Androctonus Ehrenberg, 1828

Androctonus Ehrenberg, 1828 View in CoL

( Figures 1–179 View Figures 1–4 View Figures 5–12 View Figures 13–18 View Figures 19–40 View Figure 41 View Figures 42–45 View Figures 46–52 View Figures 53–58 View Figures 59–76 View Figures 77–83 View Figures 84–87 View Figures 88–91 View Figures 92–95 View Figures 100–107 View Figures 108–129 View Figures 130–131 View Figures 132–137 View Figures 138–152 View Figures 153–164 View Figures 165–172 View Figures 173–177 View Figures 178–179 , Tables 1–2)

Androctonus Ehrenberg in Hemprich & Ehrenberg, 1828 (part), pl. II, fig. 1–2, 4–5, 8.

TYPE SPECIES. Scorpio australis Linnaeus, 1758 .

DIAGNOSIS. Adults 50–110 mm in length. Carapace subrectangular, granulate, with distinct anterior submedian, superciliary, central median, central lateral and posterior median carinae, the latter two usually not fused into a lyre configuration; five pairs of lateral eyes, in type 5 pattern ( Loria & Prendini, 2014). Sternum type 1 (Soleglad & Fet, 2003), relatively small, and markedly triangular in shape. Tergites granulate, I–VI with 1–3 carinae, VII with 5 carinae; sternites III–VI smooth, spiracles narrow, slit-like; sternite VII with 4 carinae; metasoma I with 10 carinae, II–III with 8–10 carinae, V with 5 carinae. Metasomal segments robust, wide and deep, with strong carinae; metasoma I–IV with dorsal carinae elevated posteriorly and dorsomedian furrow excavated; metasoma V with dorsal carinae granulate; posterior margins of tergite/ sternite V, and metasoma I–III with microsetal fringes. Telson with vesicle pyriform, lacking subaculear tubercle; aculeus stout, long, strongly curved. Chelicerae with typical buthid dentition (Vachon 1963), fixed finger with two denticles on ventral surface. Pedipalps orthobothriotaxic, type Aβ (Vachon 1974, 1975); femur with d 2 on dorsal surface, e 2 distal to d 5; patella with d 3 internal to dorsomedian carina; chela with Eb 2 proximal to Eb 1, eb on proximal fixed finger, db in middle region of fixed finger, db proximal to or level with est, dt and it located on distal fixed finger; pedipalp femur with dense, compact distal external macrosetal cluster (Lowe & Tang, 2024); dentate margins of fixed and movable chela fingers with 9–16 linear, non-imbricated subrows of denticles, each flanked by single internal and external accessory denticles; movable finger typically with 3 large subterminal denticles; fixed and movable fingers with two pairs of subterminal Cruz-Armas sensilla, distal pair elongate (Lowe & Fet, 2024); male fingers with or without lobe/ notch combination on basal dentate margins. Pectines with fulcra and internal fulcra, lacking accessory internal fulcra; basal teeth and basal middle lamella unmodified; pectinal tooth counts: ♂ 24–37, ♀ 19–29. Legs III– IV with tibial spurs, legs I–III typically with bristle-combs on tarsomere I; all legs with two submedian rows of macrosetae on ventral surface of tarsomere II, and prolateral and retrolateral pedal spurs. Hemispermatophore with moderately long trunk, capsule with 3-lobed sperm hemiduct, basal lobe hook-like, flagellum folded with shorter, broader pars recta, and longer, narrower pars reflecta.

REMARKS ON KARYOTYPES. We analyzed male karyotypes of three newly described Androctonus species from Morocco. The chromosomes of these species ( Figs. 173–177 View Figures 173–177 ) exhibit cytogenetic features typical of the family Buthidae : holocentric chromosomes, achiasmatic male meiosis, and a relatively low chromosome number (e. g., Mattos et al., 2013). The diploid sets of A. argoubi sp. n. (samples 1512, 1513, 1514, 1517, 1570, and holotype 2866, Figs. 173, 174 View Figures 173–177 ), A. stockmanni sp. n. ( holotype 1268, Figs. 175, 176 View Figures 173–177 ), and A. vaneki sp. n. ( holotype 2867, Fig. 177 View Figures 173–177 ) each comprise 24 chromosomes, with one pair consistently much larger than the remainder. These characteristics match the karyotypes reported for all cytogenetically studied Androctonus species across their range ( SadÍlek et al., 2015; Šťáhlavský et al., 2020), supporting the hypothesis that the genus has a stable karyotype ( SadÍlek et al., 2015). The absence of chromosomal variation is further supported by the exclusive presence of bivalents in all observed postpachytene nuclei of A. argoubi sp. n. ( Fig. 174 View Figures 173–177 ) and A. stockmanni sp. n. ( Fig. 175 View Figures 173–177 ).

REMARKS ON HEMISPERMATOPHORES. The basal lobe (hook) of the holotype of A. stockmanni was noticeably shorter and more rounded than those observed in two samples of A. sergenti (cf. Figs. 85–86 View Figures 84–87 vs. Figs. 78–79, 81–83 View Figures 77–83 ). Vachon (1952) illustrated in drawings the outlines of the basal lobes of several species of Androctonus , which showed considerable variation: A. aeneas (small, broad based; fig. 163), A. mauretanicus (small, narrow, longer; fig. 185), A. bourdoni (small, narrow, short), A. hoggarensis (small, narrow, moderate length; fig. 197), A. australis (small, narrow, longer; fig. 207), and A. amoreuxi (small, short, broad; fig. 225). Comparative studies of much larger samples are needed to test whether these interspecific differences are statistically significant and useful for species level taxonomy.

AFFINITIES. Androctonus sergenti Vachon, 1948 and the three new species described here are all characterized by having metasoma I–IV ventrally punctate, a character that is either quite well developed or at least indicated (cf. Figs. 169–172 View Figures 165–172 ). Other Androctonus species from Morocco and Western Sahara studied here lack punctae on the ventral surfaces of metasomal segments ( Fig. 164 View Figures 153–164 ). Other diagnostic characters include: base color uniformly black or blackish; metasoma I smooth dorsally without fine granulation (cf. Fig. 52 View Figures 46–52 vs. Fig. 163 View Figures 153–164 ); pedipalp chela narrow with long fingers, always narrower than patella, chela length/width ratio 6.0– 7.6 in both sexes (cf. Figs. 156–161 View Figures 153–164 vs. 153–155). The base color is uniformly yellow to reddish yellow in A. amoreuxi (Audouin, 1825) , and A. maroccanus Lourenço et al., 2009 . Metasoma I is finely granulated dorsally in A. liouvillei (Pallary, 1924) ( Fig. 163 View Figures 153–164 ). The pedipalp chela is wide with shorter fingers in at least the males (chela length/width ratio 3.5–5.4 in both sexes) of A. agrab Ythier & Lourenço, 2022 , A. bourdoni Vachon, 1948 ( Fig. 154 View Figures 153–164 ), A. gonneti Vachon, 1948 ( Fig. 155 View Figures 153–164 ), A. maroccanus Lourenço et al., 2009 , and A. mauritanicus (Pocock, 1902) ( Fig. 153 View Figures 153–164 ).