Rugodentinae Bastawade et al., 2005: 77, 82, type genus by original designation: Rugodentus Bastawade et al., 2005; Sureshan et al., 2007a: 2903, 2905, 2908; Kovařík, 2009: 6, 28; Aswathi and Sureshan, 2017: 9844.

Pandininae: Rossi, 2016a: 7–9, 19 (part).

Diagnosis: Rugodentidae can be placed unequivocally in the superfamily Scorpionoidea Latreille, 1802, based on the following combination of characters (Prendini, 2000a; Prendini et al., 2003): cheliceral movable finger with one subdistal tooth and without serrula; pedipalps with Type C trichobothrial pattern (femur with three trichobothria; patella with 19 or more; chela with 26 or more); retrolateral pedal spurs of telotarsi absent; pro- and retroventral macrosetae of telotarsi spiniform; genital opercula (♀) fused along the midline.

Rugodentidae can be separated from other scorpionoid families by the following combination of characters: pedipalp chela fingers with multiple primary rows of denticles; pedipalps Type C, neobothriotaxic major, with accessory trichobothrium in e series of patella; chela manus, trichobothrium Db located on dorsal surface; chela fixed finger, trichobothria Dt and ib located proximally, it distally, db on prolateral surface, eb midway on finger, and esb in distal third; telotarsi I–IV each with pro- and retroventral rows of spiniform macrosetae, laterodistal lobes rounded; metasomal segments I–IV each with paired ventrosubmedian carinae, segment V with transverse carina; telson vesicle not laterally compressed, without anterodorsal lateral lobes and subaculear tubercle.

Supplementary Description: The following summary is based on a reinterpretation of the description and illustrations of Bastawade et al. (2005), reproduced in figs. 11 and 12, supplemented by images 24 and 25 in Aswathi and Sureshan (2017: 9849) and live habitus (fig. 13). Carapace with median notch in anterior margin; median longitudinal sulcus narrow, with anterior bifurcation. Three pairs of lateral ocelli (MLMa, PLMa, PDMi). Median ocular tubercle raised. Cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth unequal, DE tooth considerably smaller than DI tooth (less than half its length), aligned longitudinally and not opposable. Pedipalp patella prolateral process obsolete; retrodorsal and retrolateral carinae obsolete. Pedipalp chela dorsal secondary, subdigital, digital, retromedian, ventromedian, proventral and promedian carinae obsolete; retroventral carina distinct, parallel to longitudinal axis of chela, with distal edge directed toward a point between pro- and retroventral movable finger condyles, but closer to retroventral condyle. Pedipalp chela fingers with multiple primary rows of denticles. Pedipalps Type C, neobothriotaxic major, with accessory trichobothrium in e series of patella. Pedipalp chela with trichobothrium Db located on dorsal surface of manus; Et₂ located on retrolateral surface of manus; db located on prolateral surface of fixed finger; Est located distally on manus; Dt and ib located proximally on fixed finger; eb located midway on fixed finger, proximal to or aligned with db and dorsal to esb–est–et axis; esb located in distal third of fixed finger, aligned with est–et axis; it located distally on fixed finger. Sternum longer than wide, pentagonal. Telotarsi I–IV each with pro- and retroventral rows of spiniform macrosetae; laterodistal lobes rounded; retrolateral pedal spurs absent. Metasomal segments I–IV each with paired ventrosubmedian carinae; segment V with transverse carina. Telson vesicle not laterally compressed and without anterodorsal lateral lobes; aculeus long, shallowly curved, without subaculear tubercle.

Included Taxa: This family accommodates the monotypic genus, Rugodentus Bastawade et al., 2005, stat. rev.

Distribution: This family is endemic to the state of Kerala, India (fig. 14).

Remarks: The monotypic Indian scorpionoid genus, Rugodentus, was recently synonymized with Heterometrus, creating a new homonym, Heterometrus keralaensis Bastawade et al., 2005, for which the name Heterometrus bastawadei Rossi, 2016, was devised. Rossi (2016a: 19) presented the following argument for synonymizing Rugodentus:

This argument is inconsistent with other changes in Rossi's (2016a) “revised classification” of Scorpionidae, among them the revalidation of Protophthalmus Lawrence, 1969, which contradicts the phylogenetic analyses of Lamoral (1978a, 1979), based on morphology, and subsequent analyses of Prendini et al. (2003), based on morphology and DNA sequences from the nuclear and mitochondrial genomes. Furthermore, Rossi's (2016a) arguments are unsupported by the evidence. The diagnostic characters of Rugodentus are inconsistent not only with the diagnosis of Heterometrus but with the diagnosis of Scorpionidae (Prendini, 2000a; Prendini et al., 2003), revealing that Rugodentus lacks the synapomorphies of either, and shares potential synapomorphies with several other families.

Although a comparative diagnosis was not presented, Bastawade et al. (2005: 82) justified the new subfamily, genus, and species as follows:

As correctly described and illustrated (fig. 11B) by Bastawade et al. (2005: 79, 80, figs. 2, 10, 14, 16), the dentition of the pedipalp chela fingers is indeed unlike that observed in Heterometrus or other Scorpionidae:

This was confirmed in photographs published subsequently by Aswathi and Sureshan (2017: 9847, 9849, images 24, 25), who described the dentition as “rugously granular … in a band along the interior surface.”

The dentition described and illustrated (fig. 11B) most closely resembles the “multiple rows” condition (Stockwell, 1989; Prendini, 2000a) observed in three genera of Bothriuridae Simon, 1880, i.e., the Australian Cercophonius Peters, 1861, and South American Centromachetes Lönnberg, 1897, and Urophonius Pocock, 1893, and in the Australian Urodacidae Pocock, 1893, and Malagasy Heteroscorpionidae Kraepelin, 1905, two families erroneously synonymized by Soleglad et al. (2005), as discussed by Volschenk and Prendini (2008). Additionally, the dentate margins of the fixed and movable fingers (gently scalloped, with lobes and corresponding notches conspicuously absent) of Rugodentus (Bastawade et al., 2005: 79, 80, figs. 2, 10, 14, 16; Aswathi and Sureshan, 2017: 9849, image 25) are unlike those of Heterometrus or other Scorpionidae, in which a series of pronounced lobes and corresponding notches are evident in the fixed and movable fingers (Prendini, 2000a, 2016; Prendini et al., 2003). The dentate margins of the fixed and movable fingers of Rugodentus are more akin to those observed in other families, e.g., Bothriuridae, Heteroscorpionidae, Urodacidae, and the Middle Eastern Hemiscorpiidae Pocock, 1893, another family misconstrued by Soleglad and Fet (2003a), Fet and Soleglad (2005), and Soleglad et al. (2005), as discussed by Monod and Prendini (2014).

Furthermore, unlike Heterometrus, in which the pedipalps are orthobothriotaxic (Prendini et al., 2003), the pedipalps of Rugodentus are neobothriotaxic (fig. 12), with an accessory e trichobothrium on the patella (14 trichobothria are present on the retrolateral surface in Rugodentus whereas only 13 are present in Heterometrus). Accessory e trichobothria on the pedipalp patella occur in all African scorpionid genera except Scorpio as well as in Heteroscorpionidae and Urodacidae, some Iranian species of Hemiscorpiidae, the South American bothriurid genera, Timogenes Simon, 1880, and Vachonia Abalos, 1954, the Mexican diplocentrid genus, Kolotl Santibáñez-López et al., 2014, the African hormurid genus, Hadogenes Kraepelin, 1894, and some Neotropical species of the hormurid genus, Opisthacanthus Peters, 1861 (Prendini, 2000a; Monod and Lourenço, 2005; Santibáñez-López et al., 2014b; Kovařík et al., 2017a).

Indeed, the trichobothrial pattern of the pedipalp chela appears to be unique to Rugodentus. Rugodentus (fig. 12) is unlike any other scorpionoid in the relatively distal positions of trichobothria eb and esb, situated midway on the fixed finger and in its distal third, respectively. Rugodentus is also the only scorpionoid in which trichobothrium Dt is situated proximally on the fixed finger, as in Diplocentridae Karsch, 1880, and Hemiscorpiidae, and it situated distally on the fixed finger, as in the Middle Eastern diplocentrid genus, Nebo Simon, 1878. Neither character is exhibited by Scorpionidae (Prendini, 2000a), with which Diplocentridae was erroneously synonymized by Soleglad and Fet (2003a) and Fet and Soleglad (2005). Additionally, trichobothrium Db is situated on the dorsal surface of the manus in Rugodentus, a character otherwise unique to Hemiscorpiidae and Scorpionidae, whereas db is situated on the prolateral surface of the fixed finger in Rugodentus, as in Diplocentridae, Hemiscorpiidae, Scorpionidae, and most Hormuridae Laurie, 1896, rather than on its dorsal surface, as in Bothriuridae, Heteroscorpionidae, Urodacidae, and some Hormuridae (Prendini, 2000a).

Finally, the ventral surface of metasomal segment V bears a distinct transverse carina in Rugodentus (fig. 11I), described as a “granular transverse submarginal ridge present only on inferior portion,” and illustrated by Bastawade et al. (2005: 79, 81, fig. 9). This character, observed in most genera of Diplocentridae and some Bothriuridae, e.g., Bothriurus Peters, 1861, Timogenes and Vachonia, is also absent among Scorpionidae (Prendini, 2000a, 2003).

Rugodentus lacks a key diagnostic character of Scorpionidae (Prendini, 2000a; Prendini et al., 2003): i.e., pedipalp chela dorsal secondary carina partially developed, extending partway across dorsal surface of manus, subdigital carina partially developed, extending partway across manus in opposite direction.

One character that Rugodentus shares with Scorpionidae is the presence of rounded laterodistal lobes on the leg telotarsi (fig. 11D), described as “tarsus ... lateral distal margin partly overlapping the base” and illustrated by Bastawade et al. (2005: 79, 81, fig. 4), but this character also occurs in Diplocentridae and Urodacidae (Prendini, 2000a).

Several other aspects of the habitus of Rugodentus are also noteworthy (figs. 11–13). The shape of the carapace (e.g., the truncate anterior margin) and the pedipalp chelae (described as “globular” by Aswathi and Sureshan, 2017: 9849) resemble some Bothriuridae and Urodacidae, rather than Scorpionidae, and the robust metasoma and shape of the telson also resemble some Bothriuridae, e.g., Centromachetes.

It is clear from the preceding discussion that Rugodentus is a valid genus that is neither con-familial with Scorpionidae nor with any other scorpionoid family but appears most closely allied to Diplocentridae, Scorpionidae, and Urodacidae. Kovařík (2009: 6, 28) likewise doubted the placement of Rugodentus in Scorpionidae but failed to suggest an alternative:

Subfamily Rugodentinae Bastawade et al., 2005, is hereby elevated to the rank of family, Rugodentidae Bastawade et al., 2005, genus Rugodentus Bastawade et al., 2005, and its type species Rugodentus keralaensis Bastawade et al., 2005, are revalidated, and the following new synonym is presented: Heterometrus bastawadei Rossi, 2016 = Rugodentus keralaensis Bastawade et al., 2005.

Rugodentus Bastawade et al., 2005: 77–82, type species by original designation: Rugodentus keralaensis Bastawade et al., 2005; Sureshan et al., 2007a: 2903, 2908; Kovařík, 2009: 6, 28; Javed et al., 2010a: 143; Mirza et al., 2012: 1; Aswathi and Sureshan, 2017: 9844, 9847.

Heterometrus: Rossi, 2016a: 6–9, 15, 19, 20, 25 (misidentification).

Diagnosis: As for family.

Included Species: Rugodentus keralaensis Bastawade et al., 2005, stat. rev.

Distribution: As for family.

Rugodentus keralaensis Bastawade et al., 2005: 77–82, figs. 1–16, photo 1; Sureshan et al., 2007a: 2903, 2905–2908, fig. 1; Aswathi and Sureshan, 2017: 9844, 9847, 9849, images 24, 25.

Heterometrus bastawadei Rossi, 2016a: 7–9, 25, syn. nov.

Type Material: INDIA: Kerala: Ernakulam Distr.: Holotype ♂ (ZSI 11002), Kodanad [10°11′N 76°31′E], Mallana Reserve Forest, 12.ii.1999, P.M. Sureshan.

It is unclear whether the holotype is lost. Attempts to locate it in the ZSI collections (D.B. Bastawade, in litt.) were unsuccessful.

Diagnosis: As for family.

Distribution: This species is endemic to India and currently known only from the Ernakulam District of Kerala (fig. 14).

Ecology: The known localities are situated in wet, lowland rainforest in the foothills of the Western Ghats. According to P. Jain (personal commun.), these fossorial scorpions were observed retreating into burrows at night under UV light. The habitat and habitus (Bastawade et al., 2005: 83, pl. 1; fig. 12) are consistent with the pelophilous ecomorphotype (Prendini, 2001b), like most Diplocentridae, Scorpionidae, and Urodacidae, and many Bothriuridae. This species is sympatric with the buthid Reddyanus brachycentrus (Pocock, 1899) and the scorpionid Sahyadrimetrus mathewi gen. et sp. nov.

Scorpionides Latreille, 1802: 46, 47 (part), type genus: Scorpio Linnaeus, 1758; 1804: 110 (part); 1806: 130 (part); 1810: 116, 118 (part); Leach, 1814: 412; 1815: 390; Sundevall, 1833: 29 (part).

Scorpiones: Ehrenberg in Hemprich and Ehrenberg, 1828, pl. I, figs. 1, 2 (family, part).

Buthides: C.L. Koch, 1837: 36, 37 (part).

Centrurides C.L. Koch, 1837: 38 (part), type genus: Centrurus Ehrenberg, 1829 [= Heterometrus Ehrenberg, 1828].

Pandinoidae Thorell, 1876a: 11 (part), type genus: Pandinus Thorell, 1876; 1876b: 83 (part); Karsch, 1879a: 19 (part); Thorell and Lindström, 1885: 25 (part).

Heterometridae Simon, 1879: 92, 115, type genus: Heterometrus Ehrenberg, 1828 (part) [= Scorpio Linnaeus, 1758].

Scorpionidae: Lankester, 1885: 379 (part); Pocock, 1893: 305, 306 (part); Kraepelin, 1894: 8 (part); Laurie, 1896b: 128 (part); Kraepelin, 1899: 96 (part); Pocock, 1900a: 11, 84; Kraepelin, 1901: 270 (part); 1905: 343 (part); 1913: 165; Birula, 1917a: 161 (part); 1917b: 58, 59 (part); Kopstein, 1921: 128; Pavlovsky, 1924a: 78 (part); 1925: 190, 191, 195 (part); Kopstein, 1926: 111; Fage, 1933: 27; Werner, 1934: 275 (part); Fage, 1936: 179, 181; Kästner, 1941: 232 (part); Fage, 1946: 73; Millot and Vachon, 1949: 428 (part); Petrunkevitch, 1955: 75; Bücherl, 1964: 59 (part); 1967: 114 (part); Tikader, 1973: 263; Stahnke, 1974: 339 (part); L.E. Koch, 1977: 159 (part); Lamoral, 1980: 440 (part); Levy and Amitai, 1980: 102 (part); Couzijn, 1981: 4, 5, 20; Francke, 1982: 75 (part); Tikader and Bastawade, 1983: 2, 3, 27, 32, 517, 518, fig. 45; Biswas, 1984: 318, 320, 321; Francke, 1985: 18; Lourenço, 1989: 161, 174; Sissom, 1990: 131; Nenilin and Fet, 1992: 5 (part); Kovařík, 1997: 183; 1998: 136; Lourenço, 1999: 929 (part); Fet, 2000: 427, 428 (part); Lourenço, 2000: 25; Prendini, 2000a: 34, 35, 39, 44; Indra, 2001: 55, 56; Bastawade, 2002: 295; Lourenço and Huber, 2002: 273; Kovařík, 2003: 136, table 1; Prendini et al., 2003: 185–189, 192–194, 206, 209, 210, 212, 217, 220, 223, 226, 230, 235, 250, 252–259, tables 1, 3, 4, 6, figs. 1–3, 8, appendices 1, 2; Soleglad and Fet, 2003a: 1, 3, 4, 5, 8, 29, 56, 58, 59, 61, 67, 71, 75, 84, 86, 88, 111–117, 119–121, 138, 139, 141, 145, 148, tables 1, 9, 11, figs. 114, 126, appendices A, B (part); 2003b: 2, 5, 8, 18, 25, 31, table 1, figs. 5, 22; Bastawade, 2004: 286, 290; Bastawade et al., 2004: 43, 44, 47, 49, 56, 57, figs. 6, 10; Bastawade, 2005: 411, 417; Fet and Soleglad, 2005: 9, 13 (part); Prendini, 2005: 28, 29, 32–36, tables 1, 2, figs. 1, 2; Prendini and Wheeler, 2005: 447, 448, 449, 471, 477, 481, 482, 491, tables 2, 10, appendix 1; Soleglad et al., 2005: 1, 2, 4, 6, 8, 14–17, 19, 20, 21, 23–26, 28–30, 32, 33–35, tables 1, 4, 6, figs. 37–39, 68–70, 72–74 (part); Thulsi Rao et al., 2005: 3, 8; Bastawade, 2006a: 131; Indra, 2006: 197, 199; Sureshan et al., 2007a: 2903, 2905; 2007b: 49, 52; Wirkner and Prendini, 2007: 402; Bastawade, 2008: 133, 136; Bastawade and Borkar, 2008: 212, 215, 218; Kamenz and Prendini, 2008: 6, 31, 33, table 1; Prendini and Bird, 2008: 77; Volschenk and Prendini, 2008: 242, 249; Volschenk et al., 2008: 653, 656, 669; Bastawade, 2009: 215, 219; Indra, 2009: 141; Kovařík, 2009: 6, 14, 15, 17, 18, 27, 28, 33 (part), fig. 8, pl. J; Prendini, 2009: 12; 16, 17, 22, 32, 34, 40, tables 1, 3, fig. 6; Javed et al., 2010a: 143; 2010b: 785; Michalik and Mercati, 2010: 96; Prendini, 2011: 117; Klußmann-Fricke et al., 2012: 549, 555, fig. 2; Mirza et al., 2012: 1; Di et al., 2013: 52, 86, 94; Veronika et al., 2013: 72, 76; Loria and Prendini, 2014: 5, 7, 12, 14, 16, 20, 21, 23, tables 2–4; Lourenço, 2014: 141; Amr et al., 2015: 30, 31; Rossi, 2015a: 11, 16, 21; Sharma et al., 2015: 2, 4, 5, 7, 8, table 1, figs. 1–4; Adams et al., 2016: 1, 7; Janion-Scheepers et al., 2016: 147; Rossi, 2016a: 7–9, 18, 20, 26; 2016b: 47; Aswathi and Sureshan, 2017: 9845, 9847; Esposito et al., 2017: 4, 14; Pham et al., 2017: 136; Esposito et al., 2018: 89, 90, 115, 116, appendix 1.

Pandinidae: Marx, 1890: 211.

Diagnosis: Members of the family Scorpionidae can be placed unequivocally in the superfamily Scorpionoidea on the basis of the following characters (Prendini, 2000a; Prendini et al., 2003): retrolateral pedal spurs of telotarsi absent; pro- and retroventral macrosetae of telotarsi spiniform; genital opercula (♀) fused along the midline; paraxial organ (♂) with an internobasal reflection of the sperm duct. Additional characters that, in combination, place Scorpionidae unequivocally in the Scorpionoidea, are as follows (Prendini, 2000a; Prendini et al., 2003): cheliceral movable finger with one subdistal tooth and without serrula; pedipalps with Type C trichobothrial pattern (femur with three trichobothria; patella with 19 or more; chela with 26 or more); hemispermatophore lamelliform with distinct truncal flexure.

Scorpionidae can be separated from all other scorpionoid taxa by the following character: pedipalp chela with dorsal secondary carina partially developed, extending partway across dorsal surface of manus, and subdigital carina partially developed, extending partway across in opposite direction. Scorpionidae can also be separated from all other scorpionoid taxa, except the diplocentrid subfamily Diplocentrinae Karsch, 1880, by the following characters: pedipalp chela with ventromedian carina more strongly developed than proventral and promedian carinae; promedian and proventral carinae equally developed (often obsolete); pedipalp patella with trichobothrium d₂ located on prolateral surface.

Characters separating Scorpionidae from particular scorpionoid families are as follows (Prendini, 2000a; Prendini et al., 2003): paired ventrosubmedian carinae of metasomal segments I–IV separate Scorpionidae from Hemiscorpiidae, Heteroscorpionidae and Urodacidae; rounded laterodistal lobes of the telotarsi separate Scorpionidae from Heteroscorpionidae and Hormuridae; absence of a subaculear tubercle on the telson vesicle separates Scorpionidae from Diplocentridae.

Included Taxa: As redefined in the present contribution, Scorpionidae comprises 18 genera and 152 species grouped into four extant subfamilies: Heterometrinae Simon, 1879, stat. nov.; Opistophthalminae Rossi, 2016, stat. nov.; Pandininae Thorell, 1876; Scorpioninae Latreille, 1802.

Distribution: Africa: Algeria, Angola, Benin, Botswana, Burkina Faso, Cameroon, Central African Republic, Chad, Congo, Côte d'Ivoire, Democratic Republic of Congo, Egypt, Equatorial Guinea (including Bioko Island), Eritrea, Ethiopia, Gabon, the Gambia, Ghana, Guinea, Guinea-Bissau, Kenya, Lesotho, Liberia, Libya, Malawi, Mali, Mauritania, Morocco, Mozambique, Namibia, Nigeria, Senegal, Sierra Leone, Somalia, South Africa, Sudan, Swaziland, Tanzania, Togo, Tunisia, Zambia, Zimbabwe. Asia: Bangladesh, Brunei, Cambodia, India (including the Andaman and Nicobar islands), Indonesia, Iran, Iraq, Israel and the Palestinian territories, Jordan, Kuwait, Laos, Lebanon, Malaysia, Myanmar, Nepal, the Philippines, Qatar, Saudi Arabia, Somalia, Singapore, Sri Lanka, Syria, Thailand, Turkey, Vietnam, Yemen.

Heterometrinae are endemic to Asia, with greatest species richness and endemism in India, followed by Indonesia. Opistophthalminae are endemic to Africa, with greatest species richness and endemism in South Africa, followed by Namibia. Pandininae are near endemic to Africa (including the Arabian Peninsula), with greatest species richness and endemism in Somalia. Scorpioninae is the most widespread scorpionid subfamily, extending from the West African countries of Benin and Senegal, across North Africa to Iran, but the greatest species richness and endemism occurs in Morocco. Only two scorpionid subfamilies occur in the same country and the greatest scorpionid species richness and endemism occurs in South Africa.

Heterometridae Simon, 1879: 92, 115, type genus: Heterometrus Ehrenberg, 1828 (part) [= Scorpio Linnaeus, 1758].

Pandininae: Rossi, 2016a: 19, 20, 22, 26 (part).

Heterometrini: Rossi, 2016a: 19, 20, 25, 26.

Diagnosis: Heterometrinae are the sister group of Pandininae (Prendini, 2000a; Prendini et al., 2003). Both subfamilies share the following characters, by means of which they can be separated from Opistophthalminae and Scorpioninae: presence of a stridulatory organ, comprising a “rasp” (granular tubercles) and “scraper” (stridulatory setae or scaphotrix), on opposing surfaces of the coxae of the pedipalps and the first pair of legs; similar counts of pro- and retroventral spiniform macrosetae on telotarsi I and II and on III and IV. Heterometrinae can be separated from Pandininae by means of the following characters: pedipalps orthobothriotaxic, with 26 trichobothria (patella with 13 trichobothria in e series and 3 trichobothria in v series; chela with 4 trichobothria in V series and 2 trichobothria in i series); granular tubercles of “rasp” and stridulatory setae (scaphotrix) of “scraper” situated on coxae of first leg and pedipalp, respectively. Heterometrinae can be further separated from Opistophthalminae by means of the following characters: pedipalps orthobothriotaxic, with 26 trichobothria (including 13 trichobothria in e series of patella); cheliceral coxae without stridulatory setae (scaphotrix) on prodorsal surfaces and chemoreceptive lamelliform setae (trichocopae) on promedian surfaces. Heterometrinae can be further separated from Scorpioninae by means of the following characters: digital carina of pedipalp chela obsolete; mesosomal sternite VII usually without paired ventrosubmedian and ventrolateral carinae.

Included Taxa: As redefined in the present contribution, Heterometrinae includes seven genera and 41 species: Chersonesometrus Couzijn, 1978, stat. nov. et stat. rev.; Deccanometrus, gen. nov.; Gigantometrus Couzijn, 1978, stat. nov.; Heterometrus Ehrenberg, 1828; Javanimetrus Couzijn, 1981, stat. nov. et stat. rev.; Sahyadrimetrus, gen. nov.; Srilankametrus Couzijn, 1981, stat. nov. et stat. rev.

Distribution: Asia: Bangladesh, Brunei, Cambodia, China (including Hainan Island and Tibet Autonomous Region), India (including the Andaman and Nicobar Islands), Indonesia (Babi, Bali, Bangka, Batu Islands, Belitung, Bengkalis, Java, Kalimantan, Madura, Mentawai Islands, Miang-Besar, Nias, Riau Islands, Siberut, Simeulue, Sumatra, Weh), Laos, Malaysia (mainland, Langkawi, Penang, Tinggi, and Tioman islands, Sabah and Sarawak), Myanmar (mainland, Myeik Islands), Nepal, Pakistan, the Philippines (Balabac, Luzon, Mindanao, Palawan, Panay, Tawi Tawi), Singapore, Sri Lanka, Thailand (mainland, Ko Chang, Ko Lanta, Ko Muk, Ko Samui, Phi Phi islands and Phuket), Vietnam (mainland, Phú Quoc Island).

Heterometrinae are distributed from India and Sri Lanka throughout the Southeast Asian mainland and archipelagos as far as Wallace's Line (Vachon, 1953; Couzijn, 1981; Prendini et al., 2003), where they mostly inhabit tropical and subtropical rainforests, moist and dry tropical deciduous forests, and tropical thorn forests (Couzijn, 1981; Tikader and Bastawade, 1983; Sissom, 1990). Reports of Heterometrus from the Indonesian islands of Sulawesi and Halmahera (Meise, 1932: 662; Takashima, 1945: 90), to the east of Wallace's Line, have never been confirmed (Couziijn, 1981: 175, 178).

Ecology: The ability to construct burrows has been confirmed in all genera of Heterometrinae (Couzijn, 1981; Hull-Williams, 1989; Khatavkar and More, 1990; Shivashankar and Veeresh, 1991; Shivashankar, 1992, 1994; Tare et al., 1993; Prendini et al., 2003; Javed et al., 2010a; Pande et al., 2012; Tahir and Prendini, 2014). Burrows are constructed in loamy riverbanks and other sloping ground, at the base of stones and among the roots of trees (Couzijn, 1981; Shivashankar, 1994; fig. 12). As in some species of Pandininae, composite, multi-entranced burrows containing multiple (up to 15) related individuals of different ages have been recorded in some species, e.g., Chersonesometrus madraspatensis and Deccanometrus xanthopus (Shivashankar, 1994; Prendini et al., 2003; Javed et al., 2010a). In others, e.g., Heterometrus longimanus (Herbst, 1800) and H. spinifer, mixed age groups of related and unrelated individuals cohabit with minimal aggression or cannibalism in laboratory terraria (Harrison, 1954; Polis and Lourenço, 1986) and, according to some evidence, this also occurs in the wild (Schultze, 1927).

Few ecological data are available for most Heterometrinae. However, the thickened metasoma, short, robust legs with stout, spiniform macrosetae distributed laterally and distally on the basitarsi, and curved telotarsal ungues of most genera occurring in South Asia, as well as the Southeast Asian Javanimetrus, are consistent with the fossorial, pelophilous ecomorphotype (Prendini, 2001b). Gigantometrus often inhabits termitaria and most if not all species of Heterometrus appear to be lapidicolous or corticolous as well as fossorial. For example, H. longimanus is often found under the loose bark of dead standing trees, under decaying trunks and logs, or in the cavities of rotten tree stumps (Schultze, 1927; Prendini et al., 2003). Furthermore, the morphology of the species of Heterometrus differs from that of most South Asian genera in elongation of the legs, absence of a retrolateral row of spiniform macrosetae on the leg tibia, presence of strongly curved ungues and an enlarged, curved pseudonychium (dactyl) on the telotarsi, and enlarged median ocelli. Although such characters are uncommon among scorpionids, they occur in arboreal chactids (e.g., Chactas Gervais, 1844) and hormurids (e.g., Hormurus Thorell, 1876, and some species of Opisthacanthus), and are regarded as corticolous adaptations to an arboreal habitat (Prendini, 2001b). Although some species of Chersonesometrus and Deccanometrus are dorsoventrally compressed and evidently semilithophilous, inhabiting the cracks and crevices of rock outcrops, no Heterometrinae exhibit psammophilous adaptations.

Conservation Status: Several species of Heterometrinae are harvested and consumed for food in Southeast Asia and for narcotic or medicinal purposes in India and Pakistan (Menzel and D'Aluisio, 1998; Prendini et al., 2003; Majumder and Dey, 2005; Tahir and Prendini, 2014).

