Hadruroides Pocock, 1893

Telegonus: L. Koch, 1867 (part): 235.

Hadrurus: Thorell, 1876 (part): 186; Karsch, 1879 (part): 135, 1881: 290; Boeris, 1889: 125.

Caraboctonus: Pocock, 1893a (part): 92.

Hadruroides Pocock, 1893b: 306, 309, 329, 330, type species by original designation: Hadrurus charcasus Karsch, 1879 (= Hadruroides charcasus [Karsch, 1879]); Kraepelin, 1894: 182, 206; Laurie, 1896: 130; Kraepelin, 1899: 188; Pocock, 1900: 474; Birula, 1917a: 163, 189; Birula, 1917b: 44, 48, 51; Mello-Leitão, 1945: 119; Bücherl, 1964: 61, 1967: 115; Williams, 1970: 31; Bücherl, 1971: 328; Stahnke, 1974: 122; Maury, 1975: 10, 11; Francke, 1977: 75; Francke and Soleglad, 1981: 235, 256, figs. 9, 13, 27–33, 58; Francke, 1985: 8, 17, 20; Sissom, 1990: 130, 131; Nenilin and Fet, 1992: 14; Lourenço, 1994: 157, 1995: 74–76, 1997a: 601, 602, 1998: 135; Kovařík, 1998: 135; Lourenço, 2000: 25; Sissom and Fet, 2000: 411–413; Lourenço and Dastych, 2001: 55, 56; Soleglad and Sissom, 2001: 31, fig. 34; Acosta and Ochoa, 2002: 19; Soleglad and Fet, 2003a: 2, 5, 12, 2003b: 8, 111; Fet et al., 2004: 18, 23, 24; Prendini and Wheeler, 2005: 482; Fet and Soleglad, 2005: 12; Ochoa, 2005: 65, fig. 17; Dupre´, 2007: 5, 14; Ochoa and Chaparro, 2008: 5; Francke and Prendini, 2008: 210; Fet and Soleglad, 2008: 255; Soleglad et al., 2009: 1.

DIAGNOSIS: The genus Hadruroides comprises small- to medium-sized scorpions, varying from ca. 30 mm, e.g., H. leopardus Pocock, 1900, and H. graceae, n. sp., to ca. 80 mm in total length, e.g., H. charcasus (Karsch, 1879) (Maury, 1975). The most important diagnostic characters for the genus are as follows: cheliceral movable fingers, internal surface with two subdistal teeth and one prominent basal tooth; carapace anterior margin slightly convex, with three pairs of lateral ocelli; sternum subpentagonal, with Yshaped sulcus; pedipalp chela smooth, acarinate in most species, except H. charcasus, in which granular carinae are present on internal surface; chela movable finger, median denticle row with six or seven median subrows of denticles and variable number of accessory denticles; neobothriotaxic major Type C trichobothrial pattern (Vachon, 1974): femur with three trichobothria (i, e, d); patella with 20: one internal (i), two dorsal (d 1, 2), three ventral (v 1–3), v 3 situated on external surface, and 14 external trichobothria (et 1–3, est, em 1–3, esb 1, 2, eb 1–5); chela with 26 trichobothria: 14 on manus (Et 1–5, Est, Esb, Eb 1–3, V 1–4) and 12 on fixed finger (Dt, Db, et, est, esb, eb, dt, dst, dsb, db, it, ib); leg telotarsus with ventromedian row of spinule clusters (setaceous tufts); metasomal segment V with complete, granular VL and VM carinae; telson slightly concave dorsally, with short aculeus; hemispermatophore lamelliform, with elongated crest, accompanied externally by medial slit and one free lobe, truncal flexure absent, internobasal reflection of the sperm duct absent.

