Srilankametrus caesar (C.L. Koch, 1841), 2020

Prendini, Lorenzo & Loria, Stephanie F., 2020, Systematic Revision Of The Asian Forest Scorpions (Heterometrinae Simon, 1879), Revised Suprageneric Classification Of Scorpionidae Latreille, 1802, And Revalidation Of Rugodentidae Bastawade Et Al., 2005, Bulletin of the American Museum of Natural History 2020 (442), pp. 1-480 : 392-400

Srilankametrus caesar (C.L. Koch, 1841) ,

comb. nov. et stat. rev.

Figures 10 , 27A, B , 41A, B , 56A–D , 76–78 , 199 , 225–229, table 1

Buthus caesar C.L. Koch, 1841b: 6–9 , pl. CCXCI, fig. 697; 1850: 87; Simon, 1872b: 101; Thorell, 1876b: 202; Moritz and Fischer, 1980: 311; Couzijn, 1981: 123.

Heterometrus afer: Simon, 1872b: 53 , 59, 98–101 (misidentification, part).

Pandinus caesar: Thorell, 1876b: 202 , 205; Karsch, 1884: 68.

Scorpio africanus: Pocock, 1893: 311 (misidentification).

Scorpio caesar: Pocock, 1893: 310 , 311 (part).

Scorpio ceylonicus: Kraepelin, 1894: 31 , 46–51, 54, 55, 57 (misidentification, part).

Heterometrus caesar: Kraepelin, 1899: 109 , 114 (part); Roewer, 1943: 229 (part); Takashima, 1945: 92; Bücherl, 1964: 59; Brignoli, 1985: 415.

Palamnaeus caesar: Pocock, 1900a: 86 , 97 (part).

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

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

Pandinus gravimanus: Simon, 1905: 161 (misidentification, part).

Heterometrus (Chersonesometrus) fulvipes: Couzijn, 1981: 40 , 72, 88, 123, 133–136, 139, 140, 142, 192, table 7, fig. 16b (misidentification, part); Fet, 2000: 439 (part).

Heterometrus (Chersonesometrus) granulomanus Couzijn, 1981: 19 , 22, 43, 87, 133, 141–143, 170, 192, table 7, figs. 2g, 4c, 43, 58; Tikader and Bastawade, 1983: 573, 574, 577–582 (misidentification, part), figs. 1525–1538; Kovařík, 1998: 136; Fet, 2000: 439 (part); Bastawade, 2004: 286, 291, 292 (part); Bastawade et al., 2004: 48, 54, 58, figs. 67, 68; Kovařík, 2004: 1, 29 (part); Thulsi Rao et al., 2005: 3, 9 (misidentification, part); Sureshan et al., 2007a: 2906–2908 (misidentification, part); Bastawade, 2008: 133, 136, 137 (part); Indra, 2009: 141; syn. nov.

Heterometrus (Srilankametrus) indus: Couzijn, 1981: 94 , 121, 123 (misidentification, part); Kovařík, 1998: 137 (part); Fet, 2000: 446, 447 (part).

Heterometrus (Srilankametrus) indus indus: Couzijn, 1981: 82 , 121–124, 129, 191 (misidentification, part).

Heterometrus indus: Kovařík, 2004: 2 , 17, 20, 21, 32, 52 (misidentification, part); 2009: 38, 48 (misidentification, part).

Heterometrus latimanus: Kovařík, 2004: 21 , 51, table 2 (misidentification, part); 2009: 35, 39, 47, 49, 73, 100 (misidentification, part), table 1, figs. 3, 4, 198 (misidentification).

Heterometrus madraspatensis: Kovařík, 2004: 1 , 2, 27, 29, 51, 52, table 2 (misidentification,

part); 2009: 35, 41, 48, table 1 (misidentification, part).

TYPE MATERIAL: INDIA: Buthus caesar : Lectotype [here designated]: 1 subad. ♀ (ZMB 63), “India Orientalis” [East India] [examined]. Puducherry: Puducherry Distr.: Heterometrus (Chersonesometrus) granulomanus : Holotype ♂, paratype ♀ (MNHN RS 0087), Pondichéry [Pondicherry /Puduchcheri/ Puducherry, 11°59′N 79°49′E], viii.1901, M. Maindron [examined].

