineardiscriminant analysis (LDA) of38 metricand fivemeristic variables among18 populationsofOdontobuthus. Conspecific populations in same colour.

Figure 4. Neighbour-joining phenogram generated from morphometric Mahalanobis distance matrix of 18 populations of Odontobuthus. Conspecific populations in same colour.

Figure 5. Fifty per cent majority rule consensus obtained by analysis of partial DNA sequences of mitochondrial COI gene from Odontobuthus with Bayesian inference (BI). Nodal support values above branches represent posterior probability values from BI and bootstrap values from maximum likelihood analyses, respectively. Letters A and B refer to clades also recovered by Azghadi et al. (2014).

Figure 6. Odontobuthus baluchicus. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, holotype ♀ (SMF). C, D, paratype ♂ (ZMFUM 1302). Scale bars = 10 mm.

Figure 7. Odontobuthus baluchicus. Metasomal segmentV and telson, lateral aspect (A, E), ventral aspect (B, F), metasomal segments II and III, lateral aspect (C, G), ventral aspect (D, H). A–D, holotype ♀ (SMF). E–H, paratype ♂ (ZMFUM 1302). Scale bars = 3 mm (A–D), 2.5 mm (E–H).

Figure 8. Odontobuthus baluchicus. Pedipalp femur, dorsal aspect (A, B), patella, dorsal aspect (C, D) and chela, lateral aspect (E, F). A, C, E, holotype ♀ (SMF). B, D, F, paratype ♂ (ZMFUM 1302). Scale bars = 2.5 mm.

Figure 9. Odontobuthus chabaharensis. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, holotype ♀ (SMF). C, D, paratype ♂ (ZMFUM 1325). Scale bars = 10 mm (A, B), 5 mm (C, D).

Figure 10. Odontobuthus chabaharensis. Metasomal segment V and telson, lateral aspect (A, E), ventral aspect (B, F), metasomal segments II and III, lateral aspect (C, G), ventral aspect (D, H). A, C, E, holotype ♀ (SMF). B, D, F, paratype ♂ (ZMFUM 1325). Scale bars = 2.5 mm.

Figure 11. Odontobuthus chabaharensis. Pedipalp femur, dorsal aspect (A, B), patella, dorsal aspect (C, D) and chela, lateral aspect (E, F). A, C, E, holotype ♀ (SMF). B, D, F, paratype ♂ (ZMFUM 1325). Scale bars = 2.5 mm (A–D), 2 mm (E, F).

Figure 12. Odontobuthus kermanus. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, holotype ♀ (SMF). C, D, paratype ♂ (ZMFUM 1340). Scale bars = 10 mm (A, B), 5 mm (C, D).

Figure 13. Odontobuthus kermanus. Metasomal segment V and telson, lateral aspect (A, E), ventral aspect (B, F), metasomal segments II and III, lateral aspect (C, G), ventral aspect (D, H). A, C, E, holotype ♀ (SMF). B, D, F, paratype ♂ (ZMFUM 1340). Scale bars = 3 mm (A–D), 2.5 mm (E–H).

Figure 14. Odontobuthus kermanus. Pedipalp femur, dorsal aspect (A, B), patella, dorsal aspect (C, D) and chela, lateral aspect (E, F). A, C, E, holotype ♀ (SMF). B, D, F, paratype ♂ (ZMFUM 1340). Scale bars = 2.5 mm (A, C, E), 1.5 mm (B, D), 2 mm (F).

Figure 15. Odontobuthus bidentatus. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, ♀ (ZMFUM 2067). C, D, ♂ (ZMFUM 1317). Scale bars = 10 mm.

Figure 16. Odontobuthus. Habitus, dorsal aspect (A, C, E), ventral aspect (B, D). A, B, Odontobuthus brevidigitus, ♂ (ZMFUM 1438). C–E, Odontobuthus odonturus, ♀ (ZMFUM 1439). Scale bars = 10 mm (A, B), 5 mm (C–E).

Figure 17. Odontobuthus doriae. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, ♀ (ZMFUM 2045). C, D, ♂ (ZMFUM 2016). Scale bars = 10 mm.

Figure 18. Odontobuthus tavighiae. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, ♀ (ZMFUM 1362). C, D, ♂ (ZMFUM 1364). Scale bars = 10 mm.

Figure 19. Odontobuthus tirgari. Habitus, dorsal aspect (A, C), ventral aspect (B, D). A, B, ♀ (ZMFUM 2091). C, D, ♂ (ZMFUM 2092). Scale bars = 10 mm.

Figure 20. Odontobuthus. Metasomal segment V, lateral aspect. A, Odontobuthus bidentatus ♀ (ZMFUM 2061) B, Odontobuthus brevidigitus ♂ (ZMFUM–1438). C, Odontobuthus doriae ♀ (ZMFUM 2014). D, Odontobuthus odonturus ♀ (ZMFUM 1439). E, Odontobuthus tavighiae ♀ (ZMFUM–2074). F, Odontobuthus tirgari ♂ (ZMFUM 1398). Scale bars = 2.5 mm.

Figure 21. Odontobuthus. Metasomal segment V, ventral aspect. A, Odontobuthus bidentatus ♀ (ZMFUM 2061) B, Odontobuthus brevidigitus ♂ (ZMFUM 1438). C, Odontobuthus doriae ♀ (ZMFUM 2014). D, Odontobuthus odonturus ♀ (ZMFUM–1439). E, Odontobuthus tavighiae ♀ (ZMFUM 2074). F, Odontobuthus tirgari ♂ (ZMFUM 1398). Scale bars = 2.5 mm.

Figure 22. Odontobuthus. Metasomal segments II and III, lateral aspect. A, Odontobuthus bidentatus ♀ (ZMFUM 2061). B, Odontobuthus brevidigitus ♂ (ZMFUM 1438). C, Odontobuthus doriae ♀ (ZMFUM 2014). D, Odontobuthus odonturus ♀ (ZMFUM 1439).E, Odontobuthus tavighiae ♀ (ZMFUM 2074).F, Odontobuthus tirgari ♂ (ZMFUM 1398).Scale bars = 2.5 mm.