Hadruroides appears to be most closely related to Caraboctonus. The two genera exhibit several similarities, e.g., trichobothrial pattern, ventromedian row of spinule clusters on telotarsus, dentition of chelicerae and pedipalp chela fingers. They may be distinguished by means of the ventral carinae of metasomal segment V: in Caraboctonus, the VL carinae are restricted to the posterior half of the segment and the VM carina is absent, whereas in Hadruroides, both VL and VM carinae are present and complete (except in H. tishqu, n. sp., in which the VL and VM carinae are restricted to the posterior two-thirds of the segment). The two genera may be further distinguished on the basis of pedipalp chela finger dentition: internal and external accessory denticles, flanking the median denticle row in Hadruroides, are absent in Caraboctonus. Furthermore, the hemispermatophore of Hadruroides exhibits an elongated crest and a medial slit, both absent from the hemispermatophore of Caraboctonus, which instead exhibits a digit-shaped process.

DISTRIBUTION: The genus Hadruroides is endemic to Ecuador, Peru, northern Chile, and several offshore islands, including the Galápagos (table 1). Thirteen of the 16 described species occur in Peru (figs. 1, 2). Four species inhabit dry forest in northern Peru: H. charcasus (fig. 4E), H. chinchaysuyu, n. sp. (fig. 4B); H. geckoi, n. sp. (figs. 3B, 4C); H. leopardus Pocock, 1900 (figs. 3A, 5B). Three species occur in inter-Andean valleys along the Cordillera: H. bustamantei Ochoa and Chaparro, 2008 (fig. 3D); H. carinatus Pocock, 1900 (fig. 4A); H. mauryi Francke and Soleglad, 1980. Six species inhabit desert along the Pacific coast: H. aguilari Francke and Soleglad, 1980; H. graceae, n. sp. (fig. 4D); H. juanchaparroi, n. sp. (figs. 3C, 5A); H. lunatus (L. Koch, 1867) (figs. 3G, 5C); H. tishqu, n. sp. (figs. 3E, 5E); H. vichayitos, n. sp. (figs. 3F, 5D). We exclude H. maculatus (Thorell, 1876), which is restricted to the central coastline of Ecuador, from the list of Peruvian species. Two other Hadruroides species are endemic to Ecuador: H. galapagoensis Maury, 1975; H. udvardyi Lourenco, 1995 (table 1). Most species of Hadruroides have restricted distributions, except H. charcasus and H. lunatus, which are apparently more widely distributed. We consider it necessary to reassess all previous records of the latter two species, because we suspect several are based on misidentifications. Other records of Hadruroides from the coastal desert of southern Peru and northern Chile (Ochoa, 2005; unpubl. data) correspond to new species related to H. lunatus (fig. 2), the descriptions of which are in preparation by the authors.

PERUVIAN SPECIES OF HADRUROIDES

KEY TO IDENTIFICATION OF THE SPECIES

1. Adult pedipalp chela robust, internomedian, dorsointernal, and dorsal marginal carinae well developed and comprising strong granules (fig. 4E)............. H. charcasus

– Pedipalp chela smooth, acarinate (figs. 8E, 11G).. .. .. .. ... .. .. .. ... .. .. ... 2

2. Sternite VII acarinate................ 3

– Sternite VII with two or four carinae (figs. 7C, D, 27C)........................ 6

3. Metasomal segments I–IV elongated; segment II longer than wide; segment V (♂) approximately three times longer than wide, length:width ratio: 3.18...... H. aguilari

– Metasomal segments I–IV relatively short (fig. 20E); segment II as wide as long; segment V (♂) approximately twice longer than wide, length:width ratio: 2.04–2.59 (figs. 19G, 23B)................... 4

4. Pigmentation reduced to faint spots on carapace and tergites (fig. 24B), absent on metasomal segments; segment V, VM and VL carinae reduced to posterior two-thirds of segment (fig. 23B); telson, ventral surfaces smooth (♂, ♀) (fig. 23A, C).... H. tishqu

– Pigmentation pronounced on carapace, tergites, and metasomal segments (fig. 20C, D); segment V, VM and VL carinae complete (fig. 19G); telson, ventral surfaces smooth or sparsely granular anteriorly (♂), granular (♀) ... .. .. .. ... .. .. .. ... .. .. ... 5

5. Tergites I–VI with rectangular dorsosubmedian spots of pigmentation (fig. 20C); sternite VII and metasomal segment I, ventral surfaces unpigmented; hemispermatophore broader basally with relatively long crest... H. lunatus

– Tergites I–VI with irregular dorsosubmedian spots of pigmentation (fig. 20D); sternite VII and metasomal segment I, ventral surfaces with several spots of pigmentation surrounding insertion of setae (fig. 20E); hemispermatophore slender with short crest (fig. 21)........................ H. juanchaparroi

6. Chela fixed finger (adult ♂) curved, creating distinct proximal gap with movable finger when fingers closed (figs. 8C, 11H, 27H)....... 8

– Chela fixed and movable fingers (adult ♂) straight, no proximal gap evident when fingers closed (fig. 15I).............. 7

7. Metasomal segment IV, ventral surface densely granular (fig. 15C); segment V with 7–8 ventral setae and 6–8 lateral setae; pedipalp chela length:width ratio: 5.00–5.41 (♂), 4.46– 5.12 (♀)................... H. graceae

– Metasomal segment IV, ventral surface smooth; segment V with 12–18 ventral setae and 3–5 lateral setae; pedipalp chela length:width ratio: 3.93–4.54 (♂), 4.23–4.64 (♀).................... H. leopardus

8. Metasomal segments, VSM carinae absent on segments I–III, present on IV........ 12

– Metasomal segments, VSM carinae present on segment I and, usually, II and III, absent on IV............................ 9

9. Pigmentation pronounced on carapace, tergites, and metasomal segments (figs. 8I, 9C, 12F, D); VSM carinae present on metasomal segments I–III; pedipalp chela fixed finger (adult ♂) strongly curved, creating well developed proximal gap with movable finger when fingers closed (fig. 11H)......... 10

– Pigmentation absent, except for small, faint spots on carapace and tergites (fig. 12G); VSM carinae obsolete on metasomal segment I, absent on segments II and III; pedipalp chela fixed finger (adult ♂) slightly curved, creating weak proximal gap with movable finger when fingers closed (fig. 27H)...... H. vichayitos

10. Sternite VII, VL and VSM carinae well developed; metasomal segment V, DL carinae smooth; telson, ventral surface (♀) smooth; hemispermatophore lamina apically rounded................ H. carinatus

– Sternite VII, VL carinae distinct, VSM carinae obsolete (fig. 7C); metasomal segment V, DL carinae sparsely to densely granular (figs. 8 A, D, 11A); telson, ventral surface (♀) granular (fig. 8B); hemispermatophore lamina apically acuminate (figs. 9E, 13).............. 11

11. Metasomal segment V elongated (♂): length:width ratio: 2.40–2.89 (fig. 11B); segment V, ventral surface sparsely granular, comprising scattered granules in posterior third (fig. 11B); segment V with 10–14 DL setae and 9–13 VL setae; pedipalp chela slender, length:width ratio: 4.41–4.40 (♂), 4.53–4.74 (♀) (figs. 11E, 12B); pectinal tooth count: 21– 23 (♂), 18–21 (♀); hemispermatophore lamina curved in distal half (fig. 13)..... H. geckoi

– Metasomal segment V short (♂): length:width ratio: 1.97–2.14 (fig. 8F); segment V, ventral surface densely granular (fig. 8F); segment V with 5–8 DL setae and 6–10 VL setae; pedipalp chela relatively broad, length:width ratio: 3.66–4.02 (♂), 4.0–4.18 (♀) (fig. 8E); pectinal tooth count: 17–20 (♂), 16 (♀); hemispermatophore lamina straight in distal half (fig. 9E–H)........ H. chinchaysuyu

12. Tergites I–IV each with paired dorsosubmedian and dorsolateral spots of pigmentation, forming four longitudinal stripes along mesosoma; legs I–IV each with several prolateral spots of pigmentation; pedipalp chela (♂) relatively broad, length:width ratio: 3.09–3.36............. H. bustamantei

– Tergites I–IV unpigmented, mesosoma without longitudinal stripes; legs without prolateral spots of pigmentation; pedipalp chela (♂) relatively slender, length:width ratio: 2.7– 2.8...................... H. mauryi