Several syntypes were mentioned by Koch (1841b: 9), only one of which remains in the ZMB. As noted by Tahir and Prendini (2014: 6, 20), this single specimen precisely matches two females in a similar state of preservation, suggesting they are very old, and with the same locality data, i.e., “ India,” misidentified as Heterometrus latimanus by Kovařík (2004: 21, 51, table 2; 2009: 34, 35, 39, 47, 49, 73, 100, table 1, figs. 3, 4, 198).

In the description of Heterometrus (Chersonesometrus) granulomanus, Couzijn (1981: 142) specifically mentioned a male holotype and a female allotype (i.e., a designated paratype that is the opposite sex to the holotype) from the same locality but did not refer to other material examined as paratypes. Therefore, the other specimens listed in Couzijn’s (1981) description are not paratypes of H. (C.) granulomanus contrary to Fet (2000: 439) and Kovařík (2004: 27).

DIAGNOSIS: Srilankametrus caesar may be separated from other species of Srilankametrus as follows. The carapace is vaulted, the lateral surfaces sloping steeply (fig. 27A, B), in S. caesar but slightly to markedly dorsoventrally compressed, the lateral surfaces sloping moderately to gently, in S. indus , S. pococki , and S. serratus . The interocular and posterior sutures are present, the posterior sutures extending past the median ocular tubercle and connected anteriorly to the posterior bifurcations of the interocular suture, in S. caesar whereas the sutures are absent in S. indus and S. pococki . The frontal lobes and medial region of the carapace interocular surface are granular with smooth areas (fig. 27A, B) in S. caesar , whereas the interocular surface is entirely smooth in the other species. The carapace anterolateral and mediolateral surfaces are granular in S. caesar but smooth or nearly so in S. indus . The carapace posterolateral surfaces of the female are granular in S. caesar but smooth or nearly so in S. gravimanus and S. yaleensis . The pedipalp patella dorsomedian carina of the female is entirely to predominantly costate in S. caesar but absent or obsolete in all other species. The patella retrodorsal carina of the female is absent or obsolete in S. caesar but as strongly developed as or more strongly developed than the retromedian carinae in S. gravimanus , S. serratus , and S. yaleensis . The retromedian carinae of the female are granular in S. caesar but absent or obsolete in S. gravimanus and S. yaleensis . The pedipalp chela of the adult male (fig. 228) is sparsely setose in S. caesar but moderately to densely setose in S. indus and S. pococki . The chela manus dorsal surface (between the dorsomedian and digital carinae) is curved and slightly to markedly convex in S. caesar but flat in S. pococki . The proximal margin (lobe) of the dorsal surface is moderately curved and proximal to the proximal margin of the retrolateral surface but distal to the proximal margin of the condyle (articulation with patella) in S. caesar , whereas it is markedly curved and proximal to the proximal margin of the condyle in S. couzijni . The dorsomedian carina is pronounced and costate on the chela fixed finger and distally on the manus in S. caesar , pronounced and costate on the fixed finger and distal three-quarters of the manus in S. yaleensis , and obsolete on the fixed finger and manus in S. indus , S. pococki , and S. serratus . 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 (fig. 228) of S. caesar but similar to the DSC–DC in the male of S. gravimanus . The dorsal secondary and subdigital carinae of the male (fig. 228) are entirely to predominantly granular in S. caesar but entirely to predominantly costate in S. gravimanus and S. yaleensis and absent or obsolete in S. indus and S. pococki . The digital carina is entirely to predominantly granular (figs. 228, 229) in S. caesar , entirely to predominantly costate in S. couzijni , S. gravimanus , and S. yaleensis , and absent or obsolete in S. indus and S. pococki . The retromedian carina of the male is entirely to predominantly granular in S. caesar , entirely to predominantly costate in S. couzijni , S. gravimanus , and S. yaleensis , and absent or obsolete in S. indus and S. pococki . The manus ventral surface is flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in S. caesar , but angled, with the axis of the retroventral carina ventral to the axis of the ventromedian carina in S. indus and S. pococki . Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is spiniform (fig. 56A– D) in S. caesar but setiform in all other species except S. gravimanus and S. serratus . The lateral surfaces of mesosomal tergites I–VI are granular in S. caesar but smooth in S. indus , S. pococki , and S. yaleensis . The dorsosubmedian carinae are granular or costate-granular on metasomal segments I–IV (fig. 76A) in S. caesar , costate on I–IV in S. indus , and costate on I and II or I–III and granular or costate-granular on III and IV or IV, in S. yaleensis . The ventral intercarinal surfaces of metasomal segment IV are granular in the male and female (fig. 78A) of S. caesar , but smooth in the male and female of S. indus , S. pococki , S. serratus , and S. yaleensis , and the female of S. gravimanus . The dorsolateral carinae of metasomal segment V are distinct and continuous (fig. 77A) in S. caesar but obsolete and discontinuous to absent in S. indus and S. pococki . The dorsal intercarinal surface of metasomal segment V is granular in the male and female (fig. 76A) of S. caesar but smooth in the male and female of S. indus and S. pococki , and the female of S. serratus and S. yaleensis . The telson is blackish, as dark as metasomal segment V, in S. caesar but dark reddish brown, paler than segment V, in S. serratus and S. yaleensis . The telson vesicle is elongate in S. caesar but globose in S. indus , S. pococki , and S. serratus .