Various species of Heterometrinae are readily available in Europe, the United States, China, and Japan for the exotic pet trade because their impressive size and fearsome appearance are highly prized by collectors. An extensive literature exists on the captive husbandry of Asian forest scorpions (e.g., Nemenz and Gruber, 1967; Hull-Williams, 1986; Hosoi, 1990; Dupré, 1993; Gopalakrishnakone et al., 1995; Condevaux-Lanloy, 1996; Mahsberg et al., 1999; Rubio, 2000). The following species and countries of origin have been personally recorded in the trade: C. fulvipes (India, Pakistan); C. madraspatensis (India); D. bengalensis (India); D. obscurus (India); G. swammerdami (India); H. laevigatus (Myanmar, Thailand); H. petersii (Singapore); H. silenus (Vietnam); H. spinifer (Malaysia); J. cyaneus (Indonesia). Due to the difficulties involved in the identification of Heterometrus species, especially those occurring in Southeast Asia (Dupré, 1989), species are often advertised under false or erroneous names.

In addition to being sold alive as pets, many Southeast Asian Heterometrus are dried and mounted in glass cases or set in resin to be sold as curios. Heterometrus laoticus, H. silenus, and H. spinifer have been observed for sale as curios in the United States, U.K. and South Africa.

Unfortunately, wild populations of Heterometrinae are expected to be slow to repopulate after harvesting for the following reasons. Females have gestation periods up to 12 months (Subburam and Reddy, 1978a) and produce rather small broods (30–35) compared with other scorpions (Schultze, 1927; Mathew, 1956; Polis and Sissom, 1990; Shivashankar, 1994; Condevaux-Lanloy, 1996). Young are relatively altricial, spending several months in the maternal burrow before dispersing (Schultze, 1927; Shivashankar, 1994), thereby further protracting the period before a female can give birth to her next brood. Age to sexual maturity is 4–7 yr in these scorpions (Polis and Lourenço, 1986; Polis and Sissom, 1990), during which period most juveniles experience natural predation (including cannibalism).

These factors suggest that species of Heterometrinae are extremely vulnerable to overharvesting and, unlike Pandininae, three species of which receive CITES protection (IUCN, 1994; Lourenço and Cloudsley-Thompson, 1996), no species of Heterometrinae is currently CITES listed. The apparently restricted distributional ranges of many species provide further cause for concern, given that the remaining wild populations are threatened not only by overexploitation but also by continued habitat destruction (e.g., through deforestation). Many Heterometrinae appear to be restricted to virgin habitat (e.g., Schultze, 1927) and may thus be regarded as equilibrium species.

Remarks: Rossi (2016a) recognized the Asian scorpionids as a tribe of Pandininae Thorell, 1876, for reasons unspecified. The two lineages are sister taxa, with similar levels of taxonomic and morphological diversity, hence it is appropriate to elevate Heterometrini to the rank of subfamily, i.e., Heterometrinae Simon, 1879, stat. nov.

Heterometrus (Chersonesometrus) Couzijn, 1978: 330, type species by original designation: Buthus fulvipes C.L. Koch, 1837 [= Chersonesometrus fulvipes (C.L. Koch, 1837)]; 1981: 80, 83, 131, 133, 161, 169–171, 180, 182, 184, 186–188, 192, figs. 21, 57–59, 66b, 68, 69 (part); Tikader and Bastawade, 1983: 519, 520, 573 (part); Fet, 2000: 438; Francke, 1985: 8, 18; Prendini, 2000a: 44; Prendini et al., 2003: 222, 252, appendix 1; Bastawade, 2004: 290 (part); Bastawade et al., 2004: 48 (part); Kovařík, 2004: 1; Bastawade, 2005: 417; Bastawade and Borkar, 2008: 212, 219.

Heterometrus: Kovařík, 2004: 1, 2, 7, 49, 51, tables 1, 2 (part); 2009: 34, 35, table 1 (part); Mirza et al., 2012: 1, 2 (part); Kovařík et al., 2016: 96, 100 (part); Rossi, 2016a: 6–9, 15, 19, 20, 25 (part); Aswathi and Sureshan, 2017: 9847 (part).

Diagnosis: Species of Chersonesometrus may be separated from other Asian scorpionid genera as follows. The carapace is slightly to markedly dorsoventrally compressed, the lateral surfaces sloping gently (figs. 15, 16A, B, E, F), in all except three species of Chersonesometrus, C. fulvipes, C. madraspatensis, and C. shivashankari, in which the carapace is vaulted, the lateral surfaces sloping steeply (figs. 16C, D, 17), as in Gigantometrus, Heterometrus, Javanimetrus, two species of Deccanometrus, D. latimanus and D. xanthopus, two species of Sahyadrimetrus, S. mathewi and S. scaber, and all except three species of Srilankametrus, S. indus, S. pococki, and S. serratus. The carapace anterolateral margins converge gradually anteriorly (figs. 15–17) in Chersonesometrus but are subparallel anteriorly in Javanimetrus and Srilankametrus; the mediolateral margins diverge or converge slightly posteriorly (at the posterolateral sulci) in Chersonesometrus, but converge markedly posteriorly (at the posterolateral sulci) in Gigantometrus, Heterometrus, and three species of Sahyadrimetrus, S. kanarensis, S. mathewi, and S. scaber. The carapace rostrolateral margin is entire in Chersonesometrus but distinctly incised adjacent to the posterior lateral ocelli in Heterometrus. Anterocular extensions of the superciliary carinae are present in Chersonesometrus but absent in Srilankametrus. The median ocelli are relatively small, the distance between them equal to or greater than the width of an ocellus in Chersonesometrus but relatively large, the distance between them less than the width of an ocellus in Heterometrus; the median ocular tubercle is situated posteromedially, the distance from carapace anterior margin : carapace length (CAM:CL) 0.51–0.62 in Chersonesometrus, but anteromedially to medially, CAM:CL 0.40–0.50 in all other genera. The interocular suture is present in Chersonesometrus but absent in Heterometrus, Javanimetrus, Sahyadrimetrus, one species of Deccanometrus, and two species of Srilankametrus. The carapace posterior sutures are present, extending past the median ocular tubercle, and connected anteriorly to the posterior bifurcations of the interocular suture in Chersonesometrus; present, extending to the median ocular tubercle, and connected by a short cross-suture anterior to the postocular depression in Heterometrus; and absent in Javanimetrus, Sahyadrimetrus, and two species of Srilankametrus. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in some species of Chersonesometrus, as in Gigantometrus, Javanimetrus, Sahyadrimetrus, Srilankametrus, and most Deccanometrus, but subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in other species, as in Heterometrus. The pedipalp patella dorsomedian carina of the female is entirely to predominantly granular or costate in Chersonesometrus, but absent or obsolete in Heterometrus, Javanimetrus, all except one species of Sahyadrimetrus, S. rugosus, and all except one species of Srilankametrus, S. caesar. The patella retromedian carinae of the female are granular or costate (figs. 82, 85, 93, 103, 112, 115) in Chersonesometrus, but absent or obsolete in Gigantometrus, Heterometrus, Javanimetrus, all except three species of Deccanometrus, D. bengalensis, D. obscurus, and D. phipsoni, all except two species of Srilankametrus, S. gravimanus and S. yaleensis, and three species of Sahyadrimetrus, S. kanarensis, S. mathewi, and S. scaber. The prominent spiniform granule of the patella proventral carina is absent (figs. 82, 85, 89, 93, 96, 99, 103, 107, 112, 115) in Chersonesometrus but present in Heterometrus. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are granular (figs. 82, 85, 93, 103, 112, 115) in all species of Chersonesometrus except C. nathanorum, in which the surfaces are smooth or nearly so, as in Gigantometrus, Heterometrus, Javanimetrus, Srilankametrus, all species of Deccanometrus except D. obscurus and D. phipsoni, and all species of Sahyadrimetrus except S. rugosus. The distance between the chela manus dorsomedian and promedian carinae or setal rows (DMC–PMC) is slightly to markedly greater than the distance between the promedian and proventral carinae or setal rows (PMC–PVC) in Chersonesometrus whereas the DMC–PMC is similar to the PMC–PVC in Heterometrus. The dorsomedian carina is continuous to the proximal edge of the manus in Chersonesometrus but becomes obsolete proximally in Heterometrus, Javanimetrus, Sahyadrimetrus, Srilankametrus, and all species of Deccanometrus except D. obscurus and D. phipsoni. The proximal half of the chela manus dorsal secondary carina, distal half of the subdigital carina and distal third of the digital carina are closely adjacent in Chersonesometrus whereas the dorsal secondary, subdigital and digital carinae (or setal rows) are well separated along their entire length in all the other genera except Gigantomentrus. The dorsal secondary and subdigital carinae of the male are present and entirely to predominantly granular or costate (figs. 87, 97, 100, 104, 108, 113A) in all species of Chersonesometrus, except C. bastawadei and C. fulvipes, in which the carinae are absent or obsolete (fig. 90), as in Deccanometrus, Heterometrus, Javanimetrus, Sahyadrimetrus, and two species of Srilankametrus, S. indus and S. pococki. The dorsal secondary carina comprises a double row of separate or confluent granules in Chersonesometrus whereas it comprises a single row in Srilankametrus. The chela manus retromedian carina is more pronounced than the digital carina (figs. 83, 86, 87, 90, 91, 94, 97, 98, 100, 104, 105, 108, 109, 113, 116) in Chersonesometrus whereas the digital and retromedian carinae are similarly developed in all the other genera. The retromedian carina of the male is entirely to predominantly costate in Chersonesometrus but entirely to predominantly granular in Gigantometrus, Srilankametrus caesar and S. serratus, and absent or obsolete in Deccanometrus, Heterometrus, Javanimetrus, Sahyadrimetrus, Srilankametrus indus, and S. pococki. The depression in the dorsal surface of the chela manus, proximal to the fixed finger of the adult male, is absent or obsolete (figs. 87, 90, 97, 100, 104, 108, 113A) in Chersonesometrus but present and distinct in Heterometrus. The chela manus dorsal surface is usually without reticulation in Chersonesometrus but shallowly reticulate in Javanimetrus, Sahyadrimetrus, all except two species of Deccanometrus, D. obscurus and D. phipsoni, and three species of Heterometrus, H. glaucus, H. longimanus and H. thorellii; the dorsal surface is finely to coarsely granular in Chersonesometrus but smooth in Heterometrus, Javanimetrus, two species of Sahyadrimetrus, S. kanarensis and S. tikaderi, and one species of Deccanometrus, D. ubicki. The chela manus retrolateral intercarinal surfaces are granular in Chersonesometrus but smooth or nearly so in Heterometrus, Javanimetrus, and some species of Sahyadrimetrus. The chela manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane, in all except four species of Chersonesometrus, C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis, in which it is angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina, as in Javanimetrus and two species of Srilankametrus, S. indus and S. pococki. The pro- and retrolateral surfaces of the tibiae of legs I and II each bear a row of two or three spiniform macrosetae in Chersonesometrus and scattered, setiform macrosetae, not arranged in a definite row, in Heterometrus. Macroseta st on the retroventral surface of the basitarsus of leg I is spiniform (figs. 43–45) in Chersonesometrus but usually setiform in Heterometrus, and sb on the retroventral surface of the basitarsus of leg III is usually setiform in Chersonesometrus but spiniform in Javanimetrus and Sahyadrimetrus. The pseudonychium (dactyl) of the telotarsi of legs I–IV is reduced and rounded in Chersonesometrus but prominent and acuminate in Heterometrus. The pectinal first proximal median lamella (scape) of the female is distinctly angular, > 90° but < 180° (figs. 29–31) in Chersonesometrus but straight or shallowly curved in Srilankametrus. The lengths of metasomal segments I and II are approximately equal to or less than their respective widths (figs. 58–63) in Chersonesometrus but markedly greater than their respective widths in Gigantometrus. The ventrosubmedian and ventrolateral carinae are granular or costate-granular on metasomal segment IV only and costate on segments I–III (figs. 59, 60, 62, 63) in Chersonesometrus but granular on segments I–IV, II–IV (costate on I), or III and IV (costate on I and II) in Gigantometrus. The dorsosubmedian carinae of metasomal segment V are vestigial (figs. 58, 61) in Chersonesometrus but partial in Gigantometrus and absent in Heterometrus. The telson is paler than metasomal segment V in Chersonesometrus but as dark as segment V in most species of Srilankametrus and some species of Deccanometrus and Heterometrus. The width of the telson vesicle is approximately equal to or less than the width of metasomal segment V in the female of Chersonesometrus but greater than the width of segment V in the female of Gigantometrus and the vesicle is globose in Chersonesometrus but elongate in Heterometrus, Javanimetrus, Sahyadrimetrus, and all except three species of Srilankametrus, S. indus, S. pococki, and S. serratus.

Included Species: As redefined in the present contribution, Chersonesometrus accommodates 10 species, five of which were formerly assigned to subgenus Chersonesometrus of Heterometrus by various authors (Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000), whereas four are newly described, recovered as a monophyletic group by phylogenetic analysis of morphological characters and DNA sequences from the nuclear and mitochondrial genomes (fig. 10): Chersonesometrus bastawadei, sp. nov.; Chersonesometrus beccaloniae (Kovařík, 2004), comb. nov.; Chersonesometrus fulvipes (C.L. Koch, 1837), comb. nov.; Chersonesometrus hendersoni, sp. nov.; Chersonesometrus madraspatensis (Pocock, 1900), comb. nov.; Chersonesometrus nathanorum, sp. nov.; Chersonesometrus pelekomanus (Couzijn, 1981), comb. nov. et stat. rev.; Chersonesometrus shivashankari, sp. nov.; Chersonesometrus tristis (Henderson, 1919), comb. nov.; and Chersonesometrus wroughtoni (Pocock, 1899), comb. nov.

Distribution: Chersonesometrus is endemic to India (figs. 1, 79, 80) and has been recorded in the states of Andhra Pradesh, Gujarat, Haryana, Karnataka, Madhya Pradesh, Maharashtra, National Capital Territory of Delhi, Odisha, Rajasthan, Tamil Nadu, Telangana, Uttar Pradesh, and the union territory of Puducherry (table 1). A single record of C. fulvipes from Sindh, southeastern Pakistan, on the border with India (Kovařík, 2009) requires confirmation (Tahir and Prendini, 2014).

Ecology: The species of Chersonesometrus for which data are available, occur in semiarid savanna or deciduous woodland, mostly on the Deccan Plateau or in the rain shadow of the Western Ghats (Sahyadri), at elevations of 150–1020 m above sea level. Based on morphology and available habitat data, all species of the genus appear to be pelophilous and fossorial, constructing shallow burrows under stones, or semilithophilous, inhabiting rock crevices or scrapes under stones.

Conservation Status: Chersonesometrus fulvipes and C. tristis, traded as Heterometrus mysorensis, are occasionally harvested for the commercial trade in exotic pets.

Remarks: Chersonesometrus Couzijn, 1978, stat. nov., originally created as a subgenus of Heterometrus, and subsequently synonymized with the latter (Kovařík, 2004), is hereby revalidated and elevated to the rank of genus.

Palamnaeus fulvipes madraspatensis: Pocock, 1900a: 89 (misidentification, part).

Type Material: INDIA: Karnataka: Bagalkot Distr.: Holotype ♀, paratypes: 7 ♀, 5 subad. ♂, 1 subad. ♀, 11 juv. ♂, 4 juv. ♀(AMNH), 2 juv. ♂, 1 juv. ♀ (AMCC [LP 16787]), Naganath Temple, NW of Navilahole, 15°54′N 75°44′E, 541 m, 23.xi.2009, R. Datta, narrow valley between ridges, degraded savanna with plantations, mixed broadleaf woodland with moderate to sparse grass layer on sandy loam, many loose or embedded quartzite stones, in shallow burrows (ca. 10–40 cm deep) or scrapes under stones; 1 subad. ♂ paratype (AMCC [LP 16785]), Badami, Bhootnath Temple, Badami–Mahakoota Path, 15°55′N 75°41′E, 607 m, 23.xi.2009, R. Datta, sandstone plateau, fairly degraded, mixed savanna on sandy-loam with many sandstone outcrops, loose and embedded stones, in shallow burrow (ca. 10 cm deep) under stone.

Etymology: The specific epithet is a patronym honoring Deshbhushan B. Bastawade, formerly of the Zoological Survey of India, Poona, and widely known as D.B. Bastawade, who contributed significantly to knowledge of Indian scorpions (e.g., Tikader and Bastawade, 1983) and other arachnids over the course of his career.

Diagnosis: Chersonesometrus bastawadei may be separated from other species of Chersonesometrus as follows. The carapace is slightly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 16A), in C. bastawadei, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The frontal lobes and medial region of the carapace interocular surface are granular with smooth areas in the female (fig. 16A) of C. bastawadei whereas the interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female of all other species except C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are granular in C. bastawadei but smooth or nearly so in all other species except C. fulvipes and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. bastawadei, but subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. beccaloniae, C. hendersoni, C. nathanorum, C. pelekomanus, and C. tristis. The pedipalp patella dorsal surface is convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina (fig. 82), in C. bastawadei but flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. bastawadei but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. bastawadei but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are granular (fig. 82) in C. bastawadei but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. bastawadei but immaculate and paler than the femur and patella in C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and curved, i.e., slightly to markedly convex, in the female of C. bastawadei, flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface in the male or aligned with the proximal margin of the condyle (articulation with patella) in the female (fig. 83) of C. bastawadei; moderately curved and aligned with the proximal margin of the retrolateral surface in the male or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. beccaloniae and C. nathanorum; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is greater than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. bastawadei, but greater than the DSC–DC in the male of C. beccaloniae, C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male are absent or obsolete in C. bastawadei but entirely to predominantly granular or costate in all other species except C. fulvipes. The digital carina is absent or obsolete (fig. 83) in C. bastawadei, but entirely to predominantly granular in C. beccaloniae, C. hendersoni, C. pelekomanus, C. tristis, and C. wroughtoni, and entirely to predominantly costate in C. nathanorum. The manus dorsal intercarinal surface is without reticulation in C. bastawadei but shallowly reticulate in C. nathanorum. The manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in C. bastawadei but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The legs are pale or very lightly infuscate (fig. 81A, B) in C. bastawadei but very dark or heavily infuscate in C. beccaloniae, C. nathanorum, C. pelekomanus, C. tristis, and C. wroughtoni. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 44A–D) in C. bastawadei but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are granular in the male and the lateral surfaces granular in the female of C. bastawadei, whereas the mesial surfaces are smooth in the male of C. beccaloniae, C. hendersoni, C. pelekomanus, and C. wroughtoni, the mesial surfaces smooth in the male and the lateral surfaces smooth in the female of C. tristis, and the lateral surfaces smooth in the female of C. nathanorum. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. bastawadei and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 58A) in C. bastawadei but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are smooth in the male and granular in the female (fig. 60A) of C. bastawadei but granular in the male and smooth in the female of C. beccaloniae and C. nathanorum, granular in the male of C. hendersoni, C. madraspatensis, C. pelekomanus, and C. tristis, and smooth in the female of C. wroughtoni. The dorsolateral carinae of metasomal segment V are strong and continuous in C. bastawadei but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is granular in the male and female of C. bastawadei, smooth in the female of C. wroughtoni, and smooth in the male and female of all other species except C. hendersoni and C. pelekomanus.

Description: The following description is based on the holotype and paratypes. Meristic data are presented in table 3. The adult male is unknown, hence measurements are presented only for the adult female (table 4).

Total length: Adult medium sized, maximum length, measured from anterior margin of carapace to tip of aculeus, 107 mm (100–115 mm, n = 5) (♀).

Color: Chelicerae, dorsal surfaces bicolored, mostly to entirely immaculate prodorsal half of manus paler than infuscate retrodorsal half of manus and fingers; manus markedly paler than carapace interocular surface. Carapace, pedipalps, tergites, and metasoma entirely infuscate, uniformly dark reddish black. Sternites III–VI immaculate or partially infuscate, VII mostly or entirely infuscate, dark olive to brownish black becoming darker posteriorly. Coxosternal region infuscate, brownish, maxillary lobes darker. Legs lightly infuscate, uniformly yellow to light brown contrasting with carapace, pedipalps, tergites, and metasoma (fig. 81A, B). Genital opercula and pectines uniformly pale cream. Metasomal segments I–V similar to one another; carinae infuscate darker than intercarinal surfaces; dorsal intercarinal surfaces similar to tergites; ventral intercarina surfaces darker than sternites. Telson vesicle immaculate, light yellowish, markedly paler than metasomal segment V; aculeus black.

Chelicerae: Movable finger prodistal (DI) and retrodistal (DE) teeth unequal, with DE tooth considerably smaller than DI tooth, aligned longitudinally and not opposable. Fingers and manus, proventral surfaces with long, dense vestiture of macrosetae. Coxae, prodorsal surfaces without stridulatory setae (scaphotrix); promedian surfaces without chemoreceptive lamelliform setae (trichocopae).

Carapace: Carapace slightly dorsoventrally compressed, lateral surfaces sloping gently. Anterior margin straight, frontal lobes evenly rounded or truncate, with deep median notch, without median projection or median depression (fig. 16A). Anterolateral margins converging gradually anteriorly, without distinct notch adjacent to posterior lateral ocelli; mediolateral margins diverging or converging slightly posteriorly (at posterolateral sulci). Anterior width of posterior width, 64% (62%–58%, n = 5) (♀); posterior width of length, 94% (91%–97%, n = 5) (♀). Three pairs of lateral ocelli (MLMa, PLMa, PDMi), all similar in size, two anterior pairs situated anteriorly, well separated from posterior pair, situated laterally. Median ocelli slightly larger than lateral ocelli, distance between ocelli greater than or equal to width of ocellus; median ocular tubercle situated posteromedially, distance from anterior carapace margin, 52% (51%–53%, n = 5) (♀). Median longitudinal sulcus narrow, suturiform; continuous from median notch to interocular sulcus; with obsolete anterior furcation; without anterocular depression. Anterior furcated sulci diverging broadly from anterior edge of median longitudinal sulcus. Median ocular tubercle raised, superciliary carinae higher than ocelli, extended anteriorly, subparallel. Interocular sulcus present. Circumocular depressions completely encircling median ocular tubercle, converging anteriorly. Posteromedian and posteromarginal sulci deep. Paired mediolateral and posterolateral sulci shallow. Median longitudinal suture continuous from median notch to median ocular tubercle, equally strong along entire length; not extending to anterior margin of carapace, terminating at or posterior to median notch. Anterior furcated sutures absent or obsolete. Anterocular furcated sutures absent. Interocular suture present, slender. Posterior sutures present but indistinct, converging on median ocular tubercle; connected anteriorly to posterior furcations of interocular suture and extending anteriorly beyond median ocular tubercle. Frontal lobes and medial region of interocular surface finely granular with smooth areas (fig. 16A); anterolateral, mediolateral, and posterolateral surfaces uniformly finely granular; posteromedian surface smooth.

Pedipalps: Pedipalp short, pedipalp femur length: posterior carapace width ratio (FL:PCW) < 0.77, femur length: carapace length ratio (FL:CL) < 0.74. Femur width of length, 49% (47%–51%, n = 5) (♀). Retrodorsal carina complete, costate-granular, similar to or more strongly developed than prodorsal carina. Dorsal secondary carina comprising scattered granules, becoming obsolete distally. Dorsomedian carina vestigial, reduced to prominent granule demarcated by conspicuous macroseta. Prodorsal carina complete, granular. Promedian carina complete, comprising row of spiniform or subspiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Proventral carina complete, granular, as strongly developed as or more strongly developed than retroventral carina. Ventromedian and secondary accessory carinae absent. Retroventral carina extending partly or entirely to distal edge of segment. Retromedian carinae absent, indicated only by macrosetal rows. Dorsal and prolateral intercarinal surfaces finely granular; ventral and retrolateral surfaces smooth. Patella width of length, 51% (49%–54%, n = 5) (♀). Dorsal surface convex, axis of dorsomedian carina dorsal to axis of retrodorsal carina. Dorsomedian carina entirely to predominantly costate (fig. 82). Retrodorsal carina predominantly costate-granular, more strongly developed than granular retromedian carinae. Retroventral carina distinct, costate to costate-granular. Promedian carina and promedian process absent. Other carinae absent. Dorsal and prolateral intercarinal surfaces finely granular, other surfaces smooth. Chela short, broad (fig. 83); manus height of width, 82% (78%–85%, n = 5) (♀); manus length along retroventral carina of manus width, 89% (89%–93%, n = 5) (♀); manus length along retroventral carina of movable finger length, 72% (69%–75%, n = 5) (♀). Chela sparsely setose. Manus dorsal surface (between dorsomedian and digital carinae) flat (♂) or curved, i.e., slightly to markedly convex (♀); proximal margin (lobe) of dorsal surface moderately curved and proximal to proximal margin of retrolateral surface (♂) or aligned with proximal margin of condyle (articulation with patella) (♀). Dorsomedian carina obsolete, costate on fixed finger and comprising regular row of spiniform granules along entire length of manus. Dorsal secondary, subdigital, and digital carinae absent or obsolete. Maximum distance between dorsomedian carina and setal row demarcating dorsal secondary carina (DMC–DSC) greater than maximum distance between setal rows demarcating dorsal secondary and digital carinae (DSC–DC). Retromedian carina complete, predominantly costate, more strongly developed than digital carina. Retrolateral secondary and secondary accessory carinae vestigial, proximal to condyle of movable finger. Retroventral and ventromedian carina distinct, costate, but retroventral carina more strongly developed. Proventral and promedian carinae similarly developed, each comprising row of spiniform granules and associated macrosetae. Prodorsal carina obsolete, comprising few spiniform granules and associated macrosetae. Distance between dorsomedian and promedian carinae (DMC–PMC) slightly to markedly greater than distance between promedian and proventral carinae (PMC–PVC). Manus, dorsal and retrolateral intercarinal surfaces densely and coarsely granular except for smooth depression at base of fixed finger; ventral intercarinal surface granular; prolateral intercarinal surfaces sparsely finely and coarsely granular, more densely dorsally and distally. Manus ventral surface flat, axes of retroventral and ventromedian carinae in approximately same plane. Fixed and movable fingers, intercarinal surfaces smooth; median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three situated on lobes; first (proximal) lobe of fixed finger and second lobe of movable finger larger than others, with correspondingly deeper notches in movable and fixed fingers, respectively; terminal denticles of fingers interlocking unevenly when closed, movable finger displaced retrolaterally; distinct notch near tip of fixed finger to accommodate terminal denticle of movable finger.

Trichobothria: Orthobothriotaxic, Type C, with the following segment totals (n = 20): femur, 3 (1 d, 1 i, 1 e); patella, 19 (2 d, 1 i, 3 v, 13 e, comprising 3 et, 1 est, 2 em, 2 esb, 5 eb); chela, 26 (manus, 16, comprising 2 D, 10 E, 4 V; fixed finger, 10, comprising 4 d, 4 e, 2 i). Total count of trichobothria per pedipalp: 48. Femur, i situated on dorsal surface. Patella, d₂ situated on internal surface, slightly closer to d₁ than to i; distance v₂–v₃ ca. one-third distance v₁–v₂. Chela, distance et–est less than half distance est–esb; distance est–esb greater than half distance esb–eb; est distal to dst; V₃ situated in proximal third of manus V₃, closer to V₄ than V₂.

Legs: First pair of maxillary lobes (coxapophyses) tapering anteriorly, longer than and encircling second pair. Stridulatory organs, comprising “rasp” (granular tubercles) and “scraper” (stridulatory setae or scaphotrix), present on opposing surfaces of coxae of first pair of legs and pedipalps, respectively. Legs I–IV acarinate, except femora, with distinct pro- and retrolateral surfaces demarcated by partial, granular prodorsal and complete, costate-granular proventral carinae. Femora, patellae, and tibiae, pro- and retrolateral surfaces each with scattered setiform macrosetae. Tibiae, prolateral surfaces, without spiniform macrosetae; I and II, retrolateral surfaces, each with two spiniform (t, st) macrosetae; III and IV, retrolateral surfaces, without spiniform macrosetae. Basitarsi I–IV, slightly dorsoventrally compressed; retrolateral margins similar, unmodified, rounded; prolateral pedal spurs present (fig. 44A–D); retrolateral pedal spurs absent. Basitarsi, pro- and retrolateral surfaces, each with scattered long and short setiform macrosetae, and spiniform macrosetae, more numerous on I and II than III and IV. Basitarsi, spiniform macrosetae, I, retrolateral: t, sb; retroventral: t, st, sb; proventral: t, st; II, retrolateral: t, sb; retroventral: t, st, sb; proventral: t, st; III, retrolateral: t, sb; retroventral: t, st; proventral: t; prolateral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: t, st. Telotarsi I–IV short, stout and distally broadened in dorsal and lateral views. Laterodistal lobes rounded. Dorsomedian lobes approximately equal to laterodistal lobes; each terminating in single setiform macroseta. Telotarsi, pro- and retrolateral surfaces, each with long and short, scattered macrosetae, not arranged in definite row. Telotarsi each with pro- and retroventral rows of spiniform macrosetae, two inserted on laterodistal lobes; macrosetal counts in pro- and retroventral rows similar on I and II, and on III and IV (n = 20): I, proventral: 3 (3 or 4), retroventral: 6 (5 or 6); II, 4 (3 or 4), 6 (5 or 6); III and IV, 4, 6. Telotarsal ungues short, curved, equal to subequal; pseudonychium (dactyl) reduced, rounded.

Sternum: Shape subpentagonal (fig. 30A). Median longitudinal sulcus shallow anteriorly, deep and narrow posteriorly.

Genital operculum: Genital opercula suboval, completely divided longitudinally, partially overlapping, genital papillae present (♂); subcordate, fused, genital papillae absent (♀) (fig. 30A).

Hemispermatophore: Unknown.

Pectines: Distal edge extending past distal edge of coxa IV but not reaching distal edge of trochanter IV (fig. 30A). First proximal median lamella (scape) of each pecten with mesial margin angular, approximately 90° (♂) or obtusely angular, greater than 90° but less than 180° (♀). Pectinal teeth straight and elongate (♂) or shorter and curved (♀), present along entire posterior margin; sinistral/dextral tooth count, 17/17 (16–18/16–18, n = 12) (♂), 15/15 (15 or 16/15–17, n = 12) (♀). Fulcra smooth proximally but densely setose (microsetae only) distally.

Mesosoma: Tergites each with shallow pair of submedian depressions, without obsolete median carina. Pretergites smooth and glabrous. Posttergites finely granular (♂) or smooth and glabrous (♀) medially, unevenly finely and coarsely granular to predominantly coarsely granular laterally. Sternites IV–VI, each with paired longitudinal depressions prolateral to spiracles, absent on VII. Surface, sternites III–VII, smooth; VII with pair of weakly developed, costate ventrolateral carinae only, without posteromarginal carina. Sternite VII, length of width, 65% (61%–72%, n = 5) (♀).

Metasoma and telson: Metasomal segments I–V progressively increasing in length, decreasing in width; segment V, width of segment I, width, 73% (72%–76%, n = 5) (♀). Metasoma robust, width of length, segment I, 90% (85%–98%, n = 5) (♀); II, 72% (69%–74%) (♀); III, 64% (62%–66%) (♀); IV, 53% (51%–54%) (♀); V, 39% (38%–40%) (♀). Telson vesicle, width of metasomal segment V, width, 94% (90%–95%, n = 5) (♀); vesicle globose, height of length, 55% (53%–57%, n = 5) (♀); dorsal surface flat; ventral surface evenly curved. Aculeus relatively long, gently curved, length of vesicle length, 57% (56%–60%, n = 5) (♀). Length metasoma and telson, of total length, 55% (54%–56%, n = 5) (♀). Dorsosubmedian carinae, segments I–IV, distinct, complete, granular or costate-granular, posterior spiniform granules obsolete (fig. 58A); V, vestigial. Dorsolateral carinae, segments I–V, distinct, complete, costate-granular on I–IV, costate-granular to granular on V. Median lateral carinae, segment I, incomplete, reduced to granular row in posterior half, diagonally oriented between dorsolateral and ventrolateral carinae (fig. 59A); II–IV, obsolete row of granules posteriorly; V, incomplete, granular row, restricted to anterior two-thirds. Ventrolateral and ventrosubmedian carinae, I–IV, distinct, complete, costate on I–III, granular or costate-granular on IV, more strongly developed on III and IV than on I and II (fig. 60A). Ventrolateral carinae, V, distinct, complete, comprising spiniform granules, diverging posteriorly, terminal granule slightly larger than preceding granules. Ventrosubmedian carinae, segment V, vestigial, each reduced to one or two isolated spiniform granules, demarcated by conspicuous macrosetae. Ventromedian carina, segment V comprising single row of spiniform granules, unmodified posteriorly. Anal arch, segment V, dorsal carina, costate; ventral carina comprising subspiniform granules. Dorsal intercarinal surfaces, segments I–V, granular. Lateral intercarinal surfaces, segments I–IV, granular, V, smooth. Ventral intercarinal surfaces, segments I–III, smooth, IV and V, smooth (♂) or granular (♀). Ventral surface, lateral aspect, segment IV, shallowly convex. Telson vesicle, dorsal and lateral surfaces smooth; ventral surface with four longitudinal carinae, each comprising spiniform granules.

Distribution: This species is endemic to India and presently recorded only from the state of Karnataka (fig. 80, table 1). Its distribution is allopatric with the morphologically similar C. fulvipes, C. madraspatensis, C. shivashankari, and C. wroughtoni.

Ecology: The known locality records of this species occur in semiarid savanna dominated by Acacia Martius, 1829, Euphorbia Linnaeus, 1753, and broadleaf bushes (Dichrostachys (DC.) Wight and Arnott, 1834, Grewia Linnaeus, 1753, Ziziphus Miller, 1754), with sparse grass on rocky sandstone or quartzite hills, at elevations of 540–610 m. The specimens for which data are available were collected from shallow burrows or scrapes (10–40 cm deep) under stones or boulders, embedded in hard, sandy loam. The habitat and habitus are consistent with the pelophilous, fossorial and semilithophilous ecomorphotypes (Prendini, 2001b). This species is sympatric with Charmus indicus Hirst, 1915, Hottentotta rugiscutis (Pocock, 1897) and Lychas tricarinatus (Simon, 1884).

Heterometrus indus: Roewer, 1929: 612 (misidentification, part).

Heterometrus liurus: Roewer, 1929: 613 (misidentification, part).

Heterometrus phipsoni: Roewer, 1929: 612 (misidentification, part).

Heterometrus (Chersonesometrus) palekomanus: Indra, 2001: 56, 58 (misidentification, part).

Heterometrus beccaloniae Kovařík, 2004: 1, 4, 6, 7, 51, 52, tables 1–3, fig. 9; 2009: 34–36, 47, 49, 73, 101, tables 1, 2, figs. 5–7, 217; Mirza et al., 2012: 1, 3, 6, 7, table 1.

Heterometrus (Chersonesometrus) pelekomanus: Indra, 2009: 141 (misidentification, part).

Heterometrus atrascorpius Mirza et al., 2012: 1–8, table 1, figs. 1–10; syn. nov.

Type Material: INDIA: Tamil Nadu: Nilgiris Distr.: Heterometrus atrascorpius: Holotype: 1 [subad.] ♀ (BNHS Sc-55), Bokkapuram village near Masinagudi, 11°33′N 76°40′E, 912 m, 25.i.2011, G. Desouza and N.S. Achythuan; paratypes: 1 [imm.] ♂ (BNHS Sc-56), same data except: R. Sanap and Z. Mirza, 1 imm. paratype ♀ (BNHS Sc-57), same data except: D. Joshi and S. Chikane. Heterometrus beccaloniae: Paratypes: 1 imm., 7 juv. (FKPC), Nilgiri hills, Mudumalai National Park [11°34′N 76°32′E], 6.v.2000, Rolčík. Vellore Distr.: Heterometrus beccaloniae: Holotype ♀ (FKPC), Kavalore [Kavalur, 12°35′N 78°49′E], Javadu Hills [Jawadhu Hills], 2000 ft., v.2003, T.R.S.N.

Diagnosis: Chersonesometrus beccaloniae may be separated from other species of Chersonesometrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 15A, B), in C. beccaloniae, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female (fig. 15B) of C. beccaloniae whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the female of C. bastawadei, C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are usually smooth in C. beccaloniae but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. beccaloniae but unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, and C. wroughtoni. The pedipalps of the adult male are long, with femur length: posterior carapace width ratio (FL:PCW) ≥ 0.80 and femur length: carapace length ratio (FL:CL) ≥ 0.76 (fig. 84A, B) in C. beccaloniae but short, with FL:PCW < 0.77, FL:CL < 0.74 in all other species except C. madraspatensis and C. tristis. The pedipalp patella dorsal surface is flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane (fig. 85), in C. beccaloniae but convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina, in all other species except C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. beccaloniae but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. beccaloniae but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are granular (fig. 85) in C. beccaloniae but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. beccaloniae but immaculate and paler than the femur and patella in C. shivashankari. The chela of the adult male is moderately to densely setose in C. beccaloniae but sparsely setose in C. fulvipes, C. madraspatensis, and C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and curved, i.e., slightly to markedly convex, in the female of C. beccaloniae, flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and aligned with the proximal margin of the retrolateral surface in the male (fig. 87), or aligned with or proximal to the proximal margin of the retrolateral surface in the female (fig. 86) of C. beccaloniae; moderately curved and proximal to the proximal margin of the retrolateral surface in the male, or aligned with the proximal margin of the condyle (articulation with patella) in the female of C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, C. tristis, and C. wroughtoni; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is similar to the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. beccaloniae, but greater than the DSC–DC in the male of all other species except C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male (fig. 87) are entirely to predominantly granular in C. beccaloniae, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is entirely to predominantly granular in C. beccaloniae, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The manus dorsal intercarinal surface is without reticulation in C. beccaloniae but shallowly reticulate in C. nathanorum. The manus ventral surface is angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. beccaloniae but flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in all other species except C. hendersoni, C. pelekomanus, and C. tristis. The legs are very dark or heavily infuscate (fig. 84) in C. beccaloniae but pale or very lightly infuscate in C. bastawadei, C. fulvipes, C. hendersoni, C. madraspatensis, and C. shivashankari. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 43A–D) in C. beccaloniae but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are smooth in the male and the lateral surfaces granular in the female of C. beccaloniae, whereas the mesial surfaces are granular in the male of C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari, the mesial surfaces granular in the male and the lateral surfaces smooth in the female of C. nathanorum, and the lateral surfaces smooth in the female of C. tristis. The ventral surface of mesosomal sternite VII bears two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae and a pair of weakly developed ventrolateral carinae in all other species except C. hendersoni and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV in the male but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in the female (fig. 58B) of C. beccaloniae, whereas the dorsosubmedian carinae are granular or costate-granular on segments I–IV in all other species, except C. tristis. The ventral intercarinal surfaces of segment IV are granular in the male and smooth in the female (fig. 60B) of C. beccaloniae but smooth in the male and granular in the female of C. bastawadei, C. fulvipes, and C. shivashankari, smooth in the male of C. wroughtoni, and granular in the female of C. hendersoni, C. madraspatensis, C. pelekomanus, and C. tristis. The dorsolateral carinae of metasomal segment V are strong and continuous in C. beccaloniae but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is smooth in the male and female of C. beccaloniae, granular in the male and female of C. bastawadei, C. hendersoni, and C. pelekomanus, and granular in the male of C. wroughtoni.

Distribution: This species is endemic to India and recorded from the Javadu Hills, the Nilgiri Hills, and the Yelagiri Hills in the state of Tamil Nadu (fig. 80, table 1).

Ecology: The known locality records of this species occur in semiarid savanna and deciduous woodland dominated by thorn bushes (Acacia and Ziziphus) and sparse grass on rocky granite hills, at elevations of 305–1020 m (Mirza et al., 2012). The specimens for which data are available were collected from shallow burrows or scrapes (10–20 cm long) under large stones or boulders, embedded in hard clayey or gravelly-loam soils. Burrow entrances, below stones, are crescent shaped (Mirza et al., 2012). The habitat and habitus are consistent with the pelophilous, fossorial and semilithophilous ecomorphotypes (Prendini, 2001b). This species is sympatric with the buthids, Hottentotta rugiscutis, Isometrus sankeriensis Tikader and Bastawade, 1983, and Lychas tricarinatus (Mirza et al., 2012).

Remarks: This species is newly accommodated in Chersonesometrus, stat. nov. et stat. rev., based on a phylogenetic analysis (fig. 10).

The description of H. beccaloniae was based on a female holotype. The diagnosis of H. beccaloniae compared it to H. wroughtoni, for which the adult male is unknown, but not to H. phipsoni, which was described from an adult male, occurs in geographical proximity, and resembles H. beccaloniae in several respects, including dark coloration; length, 115 mm in the type of H. beccaloniae compared with 101 mm in the type of Scorpio phipsoni; pectinal tooth count, 14–16 in H. beccaloniae compared with 15/15 in S. phipsoni; and pro- and retroventral spiniform macrosetal counts on the leg telotarsi, 5–6/6–7 on legs III and IV in H. beccaloniae compared with 5/7 on I–IV in S. phipsoni. Nevertheless, based on comparison of type and nontype material, including an adult male of C. beccaloniae, during the present investigation, C. beccaloniae is evidently distinct from both C. wroughtoni and D. phipsoni, in which the pedipalp chela manus is much more granular, among other differences recounted in the key and diagnoses.

Heterometrus atrascorpius was distinguished from H. beccaloniae by the following characters: moderate size, 91.7 mm; coloration dark blackish brown, with the exception of the vesicle, which is yellowish white, and the leg tarsi, which are paler; pectines, sternum, and pectinal basal piece yellowish brown; pedipalp chela manus tuberculate with carina; patella and femur dorsally tuberculate; pedipalp femur longer than twice as wide; manus length/width ratio 3.22; pectines with 16–16 teeth. However, as noted by Tahir and Prendini (2014), it is evident that none of the type specimens are adult. The holotype illustrated in figures 1–9 (Mirza et al., 2012: 4, 5) is subadult. Consequently, the diagnostic characters (coloration, granulation, meristic variation) presented to justify this putative new species are unreliable and the comparisons made with other species, described on the basis of adults, invalid. Based on both the description and the type locality, H. atrascorpius is assessed to be conspecific with C. beccaloniae and the following new synonym presented: Heterometrus atrascorpius Mirza et al., 2012 = Chersonesometrus beccaloniae (Kovařík, 2004), syn. nov.

Material Examined: INDIA: Tamil Nadu: Nilgiris Distr.: Ootacamund [Ooty, 11°25′N 76°42′E], 1 ♂ (ZMH); Vibudhi Malai Murugan Temple, Bokkapuram, 11°33′N 76°39′E, 1023 m, 27.xi.2004, S. Basi, rocky hill, savanna-grassland on hard clay, many granite stones and boulders, in shallow scrapes under stones, 1 ♀, 1 subad. ♂, 7 juv. ♂, 6 juv. ♀ (AMNH), 2 juv. ♂ (AMCC [LP 13133]). Vellore Distr.: Javadu Hills, S of Madanur on Highway 46 (Ambur–Vellore), 12°51′N 78°49′E, 306 m, 30.xi.2004, S. Basi, dense broadleaf savanna-woodland at base of large mountain with scattered granite stones, in shallow scrape (ca. 20 cm long), under large stone in gravelly loam, 1 ♀(AMNH); Yelagiri Hills, Mangalam Road, Yelagiri, 12°35′N 78°39′E, 1000 m, 29.xi.2004, S. Basi, savanna-grassland on rocky hillside with many large granite stones, burrow entrances under very large stones in hard gravelly loam, 1 ♀, 1 juv. ♂, 3 juv. ♀, 1 ♀ carapace and sinistral pedipalp segments (AMNH), 1 juv. ♂, 1 juv. ♀ (AMCC [LP 13134]); Yelagiri Hills, Yelagiri Road, 12°36′N 78°37′E, 773 m, 30.xi.2004, S. Basi, steep rocky slope, savanna-woodland with many granite stones embedded in hard gravelly loam, in shallow scrape (ca. 10 cm long) under stone, 1 subad. ♂ (AMNH).

Buthus fulvipes C.L. Koch, 1837: 45, 46, pl. CXXI, fig. 278; 1850: 87; Couzijn, 1981: 133, 136; Kovařík, 2004: 2, 15; 2009: 38.

Pandinus fulvipes: Karsch, 1884: 67, 68; Thorell, 1888: 329, 338.

Scorpio fulvipes: Pocock, 1893: 304–307, 309 (part); Kraepelin, 1894: 31, 43–46, 52, 59–61, 246, pl. I, figs. 22, 28, 37 (part).

Heterometrus fulvipes: Kraepelin, 1899: 109, 112, 113, 115 (part); Masi, 1912: 124 (part); Kraepelin, 1913: 166 (part); Kopstein, 1921: 119, 133 (part); 1923: 186, 187 (part); 1926: 111, 112; 1927: 107; Giltay, 1931: 9; Werner, 1934: 277 (part); Takashima, 1945: 91, 92 (part); Vyas, 1971: 65–79, figs. 1–6; Vyas and Laliwala, 1972: 267–271, figs. 1, 2; Devarajulu Naidu and Venkatachari, 1974: 539–541, figs. 1–4; Vyas, 1974: 9–14; Dalingwater, 1980: 285–289, figs. 1c, 2, 3; Brignoli, 1985: 415; Hjelle, 1990: 17, 26, 48; Polis and Sissom, 1990: 179 (part); Sissom, 1990: 80, 81, 133, figs. 3.13G, 3.28B–E; Pointer, 1991: 24; Tare et al., 1993: 102, 103, table 1; Dupré et al., 1998: 13; Prendini et al., 2003: 192, 202, 205, 208, 209, 218, 222, 252, 254, 255, 259, table 3, figs. 4, 6, 7, appendices 1, 2 (part); Kovařík, 2004: 1, 15, 16, 29, 44, 49, 51, 52, tables 2, 3 (part), fig. 14; 2009: 35–38, 48 (part), 49, 78, 100, 104, tables 1, 2 (part), figs. 32–39, 206, 207, 246 248; Javed et al., 2010a: 144, 147; Warburg, 2010: 1746, 1750, tables 2, 3 (part); 2012a: 82, 83, 85, table 1 (part); 2013: 152, 153, 15 (part); Tahir and Prendini, 2014: 2, 9.

Palamnaeus fulvipes: Pocock, 1900a: 85, 87–89, 92, 96, figs. 25A, 29B; Bardi and George, 1943: 91–115, figs. 1, 3, 4; Sareen and Monga, 1973: 281, 282, fig. 1; Cekalovic, 1982: 191 (misidentification); Warburg, 2012a: 79, 82, table 1.

Palamnaeus fulvipes bombayensis Pocock, 1900a: 89 (synonymized by Couzijn, 1981: 133–136); Tikader, 1973: 263; Couzijn, 1981: 134–136; Kovařík, 2004: 15; 2009: 37.

Heterometrus phipsoni: Kraepelin, 1913: 166 (misidentification, part); Pande et al., 2012: 2383–2385, 2387, table 1, fig. 1, images 11a, b (misidentification).

Heterometrus sp.: Dubale and Vyas, 1968: 240–244.

Heterometrus (Chersonesometrus) fulvipes: Couzijn, 1981: 18, 23, 24, 26, 40, 88, 123, 133–136, 139, 140, 142, 149, 169, 192, table 7, figs. 1b–f, 5c, 6c, 8b, 57 (part); Tikader and Bastawade, 1983: 575, 599–603, figs. 1581–1594; Kovařík, 1997: 183; 1998: 136 (part); Fet, 2000: 439 (part); Bastawade, 2002: 296; 2004: 286, 290, 291; 2005: 411, 417, 418.

Heterometrus (Chersonesometrus) granulomanus: Couzijn, 1981: 142, 143 (misidentification, part); Bastawade, 2004: 286, 291, 292 (misidentification, part); Thulsi Rao et al., 2005: 3, 9 (misidentification, part); Bastawade, 2008: 133, 136, 137 (misidentification, part).

Heterometrus (Chersonesometrus) pelekomanus: Bastawade, 2002: 296 (misidentification).

Heterometrus (Gigantometrus) swammerdami: Thulsi Rao et al., 2005: 3, 8 (misidentification, part).

Type Material: Buthus fulvipes: Holotype ?♀ [lost], “Java.” INDIA: Maharashtra: Chandrapur Distr.: Heterometrus (Chersonesometrus) fulvipes: Neotype [= Palamnaeus fulvipes bombayensis holotype]: 1 ♀ (BMNH 1913.1.6.2 ex BNHS), Chanda [Chandrapur, 19°58′N 79°18′E], Central Provinces [examined].

Buthus fulvipes was described from a holotype female with erroneous locality data (i.e., “Java”). The holotype was subsequently lost, leading Couzijn (1981: 136) to designate the holotype of Palamnaeus fulvipes bombayensis, a junior synonym, as the neotype. Couzijn (1981: 136), Fet (2000: 439), and Kovařík (2004: 15) mistakenly referred to syntypes of P. f. bombayensis. Pocock (1900a: 89) explicitly mentioned a “type from Násik.”

Diagnosis: Chersonesometrus fulvipes may be separated from other species of Chersonesometrus as follows. The carapace is vaulted, the lateral surfaces sloping steeply (fig. 17A, B), in C. fulvipes but slightly to markedly dorsoventrally compressed, the lateral surfaces sloping gently, in all other species except C. madraspatensis and C. shivashankari. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female (fig. 17B) of C. fulvipes whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the female of C. bastawadei, C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are granular in C. fulvipes but smooth or nearly so in all other species except C. bastawadei and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. fulvipes, but subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. beccaloniae, C. hendersoni, C. nathanorum, C. pelekomanus, and C. tristis. The pedipalps of the adult male are short, with femur length: posterior carapace width ratio (FL:PCW) < 0.77 and femur length: carapace length ratio (FL:CL) < 0.74 (fig. 88A, B) in C. fulvipes but long, with FL:PCW ≥ 0.80, FL:CL ≥ 0.76 in C. beccaloniae, C. madraspatensis, and C. tristis. The pedipalp patella dorsal surface is convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina (fig. 89), in C. fulvipes but flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. fulvipes but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. fulvipes but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are granular in C. fulvipes but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. fulvipes but immaculate and paler than the femur and patella in C. shivashankari. The chela of the adult male is sparsely setose in C. fulvipes but moderately to densely setose in C. beccaloniae, C. pelekomanus, and C. tristis. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and curved, i.e., slightly to markedly convex, in the female of C. fulvipes, flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface in the male (fig. 90), or aligned with the proximal margin of the condyle (articulation with patella) in the female (fig. 91) of C. fulvipes; moderately curved and aligned with the proximal margin of the retrolateral surface in the male, or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. beccaloniae and C. nathanorum; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is greater than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. fulvipes, but greater than the DSC–DC in the male of C. beccaloniae, C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male are absent or obsolete (fig. 90) in C. fulvipes but entirely to predominantly granular or costate in all other species except C. bastawadei. The digital carina is absent or obsolete in C. fulvipes, but entirely to predominantly granular in C. beccaloniae, C. hendersoni, C. pelekomanus, C. tristis, and C. wroughtoni, and entirely to predominantly costate in C. nathanorum. The manus dorsal intercarinal surface is without reticulation in C. fulvipes but shallowly reticulate in C. nathanorum. The manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in C. fulvipes but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The legs are pale or very lightly infuscate (figs. 9A, 88) in C. fulvipes but very dark or heavily infuscate in C. beccaloniae, C. nathanorum, C. pelekomanus, C. tristis, and C. wroughtoni. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 45A–D) in C. fulvipes but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are granular in the male and the lateral surfaces granular in the female of C. fulvipes, whereas the mesial surfaces are smooth in the male of C. beccaloniae, C. hendersoni, C. pelekomanus, and C. wroughtoni, the mesial surfaces smooth in the male and the lateral surfaces smooth in the female of C. tristis, and the lateral surfaces smooth in the female of C. nathanorum. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. fulvipes and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 61A) in C. fulvipes but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis, and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are smooth in the male (fig. 63A) and granular in the female of C. fulvipes but granular in the male and smooth in the female of C. beccaloniae and C. nathanorum, granular in the male of C. hendersoni, C. madraspatensis, C. pelekomanus, and C. tristis, and smooth in the female of C. wroughtoni. The dorsolateral carinae of metasomal segment V are strong and continuous in C. fulvipes but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is smooth in the male and female of C. fulvipes, granular in the male and female of C. bastawadei, C. hendersoni, and C. pelekomanus, and granular in the male of C. wroughtoni.

Distribution: Chersonesometrus fulvipes is widely distributed across northwestern India (fig. 79) and has been recorded from the following states: Gujarat, Haryana, Karnataka, Madhya Pradesh, Maharashtra, National Capital Territory of Delhi, Rajasthan, Uttar Pradesh (table 1). A single record from Sindh, southeastern Pakistan, on the border with India (Kovařík, 2009) requires confirmation (Tahir and Prendini, 2014). The distribution of C. fulvipes is parapatric with that of its sister species, C. madraspatensis, and allopatric with the morphologically similar C. bastawadei, C. shivashankari, and C. wroughtoni.

Ecology: Chersonesometrus fulvipes inhabits deciduous forest, broadleaf woodland and semiarid savanna, dominated by Acacia, Commiphora Jacquin, 1797, Sterculia Linnaeus, 1753, and Euphorbia trees, on clayey to sandy-loam soils, at elevations of 250–940 m (fig. 7A; Pande et al., 2012). As with other species of the genus, C. fulvipes, is usually found in rocky habitats, often on wooded hill slopes, gulleys, or ravines where the humidity is slightly higher (P. Jain, personal commun.). The burrowing biology of this fossorial, pelophilous species was described by Tare et al. (1993), Bastawade (2004, 2005, 2008), and Pande et al. (2012), the latter misidentified as H. phipsoni. The burrows, constructed in open ground or under moderate to large stones, have oval or crescent-shaped entrances and are almost vertical or gradually sloping to a terminal chamber at a depth of 10–30 cm. Across its distribution, C. fulvipes occurs in sympatry with the buthids Hottentotta pachyurus (Pocock, 1897), Hottentotta tamulus (Fabricius, 1798), Lychas tricarinatus, and Orthochirus bicolor (Pocock, 1897), the hormurid Liocheles nigripes (Pocock, 1897), the scorpiopid Neoscorpiops satarensis (Pocock, 1900), and the scorpionids Deccanometrus xanthopus and Gigantometrus swammerdami (Bastawade, 2004, 2005, 2008; Pande et al., 2012).

Conservation Status: Chersonesometrus fulvipes is occasionally available in the exotic pet trade under the name Heterometrus xanthopus. Material offered for sale appears to have originated from India, regardless of the alleged country of origin.

Remarks: This species was previously accommodated in subgenus Chersonesometrus of Heterometrus by various authors (e.g., Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000). The following synonym is confirmed: Palamnaeus fulvipes bombayensis Pocock, 1900 = Chersonesometrus fulvipes (C.L. Koch, 1837), comb. nov., synonymized by Couzijn (1981). Buthus caesar, synonymized with H. (C.) fulvipes by Couzijn (1981), is revalidated as Srilankametrus caesar, comb. nov. et stat. rev.

Material Examined: 1 ♂, 1 subad. ♂ (MCZ). Gujarat: Panch Mahals Distr.: Panch Mahals [22°48′N 73°34′E], W.A. Wallinger, 2 ♂ (BMNH 1896.10.20.68-69 [BNHS 421, 425]). Sunrendranagar Distr.: Karaghora [Kharaghoda, 23°11′N 71°44′E], in Kathiawar, H. Bullkley, 1 ♂ (BMNH 1896.7.30.100 [BNHS 442]); Kathiawar, 1 ♂ (BMNH 1897.9.17.16). Haryana: Faridabad Distr.: Aravalli Hills, NNE of Mangar, 28°23′N 77°11′E, 252 m, 14.xi.2006, P. Menon, savanna in ravine on summit of rocky hill, in burrows (ca. 20 cm deep) under stones, 1 ♀, 6 juv. ♀ (AMNH), 1 juv. ♀ (AMCC [LP 13990]). Madhya Pradesh: Hoshangabad Distr.: Satpura, Pipariya Road 19, between Matkuli and Pipariya, 22°39′N 78°23′E, 364 m, 6.xii.2006, P. Menon, dry broadleaf forest on plain with gulleys, excavated from burrows in open ground, ca. 15 cm long, in hard, sandy-clay bank, 1 ♀ (AMNH), 1 juv. ♂ (AMCC [LP 14031]); Raini Pani, 22°37′N 78°07′E, 385 m, 4.xii.2006, P. Menon, deciduous forest on low hills, excavated from shallow burrows (ca. 10 cm deep) in sandy-clay under stones, 1 ♀, 1 juv. ♀ (AMNH), 1 juv. ♀ (AMCC [LP 13997]); Raini Pani road, 22°37′N 78°08′E, 369 m, 5.xii.2006, P. Menon, deciduous forest along dry gulley, excavated from burrows in open ground or under sandstones, in very hard sandy-clay, 3 ♀, 1 juv. ♀ (AMNH), 1 juv. ♀ (AMCC [LP 13981]). Satna Distr.: Satna [24°36′N 80°50′E], Sarg. Maj. Prinlette, 1 juv. ♂, 1 juv. ♀ (BMNH 1896.8.15.1-2 [BNHS 593]). Maharashtra: East Khandesh, R.H. Medan, 1 ♂ [BNHS 228], 1 juv. ♀[BNHS 224] (BMNH 1896.7.30.89); East Satpura Hills [21°25′N 76°10′E], Khandesh, J.H. Du Boulay, 1 ♂[BNHS 132], 1 ♀ [BNHS 128], 1 subad. ♂ [BNHS 138], 1 subad. ♀ [BNHS 137] (BMNH 1896.7.95-98); Khandesh, R.H. Madan, 1 ♀, 1 juv. ♂, 1 juv. ♀, 1 subad. ♂ [BNHS 126] (BMNH 1896.7.30.93-94); Talgaum [Talegaon, 20°24′N 74°55′E], East Khandesh, R.H. Madan, 1 ♂ [BNHS 234], 1 ♀ [BNHS 235] (BMNH 1896.7.30.91-92). Nashik Distr.: Kalwan [20°29′N 74°02′E], Nasik, J. Duxburg, 1 ♂ (BMNH 1897.9.5.32 [BNHS 1148]); Malegaum [Malegaon, 20°33′N 74°31′E], Nasik, J. Duxburg, 1 ♀ (BMNH 1897.9.5.28 [BNHS 1146]); Nandgaum [Nandgaon, 20°19′N 74°40′E], Nasik, J. Duxburg, 1 ♀ (BMNH 1897.9.5.29 [BNHS 1150]); Sinnar [19°51′N 74°00′E], Nasik, J. Duxburg, 1 ♂ (BMNH 1897.9.5.33 [BNHS 1152]); Yeola [20°03′N 74°29′E], Nasik, J. Duxburg, 1 ♂ (BMNH 1897.9.5.30 [BNHS 1158]); Kalwan [20°29′N 74°02′E], Nasik, J. Duxburg, 1 ♂ (BMNH 1897.9.5.32 [BNHS 1148]). Satara Distr.: Khambatki Ghat, S of Khandala, on M.B.T. Road (Highway 47), 18°01′N 74°01′E, 928 m, 17.xi.2004, S. Basi, steep slopes of rocky ridge, mixed, semiarid savanna, in scrapes under basalt stones in rocky ravine, 2 pedipalp chelae (AMNH), 1 juv. ♀ (AMCC [LP 13125]), 18°01′N 74°00′E, 937 m, 30.xi.2006, P. Menon, dry savanna woodland (mostly broadleaf) on steep mountain slope with basalt outcrops, excavated from shallow burrows (ca. 10–20 cm long) under stones in clayey-loam, 2 ♂, 1 ♀, 9 juv. ♂, 5 juv. ♀ (AMNH), 1 juv. ♂, 1 juv. ♀(AMCC [LP 13957]), 902 m, 29.xi.2009, R. Datta, broadleaf woodland on fairly steep slope of large mountain, clayey-loam with many loose/embedded volcanic (basalt) stones, excavated from shallow burrows (ca. 10–15 cm deep) under embedded stones, 2 ♂, 1 ♀(AMNH), 2 juv. ♂, 1 juv. ♀ (AMCC [LP 16812]). Rajasthan: Ajmer Distr.: Ajmeer [Ajmer, 26°27′N 74°38′E], H.M. Phipson, 1 ♂ (BMNH 1893.12.20.17); Beawar [26°06′N 74°19′E], Rajputana, R.G. Robson, 4 ♂, 1 ♀(MNHN RS 0157). Jaipur Distr.: Jeypore [Jaipur, 26°55′N 75°47′E], 1 ♀ (MCZ), Jeypore [Jaipur], Rajputana, x.1912, 2 ♂, 3 ♀ (ZMH ex Mus. Calcutta). Uttar Pradesh: Agra Distr.: Agmeza [27°11′N 77°46′E], 15.v.1899, 2 ♀ (ZMH ex Mus. Brussel). Varanasi Distr.: Benares [25°20′N 83°00′E], 1 juv. ♂, 1 juv. ♀ (MCZ). PAKISTAN: [alleged], iii.2008, ex pet trade, 1 ♂(AMNH [LP 8268]), [pedipalp chela] (AMCC [LP 8268]).

Type Material: INDIA: Karnataka: Ballari Distr.: Holotype ♀, paratypes: 1 subad. ♀, 7 juv. ♂, 6 juv. ♀ (AMNH), 2 juv. ♀ (AMCC [LP 16776]), Sanapur Lake, Sanapura, 15°22′N 76°26′E, 477–478 m, 22.xi.2009, R. Datta, dry riverbed on rocky hills, slightly disturbed mixed-broadleaf savanna on coarse granitic sandy-loam with many granite outcrops and embedded stones, in scrapes or shallow burrows under stones below canopy; paratypes: 2 ♀, 2 subad. ♂, 1 subad. ♀, 6 juv. ♂, 1 juv. ♀ (AMNH), 2 juv. ♂ (AMCC [LP 16782]), Tungabhadra River, Narayanpet, 15°20′N 76°25′E, 448 m, 22.xi.2009, R. Datta, riverine dry forest with thickets of spiny bamboo and thorn trees, on granite outcrops along permanent river, dense canopy and litter in places, from shallow scrapes in coarse granitic sandy-loam, under large, flat, embedded rocks.

Etymology: The specific epithet is a patronym honoring John Robertson Henderson (1863–1925), a Scottish zoologist who served as Superintendent of the Government Museum in Madras, British India, from 1908 to 1920, in recognition of his contributions to the scorpions and other arachnids of India.

Diagnosis: Chersonesometrus hendersoni may be separated from other species of Chersonesometrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 16F), in C. hendersoni, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female (fig. 16F) of C. hendersoni, whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the female of C. bastawadei, C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are smooth in C. hendersoni but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. hendersoni but unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, and C. wroughtoni. The pedipalp patella dorsal surface is flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane (fig. 93), in C. hendersoni but convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina in all other species except C. beccaloniae, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. hendersoni but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. hendersoni but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female (fig. 93) are granular in C. hendersoni but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. hendersoni but immaculate and paler than the femur and patella in C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and female of C. hendersoni, flat in the male and curved, i.e., slightly to markedly convex, in the female of C. bastawadei, C. beccaloniae, C. fulvipes, C. nathanorum, and C. wroughtoni, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is markedly curved and proximal to the proximal margin of the condyle (articulation with patella) in the male and female (fig. 94) of C. hendersoni but moderately curved and aligned with or proximal to the proximal margin of the retrolateral surface, or aligned with the proximal margin of the condyle in all other species except C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is greater than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. hendersoni, but greater than the DSC–DC in the male of C. beccaloniae, C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male are entirely to predominantly granular in C. hendersoni, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is entirely to predominantly granular (fig. 94) in C. hendersoni, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The manus dorsal intercarinal surface is without reticulation in C. hendersoni but shallowly reticulate in C. nathanorum. The manus ventral surface is angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. hendersoni but flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in all other species except C. beccaloniae, C. pelekomanus, and C. tristis. The legs are pale or very lightly infuscate (fig. 92) in C. hendersoni but very dark or heavily infuscate in C. beccaloniae, C. nathanorum, C. pelekomanus, C. tristis, and C. wroughtoni. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 44E–H) in C. hendersoni but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are smooth in the male and the lateral surfaces granular in the female of C. hendersoni, whereas the mesial surfaces are granular in the male of C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari, the mesial surfaces granular in the male and the lateral surfaces smooth in the female of C. nathanorum, and the lateral surfaces smooth in the female of C. tristis. The ventral surface of mesosomal sternite VII bears two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. hendersoni and a pair of weakly developed ventrolateral carinae in all other species except C. beccaloniae and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 58C) in C. hendersoni but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis, and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are granular in the male and female (fig. 60C) of C. hendersoni but smooth in the male and female of C. wroughtoni, the male of C. bastawadei, C. fulvipes, and C. shivashankari, and the female of C. beccaloniae and C. nathanorum. The dorsolateral carinae of metasomal segment V are strong and continuous in C. hendersoni but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is granular in the male and female of C. hendersoni, smooth in the female of C. wroughtoni, and smooth in the male and female of all other species except C. bastawadei and C. pelekomanus.

Description: The following description is based on the holotype and paratypes. Meristic data are presented in table 3. The adult male is unknown hence measurements are presented only for the adult female (table 4).

Total length: Adult large, maximum length, measured from anterior margin of carapace to tip of aculeus, 137 mm (126–156 mm, n = 3) (♀).

Color: Chelicerae, dorsal surfaces bicolored, mostly to entirely immaculate prodorsal half of manus paler than infuscate retrodorsal half of manus and fingers; manus markedly paler than carapace interocular surface. Carapace, pedipalps, tergites, and metasoma entirely infuscate, uniformly dark reddish brown to reddish black. Sternites III–VI immaculate, VII immaculate to slightly infuscate, dark olive to blackish brown, becoming darker posteriorly. Coxosternal region infuscate, brownish, maxillary lobes darker. Legs lightly infuscate, uniformly yellow to light brown, contrasting with carapace, pedipalps, tergites, and metasoma (fig. 92). Genital opercula and pectines uniformly pale cream. Metasomal segments I–V similar to one another; carinae infuscate, darker than intercarinal surfaces; dorsal intercarinal surfaces similar to tergites; ventral intercarinal surfaces darker than sternites. Telson vesicle immaculate, light yellowish, markedly paler than metasomal segment V; aculeus black.

Chelicerae: Movable finger prodistal (DI) and retrodistal (DE) teeth subequal, with DE tooth only slightly smaller than DI tooth, and opposable, i.e., forming a bicusp. Fingers and manus, proventral surfaces with long, dense vestiture of macrosetae. Coxae, prodorsal surfaces without stridulatory setae (scaphotrix); promedian surfaces without chemoreceptive lamelliform setae (trichocopae).

Carapace: Carapace markedly dorsoventrally compressed, lateral surfaces sloping gently. Anterior margin concave, frontal lobes uneven, sloping medially toward deep median notch, without median projection or median depression (fig. 16F). Anterolateral margins converging gradually anteriorly, without distinct notch adjacent to posterior lateral ocelli; mediolateral margins diverging or converging slightly posteriorly (at posterolateral sulci). Anterior width of posterior width, 65% (64%–65%, n = 3) (♀); posterior width of length, 97% (95%–98%, n = 3) (♀). Three pairs of lateral ocelli (MLMa, PLMa, PDMi), all similar in size, two anterior pairs situated anteriorly, well separated from posterior pair, situated laterally. Median ocelli slightly larger than lateral ocelli, distance between ocelli slightly less than width of ocellus; median ocular tubercle situated posteromedially, distance from anterior carapace margin, 52% (51%–52%, n = 3) (♀). Median longitudinal sulcus narrow, suturiform; continuous from median notch to interocular sulcus; anteriorly furcated; without anterocular depression. Anterior furcated sulci diverging broadly from anterior edge of median longitudinal sulcus. Median ocular tubercle raised, superciliary carinae higher than ocelli, extended anteriorly, subparallel. Interocular sulcus present. Circumocular depressions completely encircling median ocular tubercle, converging anteriorly. Posteromedian and posteromarginal sulci deep. Paired mediolateral and posterolateral sulci shallow. Median longitudinal suture continuous from median notch to median ocular tubercle, equally strong along entire length; not extending to anterior margin of carapace, terminating at or posterior to median notch. Anterior furcated sutures absent or obsolete. Anterocular furcated sutures absent. Interocular suture present, slender. Posterior sutures present but indistinct, converging on median ocular tubercle; connected anteriorly to posterior furcations of interocular suture and extending anteriorly beyond median ocular tubercle. Frontal lobes and medial region of interocular surface finely granular with smooth areas (♂) or finely granular along median longitudinal and anterior furcated sulci only, elsewhere smooth (♀) (fig. 16F); anterolateral and mediolateral surfaces uniformly finely granular; posterolateral surfaces uniformly finely granular (♂) or smooth (♀); posteromedian surfaces smooth.

Pedipalps: Femur width of length, 47% (45%–51%, n = 3) (♀). Retrodorsal carina complete, costate-granular, similar to or more strongly developed than prodorsal carina. Dorsal secondary carina partial, reduced to granular row in anterior quarter, becoming obsolete distally. Dorsomedian carina partial, reduced to granular row in anterior quarter. Prodorsal carina complete, granular. Promedian carina complete, comprising row of spiniform or subspiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Proventral carina complete, granular as strongly developed as or more strongly developed than retroventral carina. Ventromedian and secondary accessory carinae absent. Retroventral carina extending partly or entirely to distal edge of segment. Retromedian carinae absent, indicated only by macrosetal rows. Dorsal and prolateral intercarinal surfaces finely granular; ventral and retrolateral surfaces smooth. Patella width of length, 51% (50%–53%, n = 3) (♀). Dorsal surface flat, axes of dorsomedian and retrodorsal carinae in same plane. Dorsomedian carina entirely to predominantly costate-granular (fig. 93). Retrodorsal carina predominantly costate-granular, more strongly developed than granular retromedian carinae. Retroventral carina distinct, costate-granular. Promedian carina and promedian process absent. Other carinae absent. Dorsal and retrolateral intercarinal surfaces finely granular, other surfaces smooth. Chela short, broad (fig. 94); manus height of width, 65% (60%–67%, n = 3) (♀); manus length along retroventral carina of manus width, 86% (80%–92%, n = 3) (♀); manus length along retroventral carina of movable finger length, 74% (70%–79%, n = 3) (♀). Chela moderately (♂) to sparsely (♀) setose. Manus dorsal surface (between dorsomedian and digital carinae) flat; proximal margin (lobe) of dorsal surface markedly curved and proximal to proximal margin of condyle (articulation with patella); proximal margin (lobe) of dorsal surface moderately curved and aligned with (♂) or proximal to (♀) proximal margin of retrolateral surface. Dorsomedian carina obsolete, costate on fixed finger and comprising regular row of spiniform granules along entire length of manus. Dorsal secondary, subdigital, and digital carinae incomplete, entirely to predominantly granular; dorsal secondary carina comprising double row of granules in proximal two-thirds; proximal half of dorsal secondary carina, distal half of subdigital carina and distal third of digital carina closely adjacent. Maximum distance between dorsomedian and dorsal secondary carinae (DMC–DSC) greater than maximum distance between dorsal secondary and digital carinae (DSC–DC). Retromedian carina complete, predominantly costate, more strongly developed than digital carina. Retrolateral secondary and secondary accessory carinae vestigial, proximal to condyle of movable finger. Retroventral carina distinct, costate. Ventromedian carina obsolete, costate. Proventral and promedian carinae similarly developed, each comprising row of spiniform granules and associated macrosetae. Prodorsal carina obsolete, comprising few spiniform granules and associated macrosetae. Distance between dorsomedian and promedian carinae (DMC–PMC) slightly to markedly greater than distance between promedian and proventral carinae (PMC–PVC). Manus, dorsal and retrolateral intercarinal surfaces densely and coarsely granular except for smooth depression at base of fixed finger; ventral intercarinal surface smooth or nearly so; prolateral intercarinal surfaces very sparsely and finely granular, more densely dorsally and distally. Manus ventral surface angled, axis of retroventral carina ventral to axis of ventromedian carina. Fixed and movable fingers, intercarinal surfaces smooth; median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three situated on lobes; first (proximal) lobe of fixed finger and second lobe of movable finger larger than others, with correspondingly deeper notches in movable and fixed fingers, respectively; terminal denticles of fingers interlocking unevenly when closed, movable finger displaced retrolaterally; distinct notch near tip of fixed finger to accommodate terminal denticle of movable finger.

Trichobothria: Orthobothriotaxic, Type C, with the following segment totals (n = 12): femur, 3 (1 d, 1 i, 1 e); patella, 19 (2 d, 1 i, 3 v, 13 e, comprising 3 et, 1 est, 2 em, 2 esb, 5 eb); chela, 26 (manus, 16, comprising 2 D, 10 E, 4 V; fixed finger, 10, comprising 4 d, 4 e, 2 i). Total count of trichobothria per pedipalp: 48. Femur, i situated on dorsal surface. Patella, d₂ situated on internal surface, slightly closer to d₁ than to i; distance v₂–v₃ ca. one-third distance v₁–v₂. Chela, distance et–est ca. half distance est–esb; distance est–esb greater than half distance esb–eb; est distal to dst; V₃ situated in proximal third of manus, closer to V₄ than V₂.

Legs: First pair of maxillary lobes (coxapophyses) tapering anteriorly, longer than and encircling second pair. Stridulatory organs, comprising “rasp” (granular tubercles) and “scraper” (stridulatory setae or scaphotrix), present on opposing surfaces of coxae of first pair of legs and pedipalps, respectively. Legs I–IV acarinate, except femora, with distinct pro- and retrolateral surfaces demarcated by partial, granular prodorsal and complete, costate-granular proventral carinae. Femora, patellae, and tibiae, pro- and retrolateral surfaces each with scattered setiform macrosetae. Tibiae, prolateral surfaces, without spiniform macrosetae; I and II, retrolateral surfaces, each with two spiniform (t, st) macrosetae; III and IV, retrolateral surfaces, without spiniform macrosetae. Basitarsi I–IV, slightly dorsoventrally compressed; retrolateral margins similar, unmodified, rounded; prolateral pedal spurs present (fig. 44E–H); retrolateral pedal spurs absent. Basitarsi, pro- and retrolateral surfaces, each with scattered long and short setiform macrosetae, and spiniform macrosetae, more numerous on I and II than III and IV. Basitarsi, spiniform macrosetae, I, retrolateral: t, sb; retroventral: t, st, sb; proventral: t, st; II, retrolateral: t, sb; retroventral: t, st, sb; proventral: t, st; III, retrolateral: t, sb; retroventral: t, st; proventral: t; prolateral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: t, st. Telotarsi I–IV short, stout and distally broadened in dorsal and lateral views. Laterodistal lobes rounded. Dorsomedian lobes approximately equal to laterodistal lobes; each terminating in single setiform macroseta. Telotarsi, pro- and retrolateral surfaces, each with long and short, scattered macrosetae, not arranged in definite row. Telotarsi each with pro- and retroventral rows of spiniform macrosetae, two inserted on laterodistal lobes; counts in pro- and retroventral rows similar on I and II, and on III and IV (n = 16): I, proventral: 4, retroventral: 6; II, 4 (4 or 5), 6 (6 or 7); III and IV, 5, 6 (6 or 7). Telotarsal ungues short, curved, equal to subequal; pseudonychium (dactyl) reduced, rounded.

Sternum: Shape subpentagonal (fig. 30F). Median longitudinal sulcus shallow anteriorly, deep and narrow posteriorly.

Genital operculum: Genital opercula suboval, completely divided longitudinally, partially overlapping, genital papillae present (♂); subcordate, fused, genital papillae absent (♀) (fig. 30F).

Hemispermatophore: Unknown.

Pectines: Distal edge extending past distal edge of coxa IV but not reaching distal edge of trochanter IV (♂) or reaching to distal edge of coxa IV (♀) (fig. 30F). First proximal median lamella (scape) of each pecten with mesial margin angular, approximately 90° (♂) or obtusely angular, greater than 90° but less than 180° (♀). Pectinal teeth straight and elongate (♂) or shorter and curved (♀), present along entire posterior margin; sinistral/dextral tooth count, 19/19 (17–19/18 or 19, n = 6) (♂), 16/16 (16 or 17/16 or 17, n = 10) (♀). Fulcra smooth proximally but densely setose (microsetae only) distally.

Mesosoma: Tergites each with shallow pair of submedian depressions, without obsolete median carina. Pretergites smooth and glabrous. Posttergites smooth and glabrous medially, unevenly finely and coarsely granular to predominantly coarsely granular laterally. Sternites IV–VI, each with paired longitudinal depressions prolateral to spiracles, absent on VII. Surface, sternites III–VII, smooth; VII with two pairs of moderately to strongly developed, costate ventrosubmedian and ventrolateral carinae. Sternite VII, length of width, 66% (64%–70%, n = 3) (♀).

Metasoma and telson: Metasomal segments I–V progressively increasing in length, decreasing in width; segment V, width of segment I, width, 69% (67%–71%, n = 3) (♀). Metasoma fairly robust, width of length, segment I, 83% (79%–86%, n = 3) (♀); II, 67% (63%–70%) (♀); III, 61% (56%–64%) (♀); IV, 49% (48%–51%) (♀); V, 35% (34%–37%) (♀). Telson vesicle, width of metasomal segment V, width, 102% (100%–105%, n = 3) (♀); vesicle globose, height of length, 52% (51%–53%, n = 3) (♀); dorsal surface flat; ventral surface evenly curved. Aculeus relatively long, gently curved, length of vesicle length, 67% (63%–72%, n = 3) (♀). Length metasoma and telson, of total length, 54% (53%–54%, n = 3) (♀). Dorsosubmedian carinae, segments I–IV, distinct, complete, granular or costate-granular, posterior spiniform granules obsolete (fig. 58C); V, vestigial. Dorsolateral carinae, segments I–V, distinct, complete, costate-granular on I–IV, costate-granular to granular on V. Median lateral carinae, segment I, incomplete, reduced to granular row in posterior half, diagonally oriented between dorsolateral and ventrolateral carinae (fig. 59C); II–IV, obsolete row of granules posteriorly; V, incomplete, granular row, restricted to anterior two-thirds. Ventrolateral and ventrosubmedian carinae, segments I–IV, distinct, complete, costate on I–III, granular or costate-granular on IV, more strongly developed on III and IV than on I and II (fig. 60C). Ventrolateral carinae, segment V, distinct, complete, comprising spiniform granules, diverging posteriorly, terminal granule slightly larger than preceding granules. Ventrosubmedian carinae, segment V, vestigial, each reduced to one or two isolated spiniform granules, demarcated by conspicuous macrosetae. Ventromedian carina, segment V comprising single row of spiniform granules, unmodified posteriorly. Anal arch, segment V, dorsal carina, costate; ventral carina comprising subspiniform granules. Lateral intercarinal surfaces, segments I–IV, granular, V, smooth. Ventral intercarinal surfaces, segments I–III, smooth, IV and V, granular. Telson vesicle, dorsal and lateral surfaces smooth; ventral surface with four longitudinal carinae, each comprising spiniform granules.

Distribution: This species is endemic to India and presently recorded only from the state of Karnataka (fig. 80, table 1).

Ecology: The known locality records of this species occur in semiarid savanna and deciduous riverine forest dominated by Acacia, Euphorbia, and broadleaf bushes (Grewia, Ziziphus), with bamboo, along riverbeds in rocky granite hills, at elevations of 450–480 m (fig. 7D). The specimens for which data are available were collected from shallow scrapes in coarse granitic sandy-loam soil under large, flat, embedded rocks. The habitat and habitus are consistent with the pelophilous, fossorial and semilithophilous ecomorphotypes (Prendini, 2001b). This species is sympatric with Charmus indicus, Hottentotta rugiscutis, Lychas scaber (Pocock, 1893), L. tricarinatus, and an undetermined species of Reddyanus Vachon, 1972.

Scorpio (Buthus) fulvipes: Laurie, 1890: 105.

Scorpio fulvipes: Laurie, 1891: 587–597, pl. XL, figs. 1–18; Pocock, 1893: 304–307, 309 (part); Kraepelin, 1894: 31, 43–46, 52, 59–61, 246, pl. I, figs. 22, 28, 37 (part); Laurie, 1896a: 187, 188, 192, 194, pl. IX, figs. 3, 6; Warburg, 2010: 1749; 2012b: 60.

Heterometrus fulvipes: Kraepelin, 1899: 109, 112, 113, 115 (part); Masi, 1912: 124 (part); Kraepelin, 1913: 166 (part); Giltay, 1931: 9; Sasirababu, 1961: 11–17, pl. I, figs. 1, 2 (misidentification); Pampapathi Rao, 1964: 133–141, fig. 1 (misidentification); Laverack, 1966: 241, 244, 250 (misidentification); Pampapathi Rao and Murthy, 1966: 520, 521 (misidentification); Sasirababu and Venkatachari, 1966: 22–32, figs. 1–4 (misidentification); Venkateswara Rao and Sepur Govindappa, 1967: 243–249, tables I, II, pl. XIII, figs. 1–3 (misidentification); Venkatachari and Muralikrishna Dass, 1968: 617–620, graph (misidentification); Devarajulu Naidu, 1969: 1274, figs. 1, 2 (misidentification); Chengal Raju et al., 1973: 964, 965, fig. (misidentification); Geethabali and Pampathi Rao, 1973: 189–196, figs. 2, 3, 4C, 4D, 5, 6 (misidentification); Devarajulu Naidu and Padmanabhanaidu, 1976: 1–5, tables 1, 2, figs. 1–4 (misidentification); Geethabali, 1976: 345–347, table, figs. 1–3 (misidentification); Chandrasekara Reddy and Padmanabhanaidu, 1977: 217, 218, fig. 1 (misidentification); Mastanaiah et al., 1977: 1051, 1052, fig. (misidentification); Chengal Raju et al., 1978: 111–113, table I (misidentification); Jayaram et al., 1978: 1184, 1185, table, fig. (misidentification); Mastanaiah et al., 1978: 130, 131, fig. 1 (misidentification); Subburam and Gopalakrishna Reddy 1978a: 513 514 table I (misidentification); 1978b: 665, 666, figs. 1–4 (misidentification); Jayaram and Padmanabhanaidu, 1979: 6–8, tables I, II (misidentification); Mastanaiah et al., 1979: 69, 70, fig. (misidentification); Subburam and Gopalakrishna Reddy, 1979: 853–855, table I (misidentification); Uthaman and Srinivasa Reddy, 1979: 347–352, tables 1, 2, fig. 1 (misidentification); Jayaram and Padmanabhanaidu, 1980: 109–112, table 1 (misidentification); Subburam and Gopalakrishna Reddy, 1980a: 658–661, tables I, II (misidentification); 1980b: 138, 139, figs. 1–4 (misidentification); Yellamma et al., 1980: 29–37, figs. 1–7 (misidentification); Subburam and Gopalakrishna Reddy, 1981: 275–281, figs. 1–31 (misidentification); Yellamma et al., 1982: 225–233, tables 1–3, fig. 1 (misidentification); Subburam and Gopalakrishna Reddy, 1989: 329–333, figs. 1–3 (misidentification); Root, 1990: 354, 365, 368, 396, 404–410, figs. 9.24, 9.29 (misidentification); Polis and Sissom, 1990: 179 (part); Simard and Watt, 1990: 419 (misidentification); Warburg and Polis, 1990: 229, 235 table 5.2 (misidentification); Pointer, 1991: 24; Rajyalakshmi et al., 1991: 429–432, fig. 1 (misidentification); Shivashankar and Veeresh, 1991: 697–702 (misidentification); Kalarani et al., 1992: 141–146, tables 1–3, fig. 1 (misidentification); Shivashankar, 1992: 134–138 (misidentification); 1994: 81–90, tables 1–3, figs. 1–3 (misidentification); Prendini et al., 2003: 192, 193, 202, 203, 205, 208, 209, 218, 222, 252, 254, 255, 259, tables 3, 4, figs. 4–7, appendices 1, 2 (misidentification, part); Kovařík, 2004: 1, 15, 29, 49, 51, 52, tables 2, 3 (misidentification, part); Mirza and Sanap, 2009: 488, 490 (misidentification); Javed et al., 2010b: 785, table 1 (misidentification); Warburg, 2010: 1739, 1749, 1750, table 3 (misidentification, part); 2012a: 82, 83, 85, table 1 (part); 2012b: 60, 61, 63, 64 (misidentification); 2013: 152, 153, 155 (misidentification, part).

Palamnaeus fulvipes madraspatensis Pocock, 1900a: 88, 89 (part), fig. 25B; Tikader, 1973: 263; Couzijn, 1981: 135; Fet, 2000: 439, 440; Kovařík, 2004: 29.

Heterometrus (Scorpio) fulvipes: Kraepelin, 1901: 271 (misidentification, part).

Pandinus fulvipes: Simon, 1905: 161 (misidentification).

Heterometrus (Chersonesometrus) fulvipes: Couzijn, 1981: 23, 40, 88, 123, 132–136, 139, 140, 142, 149, 169, 192, table 7, figs. 39, 57 (misidentification, part); Kovařík, 1998: 136 (part); 2002: 17 (misidentification).

Heterometrus (Chersonesometrus) madraspatensis: Tikader and Bastawade, 1983: 576, 630–635, figs. 1668–1682; Fet, 2000: 440; Indra, 2009: 142.

Heterometrus madraspatensis: Kovařík, 2004: 1, 2, 27–29, 51, 52, tables 2, 3 (part), fig. 20; 2009: 35, 36, 41, 48, 49, 85, 100, 108, tables 1, 2 (part), figs. 83–88, 208, 209, 257, 258; Javed et al., 2010b: 785, table 1; Tahir and Prendini, 2014: 9.

Heterometrus (Chersonesometrus) granulomanus: Thulsi Rao et al., 2005: 3, 9 (misidentification, part).

Type Material: INDIA: Andhra Pradesh: Kadapa Distr.: Palamnaeus fulvipes madraspatensis: Paralectotype [sex?] (BMNH), Cuddapah [Kadapa, 14°28′N 78°49′E], Popert. Tamil Nadu: Chennai Distr.: P. f. madraspatensis: Lectotype ♂, paralectotype ♂ (BMNH), Madras [Chennai, 13°05′N 80°17′E], Henderson [examined]. Telangana: Hyderabad Distr.: P. f. madraspatensis: Paralectotype [sex?] (BMNH), Secunderabad [17°26′N 78°30′E], Ricardo.

In the description of Palamnaeus fulvipes madraspatensis, Pocock (1900a: 88, 89) referred to specimens from three localities, Madras, Cuddapah, and Secunderabad, but did not specify a “type,” hence all specimens listed are syntypes. Couzijn (1981: 133–136) mistakenly referred to a holotype of P. f. madraspatensis, Tikader and Bastawade (1983: 630–635), referred to two male type specimens and mistakenly listed Madras as the type locality, and Fet (2000: 440) followed these authors in incorrectly referring to a male holotype from Madras and paratypes from Madras, Cudappah, and Secunderbad. Kovařík (2004) correctly designated a male lectotype and male paralectotype from Madras but failed to mention the specimens from Cuddapah and Secunderabad, which are therefore also paralectotypes.

Diagnosis: Chersonesometrus madraspatensis may be separated from other species of Chersonesometrus as follows. The carapace is vaulted, the lateral surfaces sloping steeply (fig. 17C, D), in C. madraspatensis, but slightly to markedly dorsoventrally compressed, the lateral surfaces sloping gently in all other species except C. fulvipes and C. shivashankari. The frontal lobes and medial region of the carapace interocular surface are granular with smooth areas in the female (fig. 17D) of C. madraspatensis whereas the interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female of all other species except C. bastawadei, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are granular in C. madraspatensis but smooth or nearly so in all other species except C. bastawadei and C. fulvipes. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. madraspatensis, but subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. beccaloniae, C. hendersoni, C. nathanorum, C. pelekomanus, and C. tristis. The pedipalps of the adult male are long, with femur length: posterior carapace width ratio (FL:PCW) ≥ 0.80 and femur length: carapace length ratio (FL:CL) ≥ 0.76 (fig. 95A, B) in C. madraspatensis but short, with FL:PCW < 0.77, FL:CL < 0.74 in all other species except C. beccaloniae and C. tristis. The pedipalp patella dorsal surface is convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina (fig. 96) in C. madraspatensis but flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. madraspatensis but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. madraspatensis but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are granular in C. madraspatensis but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. madraspatensis but immaculate and paler than the femur and patella in C. shivashankari. The chela of the adult male is sparsely setose in C. madraspatensis but moderately to densely setose in C. beccaloniae, C. pelekomanus, and C. tristis. The chela manus dorsal surface (between the dorsomedian and digital carinae) is curved, i.e., slightly to markedly convex, in the male and female of C. madraspatensis, flat in the male and curved in the female of C. bastawadei, C. beccaloniae, C. fulvipes, C. nathanorum, and C. wroughtoni, and flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface in the male (fig. 97), or aligned with the proximal margin of the condyle (articulation with patella) in the female (fig. 98) of C. madraspatensis; moderately curved and aligned with the proximal margin of the retrolateral surface in the male, or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. beccaloniae and C. nathanorum; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is similar to the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. madraspatensis, but greater than the DSC–DC in the male of all other species except C. beccaloniae, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male are entirely to predominantly granular (fig. 97) in C. madraspatensis, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is absent or obsolete in C. madraspatensis, but entirely to predominantly granular in C. beccaloniae, C. hendersoni, C. pelekomanus, C. tristis, and C. wroughtoni, and entirely to predominantly costate in C. nathanorum. The manus dorsal intercarinal surface is without reticulation in C. madraspatensis but shallowly reticulate in C. nathanorum. The manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in C. madraspatensis but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The legs are pale or very lightly infuscate (fig. 95) in C. madraspatensis but very dark or heavily infuscate in C. beccaloniae, C. nathanorum, C. pelekomanus, C. tristis, and C. wroughtoni. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 45I–L) in C. madraspatensis but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are granular in the male and the lateral surfaces granular in the female of C. madraspatensis, whereas the mesial surfaces are smooth in the male of C. beccaloniae, C. hendersoni, C. pelekomanus, and C. wroughtoni, the mesial surfaces smooth in the male and the lateral surfaces smooth in the female of C. tristis, and the lateral surfaces smooth in the female of C. nathanorum. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. madraspatensis and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 61C) in C. madraspatensis but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis, and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are granular in the male and female (fig. 63C) of C. madraspatensis but smooth in the male and female of C. wroughtoni, the male of C. bastawadei, C. fulvipes, and C. shivashankari, and the female of C. beccaloniae and C. nathanorum. The dorsolateral carinae of metasomal segment V are strong and continuous in C. madraspatensis but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is smooth in the male and female of C. madraspatensis, granular in the male and female of C. bastawadei, C. hendersoni, and C. pelekomanus, and granular in the male of C. wroughtoni.

Distribution: Chersonesometrus madraspatensis is widely distributed across eastern India (fig. 79) and has been recorded from the following states: Andhra Pradesh, Chhattisgarh, Karnataka, Odisha, Tamil Nadu, Telangana, and the union territory of Puducherry (table 1). Its distribution is parapatric with that of its sister species, C. fulvipes, and allopatric with the morphologically similar C. bastawadei, C. shivashankari, and C. wroughtoni.

Ecology: Chersonesometrus madraspatensis inhabits deciduous forest and broadleaf woodland, on sandy- to gravelly-loam soils, at elevations of 370 m. As with other species of the genus, C. madraspatensis is usually found in rocky habitats, often on wooded hill slopes, gulleys, or ravines where the humidity is slightly higher. The burrowing biology of this fossorial, pelophilous species was described by Shivashankar and Veeresh (1991) and Shivashankar (1992, 1994), misidentified as H. fulvipes. The burrows, constructed in open ground or under moderate to large stones (fig. 8E), have oval or crescent-shaped entrances and are almost vertical or gradually sloping to a terminal chamber at a depth of 10–20 cm. Across its distribution, C. madraspatensis occurs in sympatry with the buthids Hottentotta rugiscutis and Lychas tricarinatus and the scorpionid Gigantometrus swammerdami.

Conservation Status: Chersonesometrus madraspatensis is occasionally available in the exotic pet trade under the name Heterometrus xanthopus.

Remarks: Pocock (1900a: 88, 89) based two subspecies of Palamnaeus fulvipes, i.e., P. f. bombayensis and P. f. madraspatensis, on a difference in the shape of the male pedipalps:

Couzijn (1981) synonymized both subspecies with the nominotypical form of Heterometrus (Chersonesometrus) fulvipes whereas Tikader and Bastawade (1983) elevated P. f. madraspatensis to the rank of species as Heterometrus (Chersonesometrus) madraspatensis, a decision upheld by Fet (2000) and Kovařík (2004). In discussing the question, Prendini et al. (2003: 218) remarked:

Previous opinions that H. fulvipes and H. madraspatensis are conspecific (Couzijn, 1981; Prendini et al., 2003) were based on their very similar morphology. Indeed, Kovařík (2004: 15, 27) referred specimens (including adult males) from Podili (CAS) to both species. The counts of pro- and retroventral spiniform macrosetae on the leg telotarsi provided for H. fulvipes and H. madraspatensis by Tikader and Bastawade (1983) and Kovařík (2004) are identical. Tikader and Bastawade (1983) cite counts of 3/5 on legs I and II, 4/5 on III, and 4/6 on IV for H. fulvipes and counts of 3–4/5 on I and 4/6 on II–IV for H. madraspatensis whereas Kovařík (2004) cites counts of 4/5–6 on III and IV for both taxa. The primary character justifying the distinction between H. fulvipes and H. madraspatensis is the shape (width) of the male chela manus first proposed by Pocock (1900a). As described in the key of Kovařík (2004: 52) for H. fulvipes “male and female chela differently shaped but of about the same width, its length to width ratio 1.7–1.9 in both sexes,” and for H. madraspatensis “chela narrower in male than in female, its length to width ratio 2.0–2.5 in males and 1.7–1.9 in females.” Following an examination of material from across the distribution of both taxa during the present investigation, this difference, albeit subtle, appears to consistently separate H. fulvipes in the northwest of India from H. madraspatensis in the southeast (fig. 79). Additional diagnostic characters identified during the present investigation, e.g., the greater length of the male pedipalp patella and femur, the more convex female pedipalp chela manus, and the more convex carapace of H. madraspatensis, taken together with genetic divergence supporting reciprocal monophyly of samples assigned to the two taxa based on morphology, justifies the continued recognition of H. madraspatensis as distinct from H. fulvipes, hence: Chersonesometrus madraspatensis (Pocock, 1900), comb. nov.

Material Examined: INDIA: 4 ♂, 6 ♀, 2 subad. ♂, 1 subad. ♀ (AMNH); vi.1999, ex F. Somma, 1 ♀(AMCC 101695 [LP 1533]). Andhra Pradesh: Chittoor Distr.: Kothakota, on Chittoor–Tirupati Road 61, 13°23′N 79°06′E, 372 m, 30.xii.2004, S. Basi, ravine between rocky ridges, broadleaf thicket, in shallow burrow (ca. 10 cm deep) under large stone, 1 subad. ♂(AMCC [LP 13127]); Madireddy Palem [Madireddypalem, 15°37′N 79°35′E], 1.i.1967, D.E. Johnson, ex Stahnke coll., 2 ♂, 3 ♀, 2 subad. ♂ (CAS [ASU 67-053, 54, 58–60, 62, 63]). Prakasam Distr.: Podili [15°36′N 79°37′E], 1.i.1967, Johnson and Guruiah, ex Stahnke coll., 2 ♀, 1 juv. ♀ (CAS 9071141 [ASU 67-040, 56]), 13.ii.1967, D.E. Johnson, ex Stahnke coll., 2 ♂ (CAS 9071142 [ASU 67-352, 363]), 11 ♀ (CAS 9071142 [ASU 67-358, 366, 368], 9071143 [ASU 67-359–361, 364, 369–371, 373]), 1 subad. ♀ (CAS 9071143 [ASU 67-365]). Karnataka: Bengaluru Distr.: Mysore state, env. Bangalore [Bengaluru, 12°59′N 77°35′E], 8.v.1974, Geethamali, 5 ♀ (MNHN RS 7798–7802), 1 juv. ♂(MNHN RS 7803). Odisha: Sambalpur Distr.: Badrama [21°29′N 84°17′E], Orissa, 1100 ft., 20–26.ii.1975, M.L. Ripley, 1 juv. ♀ (USNM). Tamil Nadu: Chennai Distr.: Madras [Chennai, 13°05′N 80°17′E], viii.1999, T. Kolb, 1 ♀ (ZMH ex SMF); Madras, J.R. Henderson, 2 ♂, 3 ♀(BMNH 1892.2.15.1-3); Madras and Callula, J.R. Henderson, 2 ♂, 1 subad. ♂, 10 juv. ♂ (BMNH 1894.10.24.5-8). Kancheepuram Distr.: Nemmali [Nemmeli, 12°43′N 80°13′E], 45 km from Chennai, on eastern coastline, 30.v.2005, H.A. Vadivelu, A. and R. Ali, and F. Paul, 4 ♂, 4 ♀, 1 juv. ♂, 1 juv. ♀ (AMCC [LP 4324]). Villu-puram Distr.: Genji [Gingi/Gingee, 12°15′N 79°24′E], Coromandel, 25.viii–12.ix.1901, M. Maindron, 4 ♂, 6 ♀, 2 subad. ♀, 1 juv. ♀ (MNHN RS 0113), M. Maindron, 2 ♂, 3 ♀ (MNHN RS 0085).

Type Material: INDIA: Tamil Nadu: Theni Distr.: Holotype: 1 subad. ♂, paratype: 1 juv. ♀(MNHN RS 3145), India, Monts Palni, 8 km NE Ganguvarpatti [Genguvarpatti] village [10°12′N 77°44′E], 20.ii.1943, Dr. K. Lindberg, se pente en montant [upper slope].

Etymology: The specific epithet is a patronym honoring Prabala Susai Nathan (1891–1976), widely known as P. Susai Nathan, and his daughter-in-law, Theresa Rajabi Susai Nathan, widely known as T.R.S. Nathan or T.R.S.N., Indian entomologists and natural history dealers from Kurumbagaram, Puducherry, who provided many museums with important arthropod collections from India between 1945 and 1975 (Hincks, 1960; Brindle, 1965; Miles, 2015), including much of the heterometrine material in the Herbert L. Stahnke Collection, presently deposited at the CAS. Handwritten labels bearing the unpublished name “Heterometrus nathani” accompany the type material of Deccanometrus ubicki (Kovařík, 2004), comb. nov. at the CAS.

Diagnosis: Chersonesometrus nathanorum may be separated from other species of Chersonesometrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 16E) in C. nathanorum, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female of C. nathanorum, whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the female of C. bastawadei, C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are smooth in C. nathanorum but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. nathanorum but unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, and C. wroughtoni. The pedipalp patella dorsal surface is convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina (fig. 99) in C. nathanorum but flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is absent or obsolete in C. nathanorum but as strongly developed as or more strongly developed than the retromedian carinae in all other species. The retromedian carinae are costate in C. nathanorum but granular in all other species. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are smooth or nearly so in C. nathanorum but granular in all other species. The pedipalp chela is infuscate and similar in color to the femur and patella in C. nathanorum but immaculate and paler than the femur and patella in C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and curved, i.e., slightly to markedly convex, in the female of C. nathanorum, flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and aligned with the proximal margin of the retrolateral surface in the male (fig. 100), or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. nathanorum; moderately curved and proximal to the proximal margin of the retrolateral surface in the male, or aligned with the proximal margin of the condyle (articulation with patella) in the female of C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, C. tristis, and C. wroughtoni; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is similar to the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. nathanorum, but greater than the DSC–DC in the male of all other species except C. beccaloniae, C. madraspatensis, and C. tristis. The dorsal secondary and subdigital carinae of the male (fig. 100) are entirely to predominantly costate in C. nathanorum, absent or obsolete in C. bastawadei and C. fulvipes, and entirely to predominantly granular in all other species of Chersonesometrus. The digital carina is entirely to predominantly costate in C. nathanorum, entirely to predominantly granular in C. beccaloniae, C. hendersoni, C. pelekomanus, C. tristis, and C. wroughtoni, and absent or obsolete in C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The manus dorsal intercarinal surface is shallowly reticulate in C. nathanorum but without reticulation in all other species. The manus ventral surface is flat, with the axis of the retroventral and ventromedian carinae in approximately the same plane, in C. nathanorum but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The legs are very dark or heavily infuscate in C. nathanorum but pale or very lightly infuscate in C. bastawadei, C. fulvipes, C. hendersoni, C. madraspatensis, and C. shivashankari. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is setiform in C. nathanorum but spiniform in all other species except C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are granular in the male and the lateral surfaces smooth in the female of C. nathanorum, whereas the mesial surfaces are smooth in the male of C. tristis, the mesial surfaces smooth in the male and the lateral surfaces granular in the female of C. beccaloniae, C. hendersoni, C. pelekomanus, and C. wroughtoni, and the lateral surfaces granular in the female of C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. nathanorum and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 61D) in C. nathanorum but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are granular in the male (fig. 63D) and smooth in the female of C. nathanorum but smooth in the male and granular in the female of C. bastawadei, C. fulvipes, and C. shivashankari, smooth in the male of C. wroughtoni, and granular in the female of C. hendersoni, C. madraspatensis, C. pelekomanus, and C. tristis. The dorsolateral carinae of metasomal segment V are weak and discontinuous to absent in C. nathanorum but strong and continuous in all other species. The dorsal intercarinal surface of segment V is smooth in the male and female of C. nathanorum, granular in the male and female of C. bastawadei, C. hendersoni, and C. pelekomanus, and granular in the male of C. wroughtoni.

Description: The following description is based on the holotype and paratype. Meristic data are presented in table 3. Measurements are not presented as none of the type specimens are sexually mature.

Total length: Adults unknown.

Color: Chelicerae, dorsal surfaces bicolored, mostly to entirely immaculate prodorsal half of manus paler than infuscate retrodorsal half of manus and fingers; manus markedly paler than carapace interocular surface. Carapace, pedipalps, tergites, and metasoma entirely infuscate, uniformly dark brown. Sternites III–VI immaculate, VII immaculate to slightly infuscate, dark brown, becoming darker posteriorly. Coxosternal region

infuscate, brownish, maxillary lobes darker. Legs infuscate, uniformly dark brown, similar to carapace, pedipalps, tergites, and metasoma. Genital opercula and pectines uniformly pale cream. Metasomal segments I–V similar to one another; carinae infuscate, darker than intercarinal surfaces; dorsal intercarinal surfaces similar to tergites; ventral intercarinal surfaces darker than sternites. Telson vesicle immaculate, light yellowish, markedly paler than metasomal segment V; aculeus black.

Chelicerae: Movable finger prodistal (DI) and retrodistal (DE) teeth subequal, with DE tooth only slightly smaller than DI tooth, and opposable, i.e., forming a bicusp. Fingers and manus, proventral surfaces with long, dense vestiture of macrosetae. Coxae, prodorsal surfaces without stridulatory setae (scaphotrix); promedian surfaces without chemoreceptive lamelliform setae (trichocopae).

Carapace: Carapace markedly dorsoventrally compressed, lateral surfaces sloping gently. Anterior margin concave, frontal lobes uneven, sloping medially toward deep median notch, without median projection or median depression (fig. 16E). Anterolateral margins converging gradually anteriorly, without distinct notch adjacent to posterior lateral ocelli; mediolateral margins diverging or converging slightly posteriorly (at posterolateral sulci). Three pairs of lateral ocelli (MLMa, PLMa, PDMi), all similar in size, two anterior pairs situated anteriorly, well separated from posterior pair, situated laterally. Median ocelli slightly larger than lateral ocelli, distance between ocelli greater than or equal to width of ocellus; median ocular tubercle situated posteromedially, distance from anterior carapace margin, 53% (♂), 55% (♀). Median longitudinal sulcus narrow, suturiform; continuous from median notch to interocular sulcus; anteriorly furcated; without anterocular depression. Anterior furcated sulci diverging broadly from anterior edge of median longitudinal sulcus. Median ocular tubercle raised, superciliary carinae higher than ocelli, extended anteriorly, subparallel. Interocular sulcus present. Circumocular depressions completely encircling median ocular tubercle, converging anteriorly. Posteromedian and posteromarginal sulci deep. Paired mediolateral and posterolateral sulci shallow. Median longitudinal suture continuous from median notch to median ocular tubercle, equally strong along entire length; not extending to anterior margin of carapace, terminating at or posterior to median notch. Anterior furcated sutures absent or obsolete. Anterocular furcated sutures absent. Interocular suture present, slender. Posterior sutures present but indistinct, converging on median ocular tubercle; connected anteriorly to posterior furcations of interocular suture and extending anteriorly beyond median ocular tubercle. Frontal lobes and medial region of interocular surface finely granular with smooth areas (♂) (fig. 16E) or finely granular along median longitudinal and anterior furcated sulci only, elsewhere smooth (♀); anterolateral and mediolateral surfaces uniformly finely granular; posterolateral surfaces uniformly finely granular (♂) or smooth (♀); posteromedian surfaces smooth.

Pedipalps: Pedipalp short, pedipalp femur length: posterior carapace width ratio (FL:PCW) < 0.77, femur length: carapace length ratio (FL:CL) < 0.74. Femur retrodorsal carina complete, costate-granular, similar to or more strongly developed than prodorsal carina. Dorsal secondary carina comprising scattered granules, becoming obsolete distally. Dorsomedian carina vestigial, reduced to two prominent granules, one demarcated by conspicuous macroseta. Prodorsal carina complete, granular. Promedian carina complete, comprising row of spiniform or subspiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Proventral carina complete, granular, as strongly developed as or more strongly developed than retroventral carina. Ventromedian and secondary accessory carinae absent. Retroventral carina extending partly or entirely to distal edge of segment. Retromedian carinae absent, indicated only by macrosetal rows. Dorsal intercarinal surfaces finely granular; other surfaces smooth. Patella dorsal surface convex, axis of dorsomedian carina dorsal to axis of retrodorsal carina. Dorsomedian carina entirely to predominantly costate (fig. 99). Retrodorsal carina obsolete, less strongly developed than costate retromedian carinae. Retroventral carina distinct, costate. Promedian carina and promedian process absent. Other carinae absent. Intercarinal surfaces smooth. Chela short, narrow (fig. 100); moderately (♂) to sparsely (♀) setose. Manus dorsal surface (between dorsomedian and digital carinae) flat (♂) or curved, i.e., slightly to markedly convex (♀); proximal margin (lobe) of dorsal surface moderately curved and aligned with proximal margin of retrolateral surface. Dorsomedian carina obsolete, costate on fixed finger and comprising irregular row of spiniform granules along entire length of manus. Dorsal secondary, digital, and subdigital carinae incomplete, costate-granular to costate; dorsal secondary carina comprising double row of confluent granules in proximal half; proximal half of dorsal secondary carina, distal half of subdigital carina and distal third of digital carina closely adjacent. Maximum distance between dorsomedian and dorsal secondary carinae (DMC–DSC) similar to (♂) or greater than (♀) maximum distance between dorsal secondary and digital carinae (DSC–DC). Retromedian carina complete, entirely to predominantly costate, more strongly developed than digital carina. Retrolateral secondary and secondary accessory carinae vestigial, proximal to condyle of movable finger. Retroventral and ventromedian carina distinct, costate, but retroventral carina more strongly developed. Proventral and promedian carinae similarly developed, each comprising row of spiniform granules and associated macrosetae. Prodorsal carina comprising irregular row of prominent spiniform granules and associated macrosetae. Distance between dorsomedian and promedian carinae (DMC–PMC) slightly to markedly greater than distance between promedian and proventral carinae (PMC–PVC). Manus, dorsal intercarinal surfaces densely and finely granulo-reticulate except for smooth depression at base of fixed finger; retrolateral intercarinal surfaces densely and finely granular; ventral intercarinal surface smooth; prolateral intercarinal surfaces smooth, except for few scattered spiniform granules dorsally and distally. Manus ventral surface flat, axes of retroventral and ventromedian carinae in approximately same plane. Fixed and movable fingers, intercarinal surfaces smooth; median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three situated on lobes; first (proximal) lobe of fixed finger and second lobe of movable finger larger than others, with correspondingly deeper notches in movable and fixed fingers, respectively; terminal denticles of fingers interlocking unevenly when closed, movable finger displaced retrolaterally; distinct notch near tip of fixed finger to accommodate terminal denticle of movable finger.

Trichobothria: Orthobothriotaxic, Type C, with the following segment totals (n = 4): femur, 3 (1 d, 1 i, 1 e); patella, 19 (2 d, 1 i, 3 v, 13 e, comprising 3 et, 1 est, 2 em, 2 esb, 5 eb); chela, 26 (manus, 16, comprising 2 D, 10 E, 4 V; fixed finger, 10, comprising 4 d, 4 e, 2 i). Total count of trichobothria per pedipalp: 48. Femur, i situated on dorsal surface. Patella, d₂ situated on internal surface, slightly closer to d₁ than to i; distance v₂–v₃ ca. half distance v₁–v₂. Chela, distance et–est ca. half distance est–esb; distance est–esb greater than half distance esb–eb; est aligned with dst; V₃ situated in proximal third of manus, closer to V₄ than V₂.

Legs: First pair of maxillary lobes (coxapophyses) tapering anteriorly, longer than and encircling second pair. Stridulatory organs, comprising “rasp” (granular tubercles) and “scraper” (stridulatory setae or scaphotrix), present on opposing surfaces of coxae of first pair of legs and pedipalps, respectively. Legs I–IV acarinate, except femora, with distinct pro- and retrolateral surfaces demarcated by complete, costate-granular proventral carinae. Femora, patellae, and tibiae, pro- and retrolateral surfaces each with scattered setiform macrosetae. Tibiae, prolateral surfaces, without spiniform macrosetae; I and II, retrolateral surfaces, each with two spiniform (t, st) macrosetae; III and IV, retrolateral surfaces, without spiniform macrosetae. Basitarsi I–IV, slightly dorsoventrally compressed; retrolateral margins similar, unmodified, rounded; prolateral pedal spurs present; retrolateral pedal spurs absent. Basitarsi, pro- and retrolateral surfaces, each with scattered long and short setiform macrosetae, and spiniform macrosetae, more numerous on I and II than on III and IV. Basitarsi, spiniform macrosetae, I, retrolateral: t, sb; retroventral: t, st, sb; proventral: t, st; II, retrolateral: t, sb; retroventral: t, st, sb; proventral: t, st; III, retrolateral: t, sb; retroventral: t, st, sb; proventral: t; prolateral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: t, st. Telotarsi I–IV short, stout and distally broadened in dorsal and lateral views. Laterodistal lobes rounded. Dorsomedian lobes approximately equal to laterodistal lobes; each terminating in single setiform macroseta. Telotarsi, pro- and retrolateral surfaces, each with long and short, scattered macrosetae, not arranged in definite row. Telotarsi each with pro- and retroventral rows of spiniform macrosetae, two inserted on laterodistal lobes; counts in pro- and retroventral rows similar on I and II, and on III and IV (n = 4): I, proventral: 4, retroventral: 5 or 6; II, 4, 6 or 7; III, 4, 6 (6 or 7); IV, 4, 6. Telotarsal ungues short, curved, equal to subequal; pseudonychium (dactyl) reduced, rounded.

Sternum: Shape subpentagonal (fig. 30E). Median longitudinal sulcus shallow anteriorly, deep and narrow posteriorly.

Genital operculum: Genital opercula suboval, completely divided longitudinally, partially overlapping, genital papillae present (♂) (fig. 30E); subcordate, fused, genital papillae absent (♀).

Hemispermatophore: Unknown.

Pectines: Distal edge extending past distal edge of coxa IV but not reaching distal edge of trochanter IV (♂) (fig. 30E) or reaching to distal edge of coxa IV (♀). First proximal median lamella (scape) of each pecten with mesial margin angular, approximately 90° (♂) or obtusely angular, greater than 90° but less than 180° (♀). Pectinal teeth straight and elongate (♂) or shorter and curved (♀), present along entire posterior margin; sinistral/dextral tooth count, 16/16 (♂), 15/15 (♀). Fulcra smooth proximally but densely setose (microsetae only) distally.

Mesosoma: Tergites each with shallow pair of submedian depressions, without obsolete median carina. Pretergites smooth and glabrous. Posttergites finely granular medially and unevenly finely and coarsely granular laterally (♂) or entirely smooth and glabrous (♀). Sternites IV–VI, each with paired longitudinal depressions prolateral to spiracles, absent on VII. Surface, sternites III–VII, smooth; VII with pair of weakly developed, costate ventrolateral carinae only, without posteromarginal carina.

Metasoma and telson: Metasoma fairly robust. Telson vesicle globose; dorsal surface flat; ventral surface evenly curved. Aculeus relatively long, gently curved. Dorsosubmedian carinae, segments I–IV, distinct, complete, granular or costate-granular, posterior spiniform granules obsolete (fig. 61D); V, absent. Dorsolateral carinae, segments I–IV, distinct, complete, costate-granular; V, obsolete, incomplete, reduced to few spiniform granules. Median lateral carinae, segment I, incomplete, reduced to granular row in posterior half, diagonally oriented between dorsolateral and ventrolateral carinae (fig. 62D); II–IV, obsolete row of granules posteriorly; V, incomplete, granular row, restricted to anterior two-thirds. Ventrolateral and ventrosubmedian carinae, I–IV, distinct, complete, costate on I–III, granular or costate-granular on IV, more strongly developed on III and IV than on I and II (fig. 63D). Ventrolateral carinae, segment V, distinct, complete, comprising spiniform granules, diverging posteriorly, terminal granule slightly larger than preceding granules. Ventrosubmedian carinae, segment V, vestigial, each reduced to one or two isolated spiniform granules, demarcated by conspicuous macrosetae. Ventromedian carina, segment V comprising single row of spiniform granules, unmodified posteriorly. Anal arch, segment V, dorsal carina, costate; ventral carina comprising subspiniform granules. Dorsal intercarinal surfaces, segments I–V, smooth. Lateral intercarinal surfaces, segments I–IV, granular, V, smooth. Ventral intercarinal surfaces, segments I–III, smooth, IV and V, granular (♂) or smooth (♀). Ventral surface, lateral aspect, segment IV, shallowly convex. Telson vesicle, dorsal and lateral surfaces smooth; ventral surface with four longitudinal carinae, each comprising spiniform granules.

Distribution: This species is endemic to India and known only from the type locality in the state of Tamil Nadu (fig. 80, table 1).

Ecology: Although no data are available on the ecology of this species, the habitus is consistent with the fossorial, pelophilous and semilithophilous ecomorphotypes (Prendini, 2001b). The dorsoventral compression of this species suggests that it excavates burrows under stones.

Heterometrus indus: Roewer, 1943: 226 (misidentifcation, part).

Heterometrus phipsoni Roewer, 1943: 228 (misidentification).

Heterometrus (Chersonesometrus) pelekomanus Couzijn, 1981: 19, 41, 75, 86, 137–139, 169, 170, 192, table 7, figs. 2f, 41, 58 (part); Tikader and Bastawade, 1983: 577, 641–646 (part), figs. 1696–1709; Kovařík, 1998: 136; Fet, 2000: 440 (part); Bastawade, 2002: 296; Kovařík, 2002: 17; 2004: 49; Indra, 2009: 14

Heterometrus (Chersonesometrus) palekomanus Indra, 2001: 58 (part).

Heterometrus pelekomanus: Kovařík, 2004: 49; 2009: 28, pl. J.

Heterometrus wroughtoni: Kovařík, 2004: 1, 3, 7, 46, 49, 51, 52, tables 2, 3 (misidentification, part), figs. 1–7 (misidentification); 2009: 28, 35, 36, 45, 48, tables 1, 2 (misidentification, part), pl. J (misidentification); Javed et al., 2010b: 785, table 1 (misidentification).

Heterometrus swammerdami: Nagaraj et al., 2015: 1–5, table 1, fig. 1d (misidentification).

Type Material: INDIA: Heterometrus (Chersonesometrus) pelekomanus: Holotype ♂ (SMF RII/1088/19), Nilgiris, Deccan [examined]. Paratype ♀ (SMF RII/329), Bombay [Bombay Presidency], Deccan [examined].

Couzijn (1981: 137) specifically indicated a holotype and paratype in the original description:

Fet (2000) erroneously listed all material in Couzijn's (1981) description as paratypes.

Diagnosis: Chersonesometrus pelekomanus may be separated from other species of Chersonesometrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 15C, D) in C. pelekomanus, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The frontal lobes and medial region of the carapace interocular surface are granular with smooth areas in the female (fig. 15D) of C. pelekomanus whereas the interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female of all other species except C. bastawadei, C. madraspatensis, and C. shivashankari. The carapace posterolateral surfaces of the female are smooth in C. pelekomanus but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. pelekomanus but unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, and C. wroughtoni. The pedipalps of the adult male are short, with femur length: posterior carapace width ratio (FL:PCW) < 0.77 and femur length: carapace length ratio (FL:CL) < 0.74 (fig. 101) in C. pelekomanus but long, with FL:PCW ≥ 0.80, FL:CL ≥ 0.76 in C. beccaloniae, C. madraspatensis, and C. tristis. The pedipalp patella dorsal surface is flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane (fig. 103), in C. pelekomanus but convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina in all other species except C. beccaloniae, C. hendersoni, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. pelekomanus but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. pelekomanus but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female (fig. 103) are granular in C. pelekomanus but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. pelekomanus but immaculate and paler than the femur and patella in C. shivashankari. The chela of the adult male is moderately to densely setose in C. pelekomanus but sparsely setose in C. fulvipes, C. madraspatensis, and C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and female of C. pelekomanus, flat in the male and curved, i.e., slightly to markedly convex, in the female of C. bastawadei, C. beccaloniae, C. fulvipes, C. nathanorum, and C. wroughtoni, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is markedly curved and proximal to the proximal margin of the condyle (articulation with patella) in the male and female (figs. 104, 105) of C. pelekomanus but moderately curved and aligned with or proximal to the proximal margin of the retrolateral surface, or aligned with the proximal margin of the condyle in all other species except C. hendersoni. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is greater than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. pelekomanus, but greater than the DSC–DC in the male of C. beccaloniae, C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male are entirely to predominantly granular (fig. 104) in C. pelekomanus, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is entirely to predominantly granular in C. pelekomanus, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The manus dorsal intercarinal surface is without reticulation in C. pelekomanus but shallowly reticulate in C. nathanorum. The manus ventral surface is angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. pelekomanus but flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in all other species except C. beccaloniae, C. hendersoni, and C. tristis. The legs are very dark or heavily infuscate (figs. 101, 102) in C. pelekomanus but pale or very lightly infuscate in C. bastawadei, C. fulvipes, C. hendersoni, C. madraspatensis, and C. shivashankari. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 43E–H) in C. pelekomanus but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are smooth in the male and the lateral surfaces granular in the female of C. pelekomanus, whereas the mesial surfaces are granular in the male of C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari, the mesial surfaces granular in the male and the lateral surfaces smooth in the female of C. nathanorum, and the lateral surfaces smooth in the female of C. tristis. The ventral surface of mesosomal sternite VII bears two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. pelekomanus and a pair of weakly developed ventrolateral carinae in all other species except C. beccaloniae and C. hendersoni. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 58D) in C. pelekomanus but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are granular in the male and female (fig. 60D) of C. pelekomanus but smooth in the male and female of C. wroughtoni, the male of C. bastawadei, C. fulvipes, and C. shivashankari, and the female of C. beccaloniae and C. nathanorum. The dorsolateral carinae of metasomal segment V are strong and continuous in C. pelekomanus but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is granular in the male and female of C. pelekomanus, smooth in the female of C. wroughtoni, and smooth in the male and female of all other species except C. bastawadei and C. hendersoni.

Distribution: This species is endemic to India, but the full extent of its distribution is unclear. The type localities are imprecisely indicated “Nilgiris, Deccan” and “Bombay, Deccan” (presumably, the former Bombay Presidency, as this species does not occur in the vicinity of present-day Mumbai). The Nilgiri Hills occur in the state of Tamil Nadu, not Maharashtra, as implied by Couzijn (1981), and the records from Mysore, environs of Bangalore (MNHN RS 7804) in Karnataka, and the Tirumalai Hills, Tirupati (MNHN RS 4499) in Andhra Pradesh, cited by Couzijn (1981) are referable to C. tristis. Material that most closely matches the types originates from the Ballari and Chitradurga districts of Karnataka (fig. 80, table 1).

Ecology: The known locality records of this species occur in semiarid, mixed savanna dominated by Acacia, Euphorbia, and broadleaf bushes (Grewia, Ziziphus), with bamboo thickets and moderate to thick grass on rocky granite hills, at elevations of 675–1000 m. The specimens for which data are available were collected from shallow scrapes in hard sandy-clay or sandy-loam soil under large granite boulders (fig. 8C). The habitat and habitus, especially the dorsoventrally compressed carapace and pedipalp chelae, are consistent with the pelophilous, fossorial and semilithophilous ecomorphotypes (Prendini, 2001b).

Remarks: This species was previously accommodated in subgenus Chersonesometrus of Heterometrus by various authors (e.g., Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000).

Heterometrus pelekomanus, described by Couzijn (1981) and upheld by Tikader and Bastawade (1983), was mistakenly synonymized with H. wroughtoni by Kovařík (2004), based on the alleged absence of sexual dimorphism in the shape of the pedipalp chela manus, which was cited as a putative diagnostic difference between H. wroughtoni and H. mysorensis (the latter regarded as junior synonym of C. tristis in the present contribution). However, as noted below, the absence of adult males in the type series of C. wroughtoni, the adult male of which is unknown, suggests that Kovařík (2004) mistook ontogenetic character states for diagnostic character states. Comparison of types of C. pelekomanus and C. wroughtoni revealed that the two species are not closely related.

Material Examined: INDIA: Bombay [Bombay Presidency], Deccan, 1 subad. ♀, 10 juv. ♂, 8 juv. ♀(SMF RII/329). Nilgiris, Deccan, 1 ♂ (SMF RII/1088/19). Karnataka: Ballari Distr.: Jarmali, NW of Jarmali Durgamma Temple, 14°50′N 76°28′E, 675 m, 22.xi.2009, R. Datta, degraded broadleaf savanna with euphorbias on low, rocky hills with many granite outcrops and embedded stones, in scrapes under stones in coarse, granitic sandy loam, 1 juv. ♂ (AMCC [LP 16774]), 1 juv. ♂(AMNH). Chitradurga Distr.: Chitradurga, Jogi Matti Road, 14°11′N 76°24′E, 1000 m, 21.xi.2009, R. Datta, mixed savanna with broadleaf trees, bamboo thickets and Acacia on rocky hills, moderate to thick grass layer on hard sandy clay with many granite outcrops and embedded stones, in shallow scrapes under large boulders, 1 ♂, 2 ♀, 1 subad. ♀, 19 juv. ♂, 15 juv. ♀ (AMNH), 1 juv. ♂, 2 juv. ♀ (AMCC [LP 16773]).

Erroneous Record: Puducherry: Puducherry Distr.: Pondichéry [Pondicherry/Puduchcheri/Puducherry, 11°59′N 79°49′E], M. Maindron, 1 ♀ (MNHN RS 0141 [Simon coll. 19501]).

Scorpio fulvipes: Simon, 1885: 23 (misidentification); Pocock, 1893: 304–307, 309 (misidentification, part).

Palamnaeus fulvipes madraspatensis: Pocock, 1900a: 89 (misidentification, part).

Heterometrus (Scorpio) fulvipes: Kraepelin, 1901: 271 (misidentification, part).

Type Material: INDIA: Karnataka: Ballari Distr.: Holotype ♂, 1 juv. ♂, 2 juv. ♀ paratypes (BMNH), Bellahunsi [Bellahunshi, 15°10′N 76°16′E], 30.viii.1912, Fletcher, under stones; 1 ♀, 4 juv. ♀ paratypes (BMNH), near road from Hampasagar [Hampasagara/Hampasagaram, 15°07′N 76°01′E] to Hadagalli [Hadagali, 15°01′N 75°56′E], 1.ix.1912, Y.R., under stones; 2 juv. ♀ paratypes (BMNH), Hospet [15°16′N 76°24′E], 30.viii.1912, Fletcher, under stones. Andhra Pradesh: Anantapur Distr.: Paratype ♂ (MNHN RS 0083 [Simon coll. 7015]), Wagra Karoor/Wagra Karur/Wagra Karour [Vajrakarur, 15°02′N 77°23′E], Chaper, 1 ♂, 1 ♀, 2 juv. ♂ paratypes (MNHN RS 0093), same data, except: 1893, Chaper.

Etymology: The specific epithet is a patronym honoring the Indian biologist, Tirumani Shivashankar, formerly of the University of Agricultural Sciences, Arsiker, for his pioneering contributions to the knowledge of burrowing biology and sociality in the closely related species, Chersonesometrus madraspatensis (as H. fulvipes) (Shivashankar and Veeresh, 1991; Shivashankar, 1992, 1994).

Diagnosis: Chersonesometrus shivashankari may be separated from other species of Chersonesometrus as follows. The carapace is vaulted, the lateral surfaces sloping steeply (fig. 16C, D), in C. shivashankari but slightly to markedly dorsoventrally compressed, the lateral surfaces sloping gently in all other species except C. fulvipes and C. madraspatensis. The frontal lobes and medial region of the carapace interocular surface are granular with smooth areas in the female (fig. 16D) of C. shivashankari, whereas the interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female of all other species except C. bastawadei, C. madraspatensis, and C. pelekomanus. The carapace posterolateral surfaces of the female are smooth in C. shivashankari but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. shivashankari, but subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. beccaloniae, C. hendersoni, C. nathanorum, C. pelekomanus, and C. tristis. The pedipalps of the adult male are short, with femur length: posterior carapace width ratio (FL:PCW) < 0.77 and femur length: carapace length ratio (FL:CL) < 0.74 (fig. 106A, B) in C. shivashankari but long, with FL:PCW ≥ 0.80, FL:CL ≥ 0.76 in C. beccaloniae, C. madraspatensis, and C. tristis. The pedipalp patella dorsal surface is convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina (fig. 107), in C. shivashankari but flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. shivashankari but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. shivashankari but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are granular in C. shivashankari but smooth or nearly so in C. nathanorum. The pedipalp chela is immaculate and paler than the femur and patella in C. shivashankari but infuscate and similar in color to the femur and patella in all other species. The chela of the adult male is sparsely setose in C. shivashankari but moderately to densely setose in C. beccaloniae, C. pelekomanus, and C. tristis. The chela manus dorsal surface (between the dorsomedian and digital carinae) is curved, i.e., slightly to markedly convex, in the male and female of C. shivashankari, flat in the male and curved in the female of C. bastawadei, C. beccaloniae, C. fulvipes, C. nathanorum, and C. wroughtoni, and flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface in the male (fig. 108), or aligned with the proximal margin of the condyle (articulation with patella) in the female (fig. 109) of C. shivashankari; moderately curved and aligned with the proximal margin of the retrolateral surface in the male, or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. beccaloniae and C. nathanorum; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is greater than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. shivashankari, but greater than the DSC–DC in the male of C. beccaloniae, C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male (fig. 108) are entirely to predominantly granular in C. shivashankari, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is absent or obsolete in C. shivashankari, but entirely to predominantly granular in C. beccaloniae, C. hendersoni, C. pelekomanus, C. tristis, and C. wroughtoni, and entirely to predominantly costate in C. nathanorum. The manus dorsal intercarinal surface is without reticulation in C. shivashankari but shallowly reticulate in C. nathanorum. The manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in C. shivashankari but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The legs are pale or very lightly infuscate (fig. 106) in C. shivashankari but very dark or heavily infuscate in C. beccaloniae, C. nathanorum, C. pelekomanus, C. tristis, and C. wroughtoni. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is setiform (fig. 45E–H) in C. shivashankari but spiniform in all other species except C. nathanorum. The mesial surfaces of mesosomal tergites I–VI are granular in the male and the lateral surfaces granular in the female of C. shivashankari, whereas the mesial surfaces are smooth in the male of C. beccaloniae, C. hendersoni, C. pelekomanus, and C. wroughtoni, the mesial surfaces smooth in the male and the lateral surfaces smooth in the female of C. tristis, and the lateral surfaces smooth in the female of C. nathanorum. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. shivashankari and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 61B) in C. shivashankari but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are smooth in the male and granular in the female (fig. 63B) of C. shivashankari but granular in the male and smooth in the female of C. beccaloniae and C. nathanorum, granular in the male of C. hendersoni, C. madraspatensis, C. pelekomanus, and C. tristis, and smooth in the female of C. wroughtoni. The dorsolateral carinae of metasomal segment V are strong and continuous in C. shivashankari but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is smooth in the male and female of C. shivashankari, granular in the male and female of C. bastawadei, C. hendersoni, and C. pelekomanus, and granular in the male of C. wroughtoni.

Description: The following description is based on the holotype and paratypes. Meristic data are presented in table 3 and measurements in table 4.

Total length: Adult small, maximum length, measured from anterior margin of carapace to tip of aculeus, 86 mm (♂), 82 mm (♀).

Color: Chelicerae, dorsal surfaces bicolored, mostly to entirely immaculate prodorsal half of manus paler than infuscate retrodorsal half of manus and fingers; manus markedly paler than carapace interocular surface. Carapace, tergites, and metasoma entirely infuscate, uniformly dark reddish black; pedipalp trochanter, femur, and patella, entirely infuscate, uniformly dark reddish brown to reddish black, chela immaculate, manus yellowish to reddish brown with darker fingers. Sternites III–VI immaculate or partially infuscate, VII mostly or entirely infuscate, dark brownish black, becoming darker posteriorly. Coxosternal region infuscate, brownish, maxillary lobes darker. Legs immaculate, uniformly pale yellow, contrasting with carapace, pedipalps, tergites, and metasoma (fig. 106). Genital opercula and pectines uniformly pale yellow. Metasomal segments I–V entirely infuscate, similar to one another; carinae infuscate, darker than intercarinal surfaces; dorsal intercarinal surfaces similar to tergites; ventral intercarinal surfaces darker than sternites. Telson vesicle immaculate, light yellowish, markedly paler than metasomal segment V; aculeus black.

Chelicerae: Movable finger prodistal (DI) and retrodistal (DE) teeth unequal, with DE tooth considerably smaller than DI tooth, aligned longitudinally and not opposable. Fingers and manus, proventral surfaces with long, dense vestiture of macrosetae. Coxae, prodorsal surfaces without stridulatory setae (scaphotrix); promedian surfaces without chemoreceptive lamelliform setae (trichocopae).

Carapace: Carapace vaulted, lateral surfaces sloping steeply. Anterior margin straight frontal lobes evenly rounded or truncate, with deep median notch, without median projection or median depression (fig. 16C, D). Anterolateral margins converging gradually anteriorly, without distinct notch adjacent to posterior lateral ocelli; mediolateral margins diverging or converging slightly posteriorly (at posterolateral sulci). Anterior width of posterior width, 64% (♂), 65% (♀); posterior width of length, 96% (♂), 94% (♀). Three pairs of lateral ocelli (MLMa, PLMa, PDMi), all similar in size, two anterior pairs situated anteriorly, well separated from posterior pair, situated laterally. Median ocelli moderately larger than lateral ocelli, distance between ocelli slightly less than width of ocellus; median ocular tubercle situated posteromedially, distance from anterior carapace margin, 52% (♂), 50% (♀). Median longitudinal sulcus narrow, suturiform; continuous from median notch to interocular sulcus; anteriorly furcated; without anterocular depression. Anterior furcated sulci diverging broadly from anterior edge of median longitudinal sulcus. Median ocular tubercle raised, superciliary carinae higher than ocelli, extended anteriorly, subparallel. Interocular sulcus present. Circumocular depressions completely encircling median ocular tubercle, converging anteriorly. Posteromedian and posteromarginal sulci deep. Paired mediolateral and posterolateral sulci shallow. Median longitudinal suture continuous from median notch to median ocular tubercle, equally strong along entire length; not extending to anterior margin of carapace, terminating at or posterior to median notch. Anterior furcated sutures absent or obsolete. Anterocular furcated sutures absent. Interocular suture present, slender. Posterior sutures present but indistinct, converging on median ocular tubercle; connected anteriorly to posterior furcations of interocular suture and extending anteriorly beyond median ocular tubercle. Frontal lobes and medial region of interocular surface finely granular with smooth areas (fig. 16C, D); anterolateral and mediolateral surfaces uniformly finely granular; posterolateral surfaces uniformly finely granular (♂) or smooth (♀); posteromedian surfaces smooth.

Pedipalps: Pedipalp short, pedipalp femur length: posterior carapace width ratio (FL:PCW) < 0.77, pedipalp femur length: carapace length (FL:CL) < 0.74. Femur width of length, 43% (♂), 45% (♀). Retrodorsal carina complete, granular, similar to or more strongly developed than prodorsal carina. Dorsal secondary carina comprising scattered granules, becoming obsolete distally. Dorsomedian carina vestigial, reduced to prominent granule demarcated by conspicuous macroseta. Prodorsal carina complete, granular. Promedian carina complete, comprising row of spiniform or subspiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Proventral carina complete, granular, as strongly developed as or more strongly developed than retroventral carina. Ventromedian and secondary accessory carinae absent. Retroventral carina extending partly or entirely to distal edge of segment. Retromedian carinae absent, indicated only by macrosetal rows. Dorsal intercarinal surfaces finely granular; other surfaces smooth. Patella width of length, 46% (♂), 54% (♀). Dorsal surface convex, axis of dorsomedian carina dorsal to axis of retrodorsal carina. Dorsomedian carina granular to costate-granular (fig. 107). Retrodorsal carina granular, similarly developed to granular retromedian carinae. Retroventral carina obsolete, granular. Promedian carina and promedian process absent. Other carinae absent. Dorsal and retrolateral intercarinal surfaces finely granular, other surfaces smooth. Chela short, broad (figs. 108, 109); manus height of width, 206% (♂), 180% (♀); manus length along retroventral carina of manus width, 150% (♂), 116% (♀); manus length along retroventral carina of movable finger length, 66% (♂), 69% (♀). Chela sparsely setose. Manus dorsal surface (between dorsomedian and digital carinae) curved, i.e., slightly to markedly convex; proximal margin (lobe) of dorsal surface moderately curved and proximal to proximal margin of retrolateral surface (♂) or aligned with proximal margin of condyle (articulation with patella) (♀). Dorsomedian carina obsolete, costate on fixed finger and comprising regular row of spiniform granules along entire length of manus. Dorsal secondary and subdigital carinae incomplete, granular; digital carina obsolete, granular; distal half of subdigital carina and distal third of digital carina closely adjacent. Maximum distance between dorsomedian and dorsal secondary carinae (DMC–DSC) greater than maximum distance between dorsal secondary and digital carinae (DSC–DC). Retromedian carina complete, entirely to predominantly costate, more strongly developed than digital carina. Retrolateral secondary and secondary accessory carinae vestigial, proximal to condyle of movable finger. Retroventral and ventromedian carina distinct, costate, but retroventral carina more strongly developed. Proventral and promedian carinae similarly developed, each comprising few spiniform granules and associated macrosetae. Prodorsal carina obsolete, comprising few spiniform granules and associated macrosetae. Distance between dorsomedian and promedian carinae (DMC–PMC) slightly to markedly greater than distance between promedian and proventral carinae (PMC–PVC). Manus, dorsal intercarinal and retrolateral intercarinal surfaces densely and coarsely granular except for smooth depression at base of fixed finger; ventral intercarinal surface smooth; prolateral intercarinal surfaces very sparsely and finely granular, more densely dorsally and distally. Manus ventral surface flat, axes of retroventral and ventromedian carinae in approximately same plane. Fixed and movable fingers, intercarinal surfaces finely granular (♂) to smooth (♀); median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three situated on lobes; first (proximal) lobe of fixed finger and second lobe of movable finger larger than others, with correspondingly deeper notches in movable and fixed fingers, respectively; terminal denticles of fingers interlocking unevenly when closed, movable finger displaced retrolaterally; distinct notch near tip of fixed finger to accommodate terminal denticle of movable finger.

Trichobothria: Orthobothriotaxic, Type C, with the following segment totals (n = 4): femur, 3 (1 d, 1 i, 1 e); patella, 19 (2 d, 1 i, 3 v, 13 e, comprising 3 et, 1 est, 2 em, 2 esb, 5 eb); chela, 26 (manus, 16, comprising 2 D, 10 E, 4 V; fixed finger, 10, comprising 4 d, 4 e, 2 i). Total count of trichobothria per pedipalp: 48. Femur, i situated on dorsal surface. Patella, d₂ situated on internal surface, slightly closer to d₁ than to i; distance v₂–v₃ ca. one-third distance v₁–v₂. Chela, distance et–est ca. half distance est–esb; distance est–esb greater than half distance esb–eb; est aligned with dst; V₃ situated in proximal third of manus, closer to V₄ than V₂.

Legs: First pair of maxillary lobes (coxapophyses) tapering anteriorly, longer than and encircling second pair. Stridulatory organs, comprising “rasp” (granular tubercles) and “scraper” (stridulatory setae or scaphotrix), present on opposing surfaces of coxae of first pair of legs and pedipalps, respectively. Legs I–IV acarinate, except femora, with distinct pro- and retrolateral surfaces demarcated by partial, granular proventral carinae. Femora, patellae, and tibiae, pro- and retrolateral surfaces each with scattered setiform macrosetae. Tibiae, prolateral surfaces, without spiniform macrosetae; I and II, retrolateral surfaces, each with two spiniform (t, st) macrosetae; III and IV, retrolateral surfaces, without spiniform macrosetae. Basitarsi I–IV, slightly dorsoventrally compressed; retrolateral margins similar, unmodified, rounded; prolateral pedal spurs present (fig. 45E–H); retrolateral pedal spurs absent. Basitarsi, pro- and retrolateral surfaces, each with scattered long and short setiform macrosetae, and spiniform macrosetae, more numerous on I and II than III and IV. Basitarsi, spiniform macrosetae, I, retrolateral: t, sb; retroventral: t, sb; proventral: t, st; II, retrolateral: t, sb; retroventral: t, sb; proventral: t, st; III, retrolateral: t, sb; retroventral: t, st; proventral: t; prolateral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: t, st. Telotarsi I–IV short, stout and distally broadened in dorsal and lateral views. Laterodistal lobes rounded. Dorsomedian lobes approximately equal to laterodistal lobes; each terminating in single setiform macroseta. Telotarsi, pro- and retrolateral surfaces, each with long and short, scattered macrosetae, not arranged in definite row. Telotarsi each with pro- and retroventral rows of spiniform macrosetae, two inserted on laterodistal lobes; counts in pro- and retroventral rows similar on I and II, and on III and IV (n = 6): I, proventral: 3 (3 or 4), retroventral: 6 (5 or 6); II, 4 (3 or 4), 6 (5 or 6); III and IV, 4, 6 (5 or 6). Telotarsal ungues short, curved, equal to subequal; pseudonychium (dactyl) reduced, rounded.

Sternum: Shape subpentagonal (fig. 30C, D). Median longitudinal sulcus shallow anteriorly, deep and narrow posteriorly.

Genital operculum: Genital opercula suboval, completely divided longitudinally, partially overlapping, genital papillae present (♂) (fig. 30C); subcordate, fused, genital papillae absent (♀) (fig. 30D).

Hemispermatophore: Lamelliform, with complex, folded capsule and accessory distal lobe protruding between articular suture and distal lobe (hook). Distal lamina with distal crest truncate, unfolded.

Pectines: Distal edge extending past distal edge of coxa IV but not reaching distal edge of trochanter IV (fig. 30C, D). First proximal median lamella (scape) of each pecten with mesial margin angular, approximately 90° (♂) or obtusely angular, greater than 90° but less than 180° (♀). Pectinal teeth straight and elongate (♂) or shorter and curved (♀), present along entire posterior margin; sinistral/dextral tooth count, 15/16 (♂), 13–15/13–15 (n = 2) (♀). Fulcra smooth proximally but densely setose (microsetae only) distally.

Mesosoma: Tergites each with shallow pair of submedian depressions, without obsolete median carina. Pretergites smooth and glabrous. Posttergites finely granular (♂) or smooth and glabrous (♀) medially, unevenly finely and coarsely granular to predominantly coarsely granular laterally. Sternites IV–VI, each with paired longitudinal depressions prolateral to spiracles, absent on VII. Surface, sternites III–VII, smooth; VII with pair of weakly developed, costate ventrolateral carinae only, without posteromarginal carina. Sternite VII, length of width, 69% (♂), 60% (♀).

Metasoma and telson: Metasomal segments I–V progressively increasing in length, decreasing in width; segment V, width of segment I, width, 76% (♂), 70% (♀). Metasoma fairly robust, width of length, segment I, 94% (♂), 93% (♀); II, 83% (♂), 78% (♀); III, 72% (♂, ♀); IV, 61% (♂), 59% (♀); V, 41% (♂), 42% (♀). Telson vesicle, width of metasomal segment V, width, 93% (♂), 86% (♀); vesicle globose, height of length, 59% (♂), 47% (♀); dorsal surface flat; ventral surface evenly curved. Aculeus relatively long, gently curved, length of vesicle length, 64% (♂), 58% (♀). Length metasoma and telson, of total length, 55% (♂, ♀). Dorsosubmedian carinae, segments I–IV, distinct, complete, granular or costate-granular, posterior spiniform granules obsolete (fig. 61B); V, vestigial. Dorsolateral carinae, segments I–V, distinct, complete, costate to costate-granular. Median lateral carinae, segment I, incomplete, reduced to granular row in posterior half, diagonally oriented between dorsolateral and ventrolateral carinae (fig. 62B); II–IV, obsolete row of granules posteriorly; V, incomplete, granular row, restricted to anterior two-thirds. Ventrolateral and ventrosubmedian carinae, segments I–IV, distinct, complete, costate on I–III, granular or costate-granular on IV, more strongly developed on III and IV than on I and II (fig. 63B). Ventrolateral carinae, segment V, distinct, complete, comprising spiniform granules, diverging posteriorly, terminal granule slightly larger than preceding granules. Ventrosubmedian carinae, segment V, vestigial, each reduced to one or two isolated spiniform granules, demarcated by conspicuous macrosetae. Ventromedian carina, segment V comprising single row of spiniform granules, unmodified posteriorly. Anal arch, segment V, dorsal carina, costate; ventral carina comprising subspiniform granules. Dorsal and lateral intercarinal surfaces, segments I–V, smooth. Ventral intercarinal surfaces, segments I–III, smooth, IV and V, smooth (♂) or granular (♀). Ventral surface, lateral aspect, segment IV, shallowly convex. Telson vesicle, dorsal and lateral surfaces smooth; ventral surface with four longitudinal carinae, each comprising spiniform granules.

Distribution: This species is endemic to India and presently recorded only from the states of Andhra Pradesh and Karnataka (fig. 80, table 1). Its distribution is allopatric with the morphologically similar C. bastawadei, C. fulvipes, C. madraspatensis, and C. wroughtoni.

Ecology: The habitus is consistent with the fossorial, pelophilous ecomorphotype (Prendini, 2001b) and the limited data available suggest it constructs burrows under stones.

Palamnaeus tristis Henderson, 1919: 380–381, pl. XXI, figs. 3, 4.

Heterometrus tristis: Takashima, 1945: 94; Kovařík, 2004: 1, 44, 49, 51, 52, tables 2, 3; 2009: 35, 36, 45, tables 1, 2; Javed et al., 2010b: 785, table 1.

Heterometrus (Chersonesometrus) pelekomanus: Couzijn, 1981: 137, 139, 169, 170, fig. 58 (misidentification, part); Tikader and Bastawade, 1983: 577, 641, 642, 644, 646 (part); Fet, 2000: 440 (part).

Heterometrus (Chersonesometrus) tristis: Couzijn, 1981: 42, 87, 156–158, 169, 171, 192, table 7, figs. 51, 59; Tikader and Bastawade, 1983: 574, 582–587, figs. 1539–1552; Kovařík, 1998: 137; Fet, 2000: 442; Indra 2009: 141.

Heterometrus (Chersonesometrus) palekomanus: Indra, 2001: 56, 58 (misidentification, part).

Heterometrus mysorensis Kovařík, 2004: 1, 7, 29, 30, 51, 52, tables 1–3, fig. 21; 2009: 35, 36, 42, 46, 48, 49, 86, 101, 109, tables 1, 2, figs. 89–94, 215, 216, 259–262; syn. nov.

Heterometrus wroughtoni: Kovařík, 2004: 1, 7, 46, 49, 51, 52, tables 2, 3 (misidentification, part); 2009: 35, 36, tables 1, 2 (misidentification, part).

Heterometrus (Chersonesometrus) pelekomanus: Indra, 2009: 141 (misidentification, part).

Heterometrus (Chersonesometrus) tristis: Indra, 2009: 141.

Type Material: INDIA: Karnataka: Mandya Distr.: Heterometrus mysorensis: Holotype ♂(FKPC), Mysore, Maddur env. [Madduru, 12°35′N 77°03′E], ca. 900 m, 1983. Andrha Pradesh: Arcot Distr.: Palamnaeus tristis: Holotype ♂ (ZSI 2422/17), Tirupati Hills [13°39′N 79°25′E], North Arcot District, S.K. Sundaracharlu.

Although Henderson (1919: 380) listed two males from the Tirupati Hills, North Arcot District, and two females and one immature male from the Venkatagiri Hills, Nellore District, the description was based on an adult male, and the “type-specimen” explicitly identified as ZSI 2422/17. Therefore, one of the two males from the Tirupati Hills is the holotype and no other specimens are types. Couzijn (1981: 157) mistakenly listed an adult male holotype and adult male paratype from the Tirupati Hills, and an allotype female from the Venkatagiri Hills, all with the number ZSI 2160/18. Tikader and Bastawade (1983: 587), mistakenly listed a male and a female “type specimens” (ZSI 2160/18), and Fet (2000: 442) followed Couzijn (1981).

Diagnosis: Chersonesometrus tristis may be separated from other species of Chersonesometrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 15E, F) in C. tristis, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female (fig. 15F) of C. tristis, whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the female of C. bastawadei, C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are smooth in C. tristis but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. tristis but unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. bastawadei, C. fulvipes, C. madraspatensis, C. shivashankari, and C. wroughtoni. The pedipalps of the adult male are long, with femur length: posterior carapace width ratio (FL:PCW) ≥ 0.80 and femur length: carapace length ratio (FL:CL) ≥ 0.76 (fig. 110) in C. tristis but short, with FL:PCW < 0.77, FL:CL < 0.74 in all other species except C. beccaloniae and C. madraspatensis. The pedipalp patella dorsal surface is flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane (fig. 112), in C. tristis but convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina in all other species except C. beccaloniae, C. hendersoni, and C. pelekomanus. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. tristis but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. tristis but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female (fig. 112) are granular in C. tristis but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. tristis but immaculate and paler than the femur and patella in C. shivashankari. The chela of the adult male is moderately to densely setose in C. tristis but sparsely setose in C. fulvipes, C. madraspatensis, and C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and female of C. tristis, flat in the male and curved, i.e., slightly to markedly convex, in the female of C. bastawadei, C. beccaloniae, C. fulvipes, C. nathanorum, and C. wroughtoni, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface in the male (fig. 113A), or aligned with the proximal margin of the condyle (articulation with patella) in the female (fig. 113B) of C. tristis; moderately curved and aligned with the proximal margin of the retrolateral surface in the male, or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. beccaloniae and C. nathanorum; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is similar to the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. tristis, but greater than the DSC–DC in the male of all other species except C. beccaloniae, C. madraspatensis, C. nathanorum. The dorsal secondary and subdigital carinae of the male are entirely to predominantly granular (fig. 113A) in C. tristis, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is entirely to predominantly granular in C. tristis, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The manus dorsal intercarinal surface is without reticulation in C. tristis but shallowly reticulate in C. nathanorum. The manus ventral surface is angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. tristis but flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in all other species except C. beccaloniae, C. hendersoni, and C. pelekomanus. The legs are very dark or heavily infuscate (figs. 9B, 110, 111) in C. tristis but pale or very lightly infuscate in C. bastawadei, C. fulvipes, C. hendersoni, C. madraspatensis, and C. shivashankari. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 43I–L) in C. tristis but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are smooth in the male and the lateral surfaces smooth in the female of C. tristis, whereas the mesial surfaces are granular in the male of C. nathanorum, the lateral surfaces granular in the female of C. beccaloniae, C. hendersoni, C. pelekomanus, and C. wroughtoni, and the mesial surfaces granular in the male and the lateral surfaces granular in the female of C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. tristis and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are costate on metasomal segments I and II or I–III and granular or costate-granular on III and IV or IV (fig. 58E) in C. tristis but granular or costate-granular on segments I–IV in all other species, except the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are granular in the male and female (fig. 60E) of C. tristis but smooth in the male and female of C. wroughtoni, the male of C. bastawadei, C. fulvipes, and C. shivashankari, and the female of C. beccaloniae and C. nathanorum. The dorsolateral carinae of metasomal segment V are strong and continuous in C. tristis but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is smooth in the male and female of C. tristis, granular in the male and female of C. bastawadei, C. hendersoni, and C. pelekomanus, and granular in the male of C. wroughtoni.

Distribution: This species is endemic to India and has been recorded in the states of Andhra Pradesh, Karnataka, and Tamil Nadu (fig. 80, table 1). Records of H. pelekomanus from Mysore, environs of Bangalore (MNHN RS 7804) in Karnataka, and the Tirumalai Hills, Tirupati (MNHN RS 4499) in Andhra Pradesh, cited by Couzijn (1981), are referable to C. tristis.

Ecology: Chersonesometrus tristis inhabits the rocky granite or quartzite/sandstone slopes and ravines of several small mountain ranges, e.g., the Tirumalai Hills and the Venkatagiri Hills, at elevations of 150–915 m on the Deccan Plateau (fig. 7B). The known locality records occur in arid to semiarid areas of deciduous broadleaf woodland on clayey to sandy-loam soils. As noted by Henderson (1919: 380), this species inhabits “short burrows,” or scrapes, under large stones, as well as the cracks and crevices of rock and earthen walls,usually in humid, shaded locations. Specimens observed with UV light detection were abundant in suitable habitat at night and under stones during daytime (P. Jain, personal commun.). The habitat and habitus, especially the dorsoventrally compressed carapace and pedipalp chelae, are consistent with the pelophilous, fossorial and semilithophilous ecomorphotypes (Prendini, 2001b). This species is sympatric with the buthids Charmus indicus, Hottentotta rugiscutis, and Lychas tricarinatus and several species of Reddyanus across its distribution.

Conservation Status: Chersonesometrus tristis is occasionally offered in the exotic pet trade, under the name H. mysorensis.

Remarks: This species was previously accommodated in subgenus Chersonesometrus of Heterometrus by various authors (e.g., Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000).

Kovařík (2004: 29) described H. mysorensis, collected in Mysore and Chittoor, and compared it with H. phipsoni, noting that H. mysorensis “is close to H. phipsoni, but has conspicuous rows of granules (carinae) on the chela and only slightly tuberculate patella of pedipalp, whereas in H. phipsoni the chela is strongly granulose.”

Kovařík (2004: 29) did not examine the holotype of Palamnaeus tristis but suggested based on Henderson's (1919: 380) figure 3 that it could be a junior synonym of H. gravimanus “with which it shares the shape of pedipalps and the presence of carinae on the chela.” However, Couzijn (1981) and Tikader and Bastawade (1983) each presented detailed redescriptions of H. tristis, based on the holotype, accompanied by illustrations in the case of Tikader and Bastawade (1983), from which it is clear that H. mysorensis is conspecific with C. tristis, based on the similar carapace shape and granulation, chela shape, granulation, and carination, as well as metric and meristic data. For example, the total length of the male of H. mysorensis is 118 mm, the total length of the male of C. tristis is 116 mm, according to Henderson (1919), or 120 mm according to Couzijn (1981), and the total length of the female is 110 mm (Couzijn, 1981). The pectinal tooth count is 16/17 in the male of H. mysorensis, compared with 17/17 in the male and 14/15 in the female of C. tristis. The counts of pro- and retroventral spiniform macrosetae on the leg telotarsi are 5–6/6–7 on legs III and IV in H. mysorensis, 4/6, 4–5/6–7, 5/6–7, 5/7 on legs I–IV in the male and 4/5–6, 4/6, 5/6–7, 5/7 on legs I–IV in the female of C. tristis. Kovařík (2004) distinguished H. mysorensis from H. wroughtoni and H. pelekomanus, which was regarded as a junior synonym of the latter, by the sexual dimorphism in the pedipalp shape: the pedipalps of the male are longer and narrower than those of the female in H. mysorensis unlike H. pelekomanus, in which they are similar (this character cannot be evaluated for H. wroughtoni, as the adult male of that species is unknown). However, adult male specimens from the vicinity of the type locality of P. tristis, in the Chittoor area, examined during the present investigation (fig. 80), which appear to be conspecific with adult female specimens from Chittoor identified as H. mysorensis by Kovařík (2009: 42, 86, figs. 92–94) display sexually dimorphic pedipalps, proving that the character cited as diagnostic for H. mysorensis by Kovařík (2004, 2009) occurs in C. tristis. DNA sequence data obtained from samples originating from the vicinities of the type localities of H. mysorensis and C. tristis also failed to differentiate among the two species. The combination of evidence leads to the inescapable conclusion that H. mysorensis and C. tristis are conspecific and the following new synonym is therefore presented: Heterometrus mysorensis Kovařík, 2004 = Chersonesometrus tristis (Henderson, 1919), syn. nov.

Material Examined: INDIA: viii.1973, Chowdaiah, 1 subad. ♀, 2 juv. ♂ (MNHN RS 6558). Andhra Pradesh: Chittoor Distr.: Ithepalli/Agarala, NW of, 13°35′N 79°15′E, 340 m, 1.xii.2004, S. Basi, rocky granite hillslopes and ravine, broadleaf savanna with moderate grass layer on sandy loam, many large and small granite boulders, in crevices or cracks in earthen and rock walls in ravine, 2 ♂ [1 ♂ remains], 1 subad. ♂, 1 juv. ♂(AMNH), 1 juv. ♂ (AMCC [LP 13135]); Palmaner [13°12′N 78°45′E], 6 mi. W, 725 m, 26.ii.1962, E.S. Ross and D.Q. Cavagnaro, 1 juv. ♀ (CAS 9070407); Tirumalai Hills [Tirumala, 13°41′N 79°21′E], Tirupati, R.P. Sreenivasa-Reddy, 1 subad. ♀, 2 juv. ♂, 2 juv. ♀, 1 subad. (MNHN RS 4499). Kadapa Distr.: Highway 58, between Rajukunta and Rapur, 14°11′N 79°25′E, 289 m, 12.ii.2004, S. Basi, steep mountain slope, semideciduous broadleaf forest with shale outcrops and loose stones on clayey-loam, in scrape under stone, 1 juv. ♂ (AMCC [LP 13136]), 14°12′N 79°28′E, 148 m, 12.ii.2004, S. Basi, slopes of small rocky hill, semideciduous broadleaf woodland on clayey loam, many quartzite/sandstone outcrops and loose stones, in shallow scrape under stone, 1 juv. ♀ (AMNH). Karnataka: Bengaluru Distr.: Mysore state, env. Bangalore [Bengaluru, 12°59′N 77°35′E], 8.v.1974, Geethamali, 1 juv. ♀ (MNHN RS 7804). Cuddapah Distr.: Mysore, Kamalapuram [14°37′N 78°40′E], 6.ix.1912, Y.R. C.M., 1 juv. ♀ (BMNH). Mandya Distr.: Maddur [Madduru, 12°35′N 77°03′E], iii.2003 [juv. born in captivity on 26.ix.2001], 1 ♀ (AMNH), 1 juv. ♂(AMCC [LP 2275]), 1 juv. ♀ (AMCC [LP 2276]). Mysore Distr: Mysore [Mysuru, 12°18′N 76°38′E], M. Seiter, 1 ♂, 1 ♀, 1 juv. (NHMW 28529).

Palamnaeus wroughtoni Pocock, 1899: 745; 1900a: 85, 89, fig, 26; Henderson, 1919: 381; Tikader, 1973: 263.

Heterometrus wroughtoni Takashima, 1945: 93; Kovařík, 2004: 1, 7, 46, 48, 49, 51, 52, tables 2, 3 (part), fig. 31; 2009: 35, 36, 45, 48, 49, 97, 101, tables 1, 2 (part), figs. 168–173, 218, 219; Tahir and Prendini, 2014: 1, 2, 6, 9.

Heterometrus (Chersonesometrus) wroughtoni: Couzijn, 1981: 16, 44, 88, 158–160, 171, 192, table 7, figs. 52, 59; Tikader and Bastawade, 1983: 575, 593–598, figs. 1566–1580; Kovařík, 1998: 137; Fet, 2000: 443.

Heterometrus (Chersonesometrus) wrongtoni: Bastawade, 2002: 296.

Heterometrus (Chersonesometrus) wronghtoni: Bastawade and Borkar, 2008: 212, 219, 220 (part).

Type Material: INDIA: Karnataka: Belgaum Distr.: Palamnaeus wroughtoni: Lectotype ♀, paralectotypes: 3 ♀, 3 juv. ♂, 1 juv. ♀ (BMNH 1897.9.17.1-4.11.20 [BNHS 1163, 1165, 1168, 1169, 1171]), Belgaum [Belagavi, 15°52′N 74°30′E], W.A. Talbot [examined]. Maharashtra: Kolhapur Distr.: P. wroughtoni: Paralectotype ♀(BMNH 1897.9.5.2 [BNHS 1027]), Gadingal [Gadhinglaj, 16°14′N 74°21′E], Kolhapur, Wray [examined].

Pocock (1899) did not specify types among the material listed from Belgaum and Gadingal, hence all specimens in the original description are syntypes. A single type specimen from Gadingal is numbered BMNH 1897.9.5.2 [BNHS 1027]. However, Couzijn (1981: 158) mistakenly identified one specimen as the holotype: “Maharashtra: Gadingal (Gad-Hinglaj), Belgaum (1 ♀ad. Holotype, 3 ♂ juv., 4 ♀ juv., leg W.A. Talbot, BM 1897.9.17.1-4.11.20).” Tikader and Bastawade (1983: 598) again referred to “the type specimen 1 ♀” and the type locality as Belgaum. Fet (2000: 443) followed by referring incorrectly to the “holotype: ♀ (BMNH 1897.9.17.1-4.11.20), Gadingal (Gad-Hinglaj), Kolhapur District, Maharashtra; Belgaum, Karnataka, India” and seven paratypes with the same data and number. Examination of the BMNH specimens during the present investigation revealed that all specimens numbered BMNH 1897.17.1-4.11.20 are from Belgaum not Gadingal. Kovařík (2004: 46) correctly designated a lectotype female and the remaining specimens of BMNH 1897.17.1-4.11.20 as paralectotypes, but misidentified the three immature males as adults, and failed to mention the specimen from Gadingal, which is also a paralectotype.

Diagnosis: Chersonesometrus wroughtoni may be separated from other species of Chersonesometrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 16B), in C. wroughtoni, but vaulted, the lateral surfaces sloping steeply, in C. fulvipes, C. madraspatensis, and C. shivashankari. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female (fig. 16B) of C. wroughtoni, whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in the female of C. bastawadei, C. madraspatensis, C. pelekomanus, and C. shivashankari. The carapace posterolateral surfaces of the female are smooth in C. wroughtoni but granular in C. bastawadei, C. fulvipes, and C. madraspatensis. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable in C. wroughtoni, but subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable, i.e., forming a bicusp, in C. beccaloniae, C. hendersoni, C. nathanorum, C. pelekomanus, and C. tristis. The pedipalp patella dorsal surface is convex, with the axis of the dorsomedian carina dorsal to the axis of the retrodorsal carina (fig. 115) in C. wroughtoni but flat or nearly so, with the axes of the dorsomedian and retrodorsal carinae in the same plane, in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The patella retrodorsal carina of the female is as strongly developed as or more strongly developed than the retromedian carinae in C. wroughtoni but absent or obsolete in C. nathanorum. The retromedian carinae are granular in C. wroughtoni but costate in C. nathanorum. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female (fig. 115) are granular in C. wroughtoni but smooth or nearly so in C. nathanorum. The pedipalp chela is infuscate and similar in color to the femur and patella in C. wroughtoni but immaculate and paler than the femur and patella in C. shivashankari. The chela manus dorsal surface (between the dorsomedian and digital carinae) is flat in the male and curved, i.e., slightly to markedly convex, in the female of C. wroughtoni, flat in the male and female of C. hendersoni, C. pelekomanus, and C. tristis, and curved in the male and female of C. madraspatensis and C. shivashankari. The proximal margin (lobe) of the chela manus dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface in the male, or aligned with the proximal margin of the condyle (articulation with patella) in the female (fig. 116) of C. wroughtoni; moderately curved and aligned with the proximal margin of the retrolateral surface in the male, or aligned with or proximal to the proximal margin of the retrolateral surface in the female of C. beccaloniae and C. nathanorum; and markedly curved and proximal to the proximal margin of the condyle in the male and female of C. hendersoni and C. pelekomanus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is greater than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male of C. wroughtoni, but greater than the DSC–DC in the male of C. beccaloniae, C. madraspatensis, C. nathanorum, and C. tristis. The dorsal secondary and subdigital carinae of the male are entirely to predominantly granular in C. wroughtoni, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei and C. fulvipes. The digital carina is entirely to predominantly granular in C. wroughtoni, entirely to predominantly costate in C. nathanorum, and absent or obsolete in C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari. The manus dorsal intercarinal surface is without reticulation in C. wroughtoni but shallowly reticulate in C. nathanorum. The manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in C. wroughtoni but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis. The legs are very dark or heavily infuscate (fig. 81C, D) in C. wroughtoni but pale or very lightly infuscate in C. bastawadei, C. fulvipes, C. hendersoni, C. madraspatensis, and C. shivashankari. Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 44I–L) in C. wroughtoni but setiform in C. nathanorum and C. shivashankari. The mesial surfaces of mesosomal tergites I–VI are smooth in the male and the lateral surfaces granular in the female of C. wroughtoni, whereas the mesial surfaces are granular in the male of C. bastawadei, C. fulvipes, C. madraspatensis, and C. shivashankari, the mesial surfaces granular in the male and the lateral surfaces smooth in the female of C. nathanorum, and the lateral surfaces smooth in the female of C. tristis. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in C. wroughtoni and two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in C. beccaloniae, C. hendersoni, and C. pelekomanus. The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 61E) in C. wroughtoni but costate on segments I and II or I–III and granular or costate-granular on III and IV or IV in C. tristis, and the female of C. beccaloniae. The ventral intercarinal surfaces of segment IV are smooth in the male and female (fig. 63E) of C. wroughtoni but granular in the male and female of C. hendersoni, C. madraspatensis, C. pelekomanus, and C. tristis, the male of C. beccaloniae and C. nathanorum, and the female of C. bastawadei, C. fulvipes, and C. shivashankari. The dorsolateral carinae of metasomal segment V are strong and continuous in C. wroughtoni but weak and discontinuous to absent in C. nathanorum. The dorsal intercarinal surface of segment V is granular in the male and smooth in the female of C. wroughtoni, granular in female of C. bastawadei, C. hendersoni and C. pelekomanus, and smooth in the male of all other species.

Distribution: This species is endemic to India and known from only two localities in the states of Karnataka and Maharashtra (fig. 80, table 1).

Ecology: Little ecological data are available for this species, the known records of which occur on arid, rocky hills on the Deccan Plateau. This species probably constructs burrows under stones, similar to C. bastawadei and C. fulvipes. The habitus is consistent with the pelophilous, fossorial ecomorphotype (Prendini, 2001b).

Conservation Status: This species has not been seen in over a century. Its habitat has been extensively modified and it may be extinct.

Remarks: This species was previously accommodated in subgenus Chersonesometrus of Heterometrus by various authors (e.g., Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000). Heterometrus pelekomanus, described by Couzijn (1981), and upheld by Tikader and Bastawade (1983), was mistakenly synonymized with H. wroughtoni by Kovařík (2004) based on the absence of sexual dimorphism in the shape of the pedipalp chela manus, cited as a putative diagnostic difference between H. wroughtoni and H. mysorensis (= C. tristis). The absence of adult males in the type series of C. wroughtoni, the adult male of which remains unknown, suggests that Kovařík (2004) mistook ontogenetic character states for diagnostic character states.

Type species: Heterometrus (Chersonesometrus) scaber obscurus Couzijn, 1981: 87, 146–149, 170, figs. 46, 58 [= Deccanometrus obscurus (Couzijn, 1981), comb. et stat. nov.], here designated.

Heterometrus (Chersonesometrus): Couzijn, 1981: 80, 83, 131, 133, 161, 169–171, 173, 182, 184, 186–188, 192, figs. 21, 57–59, 66b, 68, 69 (part); Tikader and Bastawade, 1983: 519, 520, 573 (part); Kovařík, 2004: 1.

Heterometrus: Kovařík, 2004: 1, 2, 49, 51, tables 1, 2 (part); 2009: 34, 35, table 1 (part); Mirza and Sanap, 2009: 486 (part); Javed et al., 2010a: 143 (part); Tahir and Prendini, 2014: 1–5, 11, 20, 21, fig. 1 (part); Aswathi and Sureshan, 2017: 9847 (part).

Diagnosis: Species of Deccanometrus may be separated from other Asian scorpionid genera as follows. The carapace is slightly to markedly dorsoventrally compressed, the lateral surfaces sloping gently (figs. 19, 21), in all except two species of Deccanometrus, D. latimanus and D. xanthopus, in which the carapace is vaulted, the lateral surfaces sloping steeply (fig. 20), as in Gigantometrus, Heterometrus, Javanimetrus, three species of Chersonesometrus, C. fulvipes, C. madraspatensis, and C. shivashankari, two species of Sahyadrimetrus, S. mathewi and S. scaber, and all except three species of Srilankametrus, S. indus, S. pococki, and S. serratus. The carapace anterior margin is concave, with the frontal lobes uneven and sloping medially, in all except two species of Deccanometrus, D. latimanus and D. xanthopus, in which the anterior margin is straight, with the frontal lobes evenly rounded or truncate (fig. 20), as in Gigantometrus, Sahyadrimetrus, and some species of Chersonesometrus. The carapace anterolateral margins converge gradually anteriorly in Deccanometrus but are subparallel anteriorly in Javanimetrus and Srilankametrus; the mediolateral margins diverge or converge slightly posteriorly (at the posterolateral sulci) in Deccanometrus, but converge markedly posteriorly (at the posterolateral sulci) in Gigantometrus, Heterometrus, and three species of Sahyadrimetrus, S. kanarensis, S. mathewi, and S. scaber. The carapace rostrolateral margin is entire in Deccanometrus but distinctly incised adjacent to the posterior lateral ocelli in Heterometrus. Anterocular extensions of the superciliary carinae are present in Deccanometrus but absent in Srilankametrus. The median ocelli are relatively small, the distance between them equal to or greater than the width of an ocellus in Deccanometrus but relatively large, the distance between them less than the width of an ocellus in Heterometrus; the median ocular tubercle is situated anteromedially to medially, the distance from carapace anterior margin : carapace length (CAM:CL) 0.40–0.50 in Deccanometrus, but posteromedially, CAM:CL 0.51–0.62 in Chersonesometrus. The interocular suture is present in all species of Deccanometrus except D. liurus, in which it is absent, as in Heterometrus, Javanimetrus, Sahyadrimetrus, and two species of Srilankametrus. The carapace posterior sutures are present, extending past the median ocular tubercle, and connected anteriorly to the posterior bifurcations of the interocular suture in all species of Deccanometrus except D. liurus, in which the posterior sutures and posterior bifurcations of the interocular suture are disconnected; present, extending to the median ocular tubercle, and connected by a short cross-suture anterior to the postocular depression in Heterometrus; and absent in Javanimetrus, Sahyadrimetrus, and two species of Srilankametrus. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are unequal, with the DE tooth considerably smaller than the DI tooth, aligned longitudinally and not opposable, in all Deccanometrus except D. liurus and D. ubicki, in which the DI and DE teeth are opposable, i.e., forming a bicusp, resembling Heterometrus and some species of Chersonesometrus, in which the DI and DE teeth are subequal, with the DE tooth only slightly smaller than the DI tooth, and opposable. The pedipalp patella dorsomedian carina of the female is entirely to predominantly granular or costate (fig. 142) in all except three species of Deccanometrus, D. latimanus, D. liurus, and D. ubicki, in which the carina is absent or obsolete (figs. 124, 128, 140), as in Heterometrus, Javanimetrus, all except one species of Sahyadrimetrus, S. rugosus, and all except one species of Srilankametrus, S. caesar. The patella retromedian carinae of the female are absent or obsolete (figs. 124, 128, 140, 142) in all except three species of Deccanometrus, D. bengalensis, D. obscurus, and D. phipsoni, in which the carinae are granular or costate, as in Chersonesometrus, three species of Sahyadrimetrus, S. barberi, S. rugosus, and S. tikaderi, and two species of Srilankametrus, S. gravimanus and S. yaleensis. The prominent spiniform granule of the patella proventral carina is absent in Deccanometrus (figs. 121, 124, 128, 132, 136, 140, 142) but present in Heterometrus. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are smooth or nearly so (figs. 124, 128, 140, 142) in all species of Deccanometrus except D. obscurus and D. phipsoni, in which the surfaces are granular, as in all species of Chersonesometrus except C. nathanorum. The pedipalp chela of the adult male is moderately to densely setose (figs. 122B–D, 125, 133, 137, 141A) in all species of Deccanometrus except D. xanthopus, in which the chela is sparsely setose (fig. 143A), as in Gigantometrus, Heterometrus, Javanimetrus, most species of Srilankametrus, and some species Chersonesometrus. The distance between the chela manus dorsomedian and promedian carinae or setal rows (DMC–PMC) is slightly to markedly greater than the distance between the promedian and proventral carinae or setal rows (PMC–PVC) in Deccanometrus whereas the DMC–PMC is similar to the PMC–PVC in Heterometrus. The dorsomedian carina becomes obsolete proximally in all except two species of Deccanometrus, D. obscurus and D. phipsoni, in which the carina is continuous to the proximal edge of the manus (figs. 133, 134, 137, 138), as in Chersonesometrus and Gigantometrus. The dorsal secondary, subdigital, and digital carinae (setal rows) are well separated along their entire length in Deccanometrus, whereas the proximal half of the dorsal secondary carina, distal half of the subdigital carina and distal third of the digital carina are closely adjacent in Chersonesometrus and Gigantomentrus. The dorsal secondary and subdigital carinae of the male are absent or obsolete in Deccanometrus but present and entirely to predominantly granular or costate in Gigantometrus, all species of Chersonesometrus except C. bastawadei and C. fulvipes, and all species of Srilankametrus except S. indus and S. pococki. The chela manus digital and retromedian carinae are similarly developed (figs. 122, 125, 126, 129, 133, 134, 137, 138, 141, 143, 144) in Deccanometrus, whereas the retromedian carina is more pronounced than the digital carina in Chersonesometrus. The retromedian carina of the male is absent or obsolete in Deccanometrus but entirely to predominantly granular in Gigantometrus, Srilankametrus caesar, and S. serratus, and entirely to predominantly costate in Chersonesometrus, Srilankametrus couzijni, S. gravimanus, and S. yaleensis. The depression in the dorsal surface of the chela manus, proximal to the fixed finger of the adult male, is absent or obsolete (figs. 122B–D, 125, 133, 137, 141A, 143A) in Deccanometrus but present and distinct in Heterometrus. The chela manus dorsal surface is shallowly reticulate (figs. 122, 125, 126, 129, 141, 143, 144) in all except two species of Deccanometrus, D. obscurus and D. phipsoni, in which the surface is without reticulation (figs. 133, 134, 137, 138), as in Gigantometrus, Srilankametrus, most species of Chersonesometrus, and all except three species of Heterometrus, H. glaucus, H. laevigatus, and H. thorellii; the dorsal surface is finely to coarsely granular in all species of Deccanometrus except D. ubicki, in which the surface is smooth, as in Heterometrus, Javanimetrus, and two species of Sahyadrimetrus, S. kanarensis and S. tikaderi. The chela manus retrolateral intercarinal surfaces are granular (figs. 122, 125, 126, 129, 133, 134, 137, 138, 143) in all species of Deccanometrus except D. ubicki, in which the surfaces are smooth or nearly so (fig. 141), as in Heterometrus, Javanimetrus and some species of Sahyadrimetrus. The chela manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in Deccanometrus but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in Javanimetrus, four species of Chersonesometrus, C. beccaloniae, C. hendersoni, C. pelekomanus, and C. tristis, and two species of Srilankametrus, S. indus and S. pococki. The pro- and retrolateral surfaces of the tibiae of legs I and II each bear a row of two or three spiniform macrosetae (figs. 46–48) in Deccanometrus and scattered, setiform macrosetae, not arranged in a definite row, in Heterometrus. The pseudonychium (dactyl) of the telotarsi of legs I–IV is reduced and rounded (figs. 46–48) in Deccanometrus but prominent and acuminate in Heterometrus. Macroseta st on the retroventral surface of the basitarsus of leg I is spiniform (figs. 46–48) in Deccanometrus but usually setiform in Heterometrus, and sb on the retroventral surface of the basitarsus of leg III is setiform in Deccanometrus but spiniform in Javanimetrus and Sahyadrimetrus. The pectinal first proximal median lamella (scape) of the female is distinctly angular, ≥ 90° but < 180° (figs. 33–35) in Deccanometrus but straight or shallowly curved in Srilankametrus. The mesial surfaces of mesosomal tergites I–VI of the male are smooth (figs. 119, 123A, B, 130, 135A, B, 139A, B) in all except one species of Deccanometrus, D. xanthopus, in which the surfaces are granular (fig. 127C, D), as in Gigantometrus, some species of Chersonesometrus, and two species of Sahyadrimetrus, S. mathewi and S. scaber. The lengths of metasomal segments I and II are approximately equal to or less than their respective widths (figs. 64–66) in Deccanometrus but markedly greater than their respective widths in Gigantometrus. The ventrosubmedian and ventrolateral carinae are granular or costate-granular on metasomal segment IV only and costate on segments I–III (figs. 65, 66) in Deccanometrus but granular on segments I–IV, II–IV (costate on I), or III and IV (costate on I and II) in Gigantometrus. The ventral intercarinal surfaces of metasomal segment IV are smooth in Deccanometrus but sparsely granular in Javanimetrus. The dorsosubmedian carinae of metasomal segment V are vestigial (fig. 64) in Deccanometrus but partial in Gigantometrus and absent in Heterometrus. The dorsal intercarinal surface of segment V is smooth in Deccanometrus but granular in Gigantometrus, and some species of Chersonesometrus and Srilankametrus. The width of the telson vesicle is approximately equal to or less than the width of metasomal segment V in the female of Deccanometrus but greater than the width of segment V in the female of Gigantometrus and the vesicle is globose in Deccanometrus but elongate in Heterometrus, Javanimetrus, Sahyadrimetrus, and all except three species of Srilankametrus, S. indus, S. pococki, and S. serratus.

Etymology: The generic name refers to the Deccan Plateau, covering an area of approximately 422,000 km² between the Western Ghats and the Eastern Ghats of the Indian subcontinent, and traversing the central Indian states of Maharashtra, Karnataka, Telangana, and Andhra Pradesh. Along with many other taxa, most species of the genus are endemic to or associated with the Deccan Plateau.

Included Species: Deccanometrus, gen. nov., is hereby created to accommodate seven species, formerly assigned to subgenera Chersonesometrus, Heterometrus, and Srilankametrus of Heterometrus by various authors (Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000), one of which was formerly recognized at the rank of subspecies, recovered as a monophyletic group by phylogenetic analysis of morphological characters and DNA sequences from the nuclear and mitochondrial genomes (fig. 10): Deccanometrus bengalensis (C.L. Koch, 1841), comb. nov.; Deccanometrus latimanus (Pocock, 1894), comb. nov.; Deccanometrus liurus (Pocock, 1897), comb. nov.; Deccanometrus obscurus (Couzijn, 1981), comb. et stat. nov.; Deccanometrus phipsoni (Pocock, 1893), comb. nov.; Deccanometrus ubicki (Kovařík, 2004), comb. nov.; and Deccanometrus xanthopus (Pocock, 1897), comb. nov.

Distribution: Deccanometrus, gen. nov., is distributed across the Indian subcontinent (figs. 3, 117, 118, table 1), with records confirmed from China (in the Tibet Autonomous Region only), India (Andhra Pradesh, Assam, Chhattisgarh, Gujarat, Haryana, Himachal Pradesh, Karnataka, Jharkhand, Madhya Pradesh, Maharashtra, Odisha, Rajasthan, Tamil Nadu, Telangana, Uttarakhand, Uttar Pradesh, West Bengal, and the union territory of Puducherry), Nepal and Pakistan (in Khyber Pakhtoon Khwa and perhaps Azad Kashmir). The presence of this genus in Bangladesh, while likely, requires confirmation.

Ecology: The species of Deccanometrus for which data are available, occur in habitats ranging from evergreen forest, through deciduous woodland to semiarid savanna, at elevations of 300–1030 m above sea level. Based on morphology and available habitat data, all species of the genus appear to be pelophilous and fossorial, constructing burrows under stones or, in the case of D. xanthopus, in open ground. Some species, e.g., D. liurus and D. phipsoni, are markedly dorsoventrally compressed, suggesting a semilithophilous habit.

Conservation Status: Deccanometrus bengalensis and D. latimanus are harvested, dried, and consumed for medicinal purposes in India (Majumder and Dey, 2005) and Pakistan (Tahir and Prendini, 2014), respectively, and D. bengalensis is occasionally harvested for the commercial trade in exotic pets.

Buthus bengalensis C.L. Koch, 1841b: 3–5, pl. CCXC, fig. 696; 1850: 88; Thorell, 1876b: 220; Moritz and Fischer, 1980: 310; Couzijn, 1981: 142; Kovařík, 2004: 7, 9.

Palamnaeus spinifer: Pocock, 1892: 38–41 (misidentification, part); 1894c: 316 (misidentification, part).

Scorpio bengalensis: Pocock, 1892: 39; 1893: 310, 312; Kraepelin, 1894: 25, 31, 37, 46, 47, 49, 51–53, pl. I, fig. 30 (part); Pocock, 1894a: 72.

Heterometrus bengalensis: Kraepelin, 1899: 110, 114, 115 (part); Werner, 1902: 601; Kraepelin, 1913: 166 (part); Borelli, 1915: 463; Lampe, 1918: 198; Roewer, 1943: 228; Takashima, 1945: 93; Pérez Minnocci, 1974: 38; Brignoli, 1985: 415; Hjelle, 1990: 51, fig. 2.24; McCormick and Polis, 1990: 298; Kovařík, 2004: 1, 7–9 (part), 44, 51–53, tables 2, 3, fig. 10; 2009: 34–36, 48, 49, 74, 99, tables 1, 2, figs. 8–13, 180, 181; Warburg, 2012a: 85.

Palamnaeus bengalensis: Pocock, 1900a: 85, 94–96; Kanungo, 1954: 536–539, figs. 1, 2; Srivastava, 1955: 85–91, text-figs. 1–5; Sinha, 1966: 221, 222, table; Srivastava and Kanungo, 1966: 629–632, tables 1–3; Sinha and Kanungo, 1967: 386–390, figs. 1–3; Kovařík, 2004: 44; Warburg, 2012a: 82, 85, table 1.

Palamnaeus phipsoni: Pocock, 1900a: 85, 94–96 (misidentification, part).

Heterometrus (Scorpio) bengalensis: Kraepelin, 1901: 271.

Heterometrus longimanus: Kraepelin, 1913: 166 (misidentification, part); Kopstein, 1921: 129 (misidentification, part).

Heterometrus longimanus longimanus: Giltay, 1935: 2 (part); Takashima, 1945: 90 (part).

Heterometrus (Heterometrus) bengalensis: Couzijn, 1981: 38, 73, 75–77, 80, 85, 88, 89, 117–120, 167, 173, 174, 177, 178, 183, 184, 189–191, table 7, figs. 17b, 18, 19, 21, 62, 63, 71 (part); Tikader and Bastawade, 1983: 522, 538–544 (part), figs. 1445–1457; Biswas, 1984: 320, 321; Kovařík, 1998: 137 (part); Fet, 2000: 432 (part); Majumder and Dey, 2005: 13, 14.

Heterometrus (Chersonesometrus) fastigiosus Couzijn, 1981: 22, 41, 87, 120, 140–142, 169, 170, 192, table 7, figs. 4d, 42, 57 (synonymized by Kovařík, 2004: 1, 7, 9, 44); Tikader and Bastawade, 1983: 574, 587–593, figs. 1553–1565; Kovařík, 1998: 136; Fet, 2000: 432, 438, 439; Kovařík, 2002: 3, 17; 2004: 1, 9, 44.

Heterometrus (Chersonesometrus) phipsoni: Couzijn, 1981: 153, 155, 157–159, 169, 180 (misidentification, part); Fet, 2000: 442 (part).

Heterometrus (Chersonesometrus) phipsoni phipsoni: Couzijn, 1981: 43, 87, 149–151, 153, 170, 192, table 7 (misidentification, part).

Heterometrus cf. (Chersonesometrus) phipsoni: Kovařík, 1997: 183 (misidentification).

Heterometrus fastigiosus: Kovařík, 2004: 7, 9; Zhu and Yang, 2007: 95.

Heterometrus nepalensis Kovařík, 2004: 1, 7, 29, 31, 32, 51, 52, tables 1, 2, fig. 22; 2009: 35, 42, 48, 49, 79, 100, table 1, figs. 47–49, 205; syn. nov.

Heterometrus phipsoni: Kovařík, 2004: 1, 4, 29, 34, 44, 49, 51, 52, tables 2, 3 (part); 2009: 35, 36, 42, 43, 48, tables 1, 2 (part).

Heterometrus tibetanus Lourenço et al., 2005: 1, 5, 10–14, table 1, figs. 18–35; Zhu and Yang, 2007: 95; Kovařík, 2009: 35, 45, table 1; Di et al., 2013: 52, 86, 88, 94, 95, fig. 138; syn. nov.

Heterometrus phipsoni: Mirza and Sanap, 2009: 488 (part); Aswathi and Sureshan, 2017: 9847 (part).

Type Material: CHINA: Tibet Autonomous Region: Ngari Prefecture: Heterometrus tibetanus: Holotype ♂ (MNHN RS 8608) [examined], 2 ♂ paratypes (MNHN) [1 ♂ paratype (MNHN RS 8726) examined], Tibet, south region of Pulan [Purang/Burang Xian, 30°15′N 81°10′E], low valley of Kongque He River, near border with Nepal, vii.1931. INDIA: Assam: Heterometrus (Chersonesometrus) fastigiosus: Holotype ♂, paratypes: 1 ♀, 1 subad. ♂ (SMF II/8886/228), Assam [examined]. West Bengal: Buthus bengalensis: Lectotype [= Heterometrus (Chersonesometrus) fastigiosus paratype]: 1 ♂(ZMB 70/2), paralectotypes [= H. (C.) fastigiosus paratypes]: 1 ♂ (ZMB 70/1), 3 ♂ (ZMB 70/4, 5, 7), 1 ♀ (ZMB 70/3), 1 subad. ♀ (ZMB 70/6), Bengal, Lam [examined]. NEPAL: Heterometrus nepalensis: Holotype ♀ [not ♂] (FKPC), Royal Chitwan National Park, Island Jungle Resort [27°38′N 84°10′E], Nepal, vi.1993, Csorba.

Buthus bengalensis was based on seven syntypes deposited at the ZMB (C.L. Koch, 1841b). According to Couzijn (1981: 120, 142) these syntypes were destroyed during World War II (unlikely as the arachnid collection at ZMB was not destroyed in the war, unlike those of some other German museums). Couzijn (1981: 142) examined two of the specimens, but speculated that they did not match Koch's (1841) original description: “they do not fit the description of B. bengalensis by the absence of external keels on the pedipalp hands and the different internal denticulation on the fourth pedipalp segment.” Couzijn (1981) designated the two syntypes of B. bengalensis as paratypes of a new species, Heterometrus (Chersonesometrus) fastigiosus, and designated a neotype, deposited at MNHN, for H. bengalensis, which was in fact conspecific with another species, Heterometrus thorellii.

Kovařík (2004: 9) considered Couzijn's (1981) taxonomic decision “entirely invalid and incomprehensible.” As all seven syntypes of B. bengalensis are still extant at the ZMB, Kovařík (2004) designated one as lectotype, invalidating Couzijn's (1981) designation of the neotype in accordance with ICZN Article 75.8.

Diagnosis: Deccanometrus bengalensis may be separated from other species of Deccanometrus as follows. The carapace is slightly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 19A, B), in D. bengalensis but vaulted, the lateral surfaces sloping moderately, in D. latimanus and D. xanthopus. The median notch in the carapace anterior margin is deeply excavated in D. bengalensis but shallow in D. liurus. The superciliary carinae are higher than the median ocelli in D. bengalensis, but lower than the ocelli in all other species except D. latimanus, D. liurus, and D. ubicki. The interocular suture is present in D. bengalensis but absent in D. liurus. The carapace interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the male (fig. 19A), and the frontal lobes and medial region are granular, with smooth areas, in the female (fig. 19B) of D. bengalensis, whereas the interocular surface is granular along the median longitudinal and anterior bifurcated sulci only in the female of D. latimanus, D. liurus, D. obscurus, and D. phipsoni, and the frontal lobes and medial region are granular, with smooth areas, in the male of D. ubicki or partially to entirely granular in the male of D. xanthopus. The carapace posterolateral surfaces of the female are granular in D. bengalensis but smooth or nearly so in D. obscurus, D. phipsoni, D. ubicki, and D. xanthopus. The cheliceral movable finger prodistal (DI) and retrodistal (DE) teeth are aligned longitudinally and not opposable in D. bengalensis but opposable, i.e., forming a bicusp, in D. liurus and D. ubicki. The pedipalps of the adult male are long, with femur length: posterior carapace width ratio (FL:PCW) ≥ 0.80 and femur length: carapace length ratio (FL:CL) ≥ 0.76 (fig. 119) in D. bengalensis but short, with FL:PCW < 0.77, FL:CL < 0.74 in D. latimanus. The pedipalp femur prodorsal carina is present and distinct in D. bengalensis but absent or obsolete in D. liurus and D. xanthopus. The pedipalp patella dorsomedian carina of the female is entirely to predominantly granular in D. bengalensis but absent or obsolete in D. latimanus, D. liurus, and D. ubicki. The patella retrodorsal carina of the female is absent or obsolete in D. bengalensis but as strongly developed as or more strongly developed than the retromedian carinae in D. obscurus and D. phipsoni. The retromedian carinae of the female are granular in D. bengalensis but absent or obsolete in D. latimanus, D. liurus, D. ubicki, and D. xanthopus. The patella dorsal, retrodorsal, and retroventral intercarinal surfaces of the female are smooth or nearly so in D. bengalensis but granular in D. obscurus and D. phipsoni. The pedipalp chela is infuscate and similar in color to the femur and patella in D. bengalensis but immaculate and paler than the femur and patella in D. latimanus, D. liurus, and D. xanthopus. The chela of the adult male is moderately to densely setose (fig. 122B–D) in D. bengalensis but sparsely setose in D. xanthopus. The chela manus promedian carina of the female is granular in D. bengalensis but absent or obsolete in D. latimanus and D. xanthopus. The maximum distance between the dorsomedian and dorsal secondary carinae (DMC–DSC) of the chela manus is less than the maximum distance between the dorsal secondary and digital carinae (DSC–DC) in the male and female of D. bengalensis whereas the DMC–DSC is similar to the DSC–DC in the male and female of D. xanthopus, and the female of D. obscurus, and greater than the DSC–DC in the male and female of D. latimanus. The chela manus dorsal intercarinal surface is finely to coarsely granular and shallowly reticulate (fig. 122) in D. bengalensis, smooth and shallowly reticulate in D. ubicki, and finely to coarsely granular, without reticulation in D. obscurus and D. phipsoni. The manus retrolateral intercarinal surfaces are granular in D. bengalensis but smooth or nearly so in D. ubicki. The legs are very dark or heavily infuscate (figs. 119, 120) in D. bengalensis but pale or very lightly infuscate in D. latimanus, D. liurus, D. ubicki, and D. xanthopus. Macroseta st on the retroventral surfaces of the basitarsi is spiniform on legs I and II and setiform on III (fig. 46A–D) in D. bengalensis, spiniform on legs I–III in D. obscurus, D. phipsoni, and D. ubicki, and setiform on legs I–III in D. latimanus, D. liurus, and D. xanthopus; sb on the retroventral surfaces of the basitarsi is spiniform on legs I and II in D. bengalensis, setiform on legs I and II in D. latimanus, and setiform on leg I, and setiform or spiniform on leg II in D. xanthopus; st on the proventral surface of the basitarsus of leg III is spiniform in D. bengalensis but setiform in D. obscurus and D. phipsoni; sb on the retrolateral surface of the basitarsus of leg III is spiniform in D. bengalensis but setiform in D. latimanus. The laterodistal lobes on the telotarsi of legs I–IV each bear only two spiniform macrosetae in D. bengalensis but three or more in D. latimanus. The angle of the pectinal first proximal median lamella (scape) is > 90° but < 180° in the male and female (fig. 33A, B) of D. bengalensis, and approximately 90° in the male of D. latimanus, D. liurus, and D. xanthopus, and the male and female of D. ubicki. The mesial surfaces of mesosomal tergites I–VI are smooth in the male and the lateral surfaces granular in the female of D. bengalensis, whereas the mesial surfaces are granular in the male of D. xanthopus, and the lateral surfaces smooth in the female of D. latimanus, D. liurus, D. ubicki, and D. xanthopus. The ventral surface of mesosomal sternite VII bears a pair of weakly developed ventrolateral carinae in D. bengalensis but two pairs of moderately to strongly developed ventrosubmedian and ventrolateral carinae in D. latimanus, D. liurus, D. ubicki, and D. xanthopus. The dorsosubmedian carinae of metasomal segments I–IV are granular or costate-granular (fig. 64A) in D. bengalensis, costate in D. latimanus, and obsolete in D. liurus. The ventrosubmedian and ventrolateral carinae of metasomal segments I–IV are more strongly developed on segments III and IV than on I and II (fig. 66A) in D. bengalensis but more strongly developed on segments I and II than on III and IV in D. latimanus, D. liurus, D. ubicki, and D. xanthopus. The lateral carinae are partial on metasomal segments I and V (fig. 65A) in D. bengalensis but vestigial on I in D. liurus and D. ubicki, and absent or obsolete on V in D. latimanus, D. liurus, and D. ubicki. The dorsolateral carinae of metasomal segment V are strong and continuous in D. bengalensis but weak and discontinuous to absent in D. latimanus, D. liurus, and D. xanthopus. The width of the telson vesicle is approximately equal to or less than the width of metasomal segment V in the male (figs. 64A, 66A) of D. bengalensis but greater than the width of segment V in the male of D. latimanus, D. liurus, D. ubicki, and D. xanthopus.

Distribution: This species has been recorded in China (in the Tibet Autonomous Region only), India (in the states of Assam, Haryana, Himachal Pradesh, Jharkhand, Madhya Pradesh, Odisha, Rajasthan, Uttar Pradesh, Uttarakhand, and West Bengal) and Nepal (fig. 117, table 1). Its presence in Bangladesh, based on a record from the Ganges Delta (Kraepelin, 1913), while likely, requires confirmation. Records of this species and its junior synonym, H. fastigiosus, from Maharashtra, India (Tikader and Bastawade, 1983; Bastawade, 2002) are referable to D. phipsoni.

Ecology: Surprisingly little is known about the ecology of this widespread species. The known records occur in areas of deciduous forest on the plains south of the Himalayas and valleys in the Himalayan foothills. Specimens from Rajaji National Park in Uttar Pradesh, India, were found in a deciduous forest with Sal, Shorea robusta C.F. Gaertner, 1805 (Biswas, 1984). The burrowing biology of this species was described by Kanungo (1954). The burrows are constructed in open ground on shaded slopes of low, vegetated ridges (“bunds”). The crescentic entrance, 5.1–7.6 cm × 1.3–3.8 cm, leads via a sinuous passage, almost circular in cross-section, to a terminal chamber, ca. 30 cm below the surface. As with other species of the genus, the habitus is consistent with the fossorial, pelophilous ecomorphotype (Prendini, 2001b). Deccanometrus bengalensis is sympatric with the buthid Hottentotta tamulus, and the hormurid Liocheles nigripes (Kanungo, 1954).

Conservation Status: Deccanometrus bengalensis is harvested for medicinal purposes (Majumder and Dey, 2005) and occasionally for the exotic pet trade.

Remarks: This species, previously placed in the nominal subgenus of Heterometrus by various authors (e.g., Couzijn, 1981; Tikader and Bastawade, 1983; Fet, 2000), is newly accommodated in Deccanometrus, gen. nov., based on a phylogenetic analysis (fig. 10).

Couzijn (1981: 117) examined the syntypes of Palamnaeus costimanus var. γ glaucus, allegedly from “Bengal,” and synonymized the taxon with Heterometrus (H.) bengalensis. Fet (2000: 432) and Kovařík (2004: 7) followed Couzijn (1981). Reexamination of the types during the present investigation demonstrated that they are neither conspecific with one another nor with D. bengalensis, however.

Kovařík (2004) examined the holotype and paratypes of H. (C.) fastigiosus, concluded they were conspecific with H. bengalensis, and synonymized H. (C.) fastigiosus with the latter. Based on Koch's (1841b) original description of Buthus bengalensis and examination of the types of B. bengalensis and H. (C.) fastigiosus, Kovařík's (2004) conclusions, that Couzijn (1981) erroneously synonymized H. thorellii with H. bengalensis and redescribed B. bengalensis as H. (C.) fastigiosus, are upheld. Koch's (1841b) original illustration of the male and the pedipalp chela of the female of B. bengalensis reveals that the type specimens do not match H. thorellii, i.e., Couzijn's (1981) concept of D. bengalensis, because the chela manus is granular and relatively shorter than that of H. thorellii, and a prominent spiniform granule is absent from the proventral surface of the patella. Therefore, the following synonym is confirmed: Heterometrus (Chersonesometrus) fastigiosus Couzijn, 1981 = Deccanometrus bengalensis (C.L. Koch, 1841), comb. nov., synonymized by Kovařík (2004).

Heterometrus nepalensis is known only from the holotype, a dry specimen that is allegedly male, but judging from Kovařík's (2009: 79) figures 47–49 closely matches the female of D. bengalensis. Although Kovařík (2004: 32) cites a close relationship to Srilankametrus indus (as H. indus), the following putatively diagnostic characters are a precise match for D. bengalensis (Kovařík, 2004: 9, 32), which occurs in close geographical proximity to the type locality of H. nepalensis: carapace smooth, with granules only at margins; pedipalp patella without pronounced internal tubercle; chela hirsute, “lobiform,” its adult length to width ratio 2.2, as in the female of D. bengalensis, for which Kovařík (2004: 9) gives a ratio of 2.2, and Tikader and Bastawade (1983: 541) describe the chela manus of D. bengalensis as “lobate”; pectinal tooth count 14–15 compared with 14–17 in both sexes of D. bengalensis (Tikader and Bastawade, 1983; Kovařík, 2004); “spine formula” of legs III and IV, 4/5 compared with 4–6/4–6 in D. bengalensis, based on data in Tikader and Bastawade (1983) and Kovařík (2004); telson bulbous, vesicle longer than aculeus. Many specimens of D. bengalensis (e.g., figs. 119, 120) are “uniformly reddish black, only telson lighter colored” (Kovařík, 2004: 32) and the length of the holotype of H. nepalensis (83 mm) falls within the range of 78–115 mm for D. bengalensis based on data from Couzijn (1981), Tikader and Bastawade (1983) and Kovařík (2004). The granular pedipalp chela manus described for H. nepalensis is also observed in D. bengalensis and was described as “covered with obsolete granules” by Tikader and Bastawade (1983: 542) and as “sparsely tuberculate, uneven” by Kovařík (2004: 9). Finally, the suggestion that sexual dimorphism in the proportions of the pedipalps is “probably not noticeable” (Kovařík, 2004: 32) rests on the assumption that the holotype is an adult male, which has not been demonstrated.

Heterometrus tibetanus was described from three males (Lourenço et al., 2005). The diagnosis of H. tibetanus compares it with H. nepalensis and lists the following differences: much paler coloration in H. tibetanus than H. nepalensis which is reddish black; pedipalps and chela strongly elongated in H. tibetanus but hirsute “lobiform” chela in H. nepalensis; second metasomal segment longer than wide in H. tibetanus but wider than long in H. nepalensis; telson aculeus same length as vesicle in H. tibetanus but longer than aculeus in H. nepalensis. Setting aside the low sample sizes for both species (n = 1 for H. nepalensis; n = 3 for H. tibetanus), it is apparent that several putative diagnostic differences, e.g., the differences in the shape of the pedipalps, metasoma, and telson, reflect sexual dimorphism. Although the diagnoses of H. nepalensis and H. tibetanus do not compare either species to D. bengalensis, the figures and diagnostic characters closely match the latter, with which they are clearly conspecific. Kovařík (2009: 45) suggests further that H. nepalensis may be a senior synonym of H. tibetanus. As H. tibetanus is undoubtedly a male of D. bengalensis, Kovařík's (2009) observation reinforces the conclusion that H. nepalensis is a female, conspecific with D. bengalensis.

The total length of H. tibetanus (85–88 mm) is close to the length of the holotype of H. nepalensis (83 mm) and falls within the range of D. bengalensis, which is 78–115 mm based on data from Tikader and Bastawade (1983) and Kovařík (2004). The pectinal tooth count of H. tibetanus (15–16 teeth) falls within the range of D. bengalensis, which has 14–17 teeth in both sexes (Tikader and Bastawade, 1983; Kovařík, 2004). The character “pedipalps, especially the chela strongly elongated and rather slender” (Lourenço et al., 2005: 11) exactly matches the male of D. bengalensis as do the characters “chela manus with strong spinoid granules on the dorsointernal aspect” and “second metasomal segment longer than wide (7.1/5.9).” The coloration “reddish-yellow to reddish-brown ... legs, chelicerae and telson paler than the body” is commonly observed in D. bengalensis which varies from reddish yellow to reddish black across its distribution; Tikader and Bastawade (1983: 539) described the color as “entirely brown with reddish tint.” The counts of proventral and retroventral spiniform macrosetae on the telotarsi, 3/4 on legs I and II in H. tibetanus (Lourenço et al., 2005), and 4/5 on legs III and IV in H. nepalensis (Kovařík, 2004) fall within the ranges of corresponding counts for D. bengalensis, i.e., 3/5 on I and II, and 4–6/4–6 on III and IV (Tikader and Bastawade, 1983; Kovařík, 2004). Based on the evidence, there is little doubt that H. nepalensis and H. tibetanus are conspecific with D. bengalensis and represent the sexually dimorphic female and male of that species, respectively. This conclusion was reinforced by examination of the type material of D. bengalensis and H. tibetanus. The following new synonyms are proposed accordingly: Heterometrus nepalensis Kovařík, 2004 = Deccanometrus bengalensis (C.L. Koch, 1841), syn. nov.; Heterometrus tibetanus Lourenço et al., 2005 = D. bengalensis, syn. nov.

Material Examined: INDIA: Dr. C.J. Marinkelle, ex Stahnke coll., 1 ♀ (CAS 9071132 [ASU 61-11/65-0121]); northern India, 22.xi.1873, M.M. Carleton, 1 ♀, 1 subad. ♀ (MCZ 88170). Assam: Coremia, Assam, O.P. Cambridge, 1 ♂ (BMNH 1896.10… [illegible]). Haryana: Ambala Distr.: Amballa [Ambala, 30°23′N 76°47′E], plains 70 mi. SW, M.M. Carleton, 1874, 1 ♂, 1 ♀, 1 subad. ♂, subad. ♀ (MCZ 88169). Karnal Distr.: Nokheri [Nilokheri, 29°50′N 76°56′E], Faunthorpe-Vernay Expedition, 29.iv.1923, 1 ♀(AMNH [24381]). Himachal Pradesh: Kullu Distr.: Koolloo [Kullu, 31°57′N 77°07′E], 1876, 1 ♀ (MCZ 88164). Jharkhand: Chota Nagpur [Chota Nagpur Plateau, ca. 23°N 85°E], D. Winffin, 1 ♂ (BMNH 1896.9.26-92 [BNHS 662]). Dumka Distr.: KathiKund (Bihar) [Kathikund, 24°22′N 87°25′E], viii.1967, R. Heim, voisinage immediate de la fôret (Shorea), 1 ♂(MNHN RS 5325). Madhya Pradesh: Gwalior Distr.: Gwalior [26°13′N 78°11′E], 1 ♂ (BMNH 1896.12.15-19 [BNHS 185]). Satna Distr.: Satna [24°36′N 80°50′E], central India, R. Wroughton, 1 ♂ [BNHS 576], 1 ♀, 1 subad. ♀ [BNHS 814] (BMNH 1896.8.15.4). Odisha: Majurbanj Distr.: Rangamatia Village, near Baripada, 21°55′N 86°45′E, T. Kroes, 1 ♂, 1 ♀ (AMNH), 3 juv. (AMCC [LP 14238]). Uttar Pradesh: Allahabad Distr.: Allahabad [25°26′N 81°51′E], E.W. Oates, 2 subad. ♂, 2 subad. ♀ (BMNH 1897.7.5.10.218-221). Unnao Distr.: Shakpur [26°32′N 80°28′E], near Unao, Punjab, H.W. Waite, 15.vi.1912, 1 ♂ (BMNH 1913.1.6.3). Uttarakhand: Dehradun Distr.: Dehra Dun [30°19′N 78°02′E], 1 ♂ (BMNH 1897.9.17.5 [BNHS 1236]), F. Gleadow, 1 ♂ (BMNH 1896.10.20.70 [BNHS 843]), Gamble, 2 ♂, 1 ♀ (BMNH 1896.2.2.1-2). Pauri Garhwal Distr.: Munala [Mundala, 29°48′N 78°31′E], A.N. Cassia, 1 juv. ♂, 1 juv. ♀ (BMNH 1896.9.26.85,86 [BNHS 764]). West Bengal: Bengal, 1 ♂, 1 ♀ (BMNH 1860.15); West Bengal, 13.iii.1968, received from Zafer's Export Centre, ex Stahnke coll., 8 ♂, 5 ♀, 1 subad. ♂, 1 subad. ♀(CAS [ASU 68-0178, 180, 182, 184, 186, 187, 191, 192, 194, 224, 226, 245, 251–253]), 13.iii.1968, Zafer's Export Centre, ex Stahnke coll., 2 ♀ (CAS 9071130 [ASU 68-0227], 9071131 [ASU 68-0223]). Kolkata Distr.: Calcutta [Kolkata, 22°34′N 88°22′E], Barabank?, B.V. College, rec. P.K. Sina, 17.viii.1965, ex Stahnke coll., 1 ♀(CAS 9071134 [ASU 60-271/75-0321]). Paschim Medi-nipur Distr.: Kharqpur [Kharagpur, 22°20′N 87°20′E], 30 m, 20.vi.1960, L. Murmu, in yard, 1 ♂ (CAS 9071140), 17.ii.1961, C.L. Kau, engine room by tin, 1 ♀(CAS 9071133). Purba Bardhaman Distr.: Talpata village [23°11′N 88°07′E], near Saktigarh [23°12′N 87°58′E], ca. 100 km W of Calcutta, xi.1973, Prof. Lahiri, 1 ♀, 2 subad. ♂ (MNHN).