DESCRIPTION: In addition to the original description of Buthus caesar , the holotype and paratype of H. (C.) granulomanus were described and illustrated by Couzijn (1981) and again by Tikader and Bastawade (1983).

DISTRIBUTION: This species is endemic to India. The only material with locality data is labelled as originating from the union territory of Puducherry (fig. 199, table 1). If accurate, this species would appear to be allopatric with all other species of Srilankametrus . Records from Ceylon ( Sri Lanka) attributed to this species ( Pocock, 1894a; Kraepelin, 1899) refer to S. serratus .

ECOLOGY: The only known locality record occurs in deciduous forest at low elevation. The habitus is consistent with the fossorial, pelophilous ecomorphotype ( Prendini, 2001b). The spe- cies probably excavates deep burrows in sandy-loam soil, in open ground.

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

REMARKS: Buthus caesar was described from an unspecified number of syntypes from “ India Orientalis” ( Koch, 1841b). The identity of this species has remained enigmatic since its description. According to Fet (2000: 445), Kraepelin (1894: 46) first synonymized B. caesar with S. indus whereas, according to Kovařík (2004, 2009), Simon (1872b: 99) was the first to do so. However, Simon (1872b: 99) synonymized B. caesar with specimens misidentified as Heterometrus afer (nec Scorpio afer Linnaeus, 1758 ) that were conspecific with Heterometrus (Chersonesometrus) scaber , as discussed by Couzijn (1981: 145, 146). Kraepelin (1894: 46) synonymized B. caesar with Scorpio ceylonicus , which was synonymized with S. indus , notwithstanding that S. indus was the older name. Therefore, it appears Couzijn (1981: 123, 134) was the first to synonymize B. caesar with H. indus despite stating that a syntype of B. caesar in the ZMB “evidently belongs to H. fulvipes (Koch) and not to H. indus (de Geer) , nor to S. caesar (Koch) ,” an opinion repeated by Fet (2000: 439, 447). Other authors confused B. caesar with S. serratus ( Pocock, 1894a, 1900a; Kraepelin, 1899; Roewer, 1943; Takashima, 1945).

Despite immaturity, poor condition, and the absence of precise locality data, the sole remaining type (here designated as the lectotype) of Buthus caesar is clearly conspecific with type and nontype material of H. (C.) granulomanus (MNHN RS 0079, 0087, 3246) and several specimens, misidentified as H. (C.) fulvipes by Couzijn (1981: 134), with the same collection data (MNHN RS 6168) as well as material, in some cases identified as Palamnaeus caesar , from other collections (BMNH, ZMB 2282, 2283, 7244), based on the shape and macrosculpture of the carapace, pedipalp chela, metasoma, and telson. These specimens also match the original description and illustration of B. caesar in coloration, being completely black with pale tarsi (unlike Chersonesometrus fulvipes , in which the legs and telson are pale yellow); surface macrosculpture, differing from S. indus (and other species of Srilankametrus ) in the granular frontal lobes of the carapace (“vorn an den zwei vorderrandslappen niedergedruckt und flach, auf dieser flache unordentlich zerstreute ungleichgrosse kleine kornchen”) and from Sahyadrimetrus scaber in the granular pedipalp chela manus; and the shape of the pedipalp chela with its bent, flat and broad fixed finger (“die finger etwas gebogen, sehr flach, breit”) ( Koch, 1841b: 6–9, pl. CCXCI, fig. 697). Based on these differences, S. caesar is not synonymous with S. scaber , as proposed by Simon (1872b) [misidentified as H. afer ], S. indus , as proposed by Kraepelin (1894: 46) [as S. ceylonicus ], or C. fulvipes , as proposed by Couzijn (1981). It is also distinct from S. serratus , for which it was mistaken by Pocock (1894a, 1900a), Kraepelin (1899), and others ( Roewer, 1943; Takashima, 1945). Srilankametrus caesar (C.L. Koch, 1841) , comb. nov. et stat. rev., is therefore revalidated, and the following new synonym presented: Heterometrus (Chersonesometrus) granulomanus Couzijn, 1981 = Srilankametrus caesar (C.L. Koch, 1841) , syn. nov.

Couzijn (1981) based H. (C.) granulomanus on a holotype and paratype (MNHN RS 0087) that were subsequently redescribed and illustrated by Tikader and Bastawade (1983: 577−581). The species was listed as valid by subsequent authors ( Fet, 2000; Kovařík, 1998, 2002) until synonymized with H. madraspatensis by Kovařík (2004: 29) based on an appeal to authority: “ Heterometrus (Chersonesometrus) granulomanus …in my opinion is a synonym of H. madraspatensis .” Interestingly, two females in a similar state of preservation, with the same locality data as the holotype of B. caesar , and precisely matching the paratype and nontype females of H. (C.) granulomanus (MNHN RS 0079, 0087, 3246) were misidentified as H. latimanus by Kovařík (2004: 21, 51, table 2; 2009: 34, 35, 39, 47, 49, 73, 100, table 1, figs. 3, 4, 198), as noted by Tahir and Prendini (2014: 6, 20). Setting aside that Couzijn (1981: 142) synonymized P. madraspatensis with H. (C.) fulvipes and considered the latter closely related to H. (C.) granulomanus — even misidentifying as H. (C.) fulvipes several specimens with the same collection data as the types of H. (C.) granulomanus (MNHN RS 6168) ( Couzijn, 1981: 134)—these species share little in common. Srilankametrus caesar differs from C. fulvipes and C. madraspatensis in coloration, carination of the retrolateral surface of the pedipalp chela manus, and macrosetae of the leg basitarsi. Accordingly, Kovařík’s (2004) synonymy of H. (C.) granulomanus with H. madraspatensis is rejected. Based on phylogenetic analysis (fig. 10), S. caesar is most closely related to S. couzijni , S. gravimanus , and S. yaleensis , from which it differs in the surface macrosculpture of the carapace and pedipalp chela, shape of the pedipalp chela, and macrosetae of the leg basitarsi.

As discussed below, a nontype male from Palni Hills, India (SMF 5332/1), listed in the original description of H. (C.) granulomanus , and which closely matches an adult male from Tanjore, India (BMNH 1896.7.30.121), misidentified as H. gravimanus by Pocock (1900a), Tikader and Bastwawade (1983) and Kovařík (2004), is newly described as the paratype of S. couzijni .

MATERIAL EXAMINED: 1 ♀, 1 subad. ♀, 2 ♂, 1 juv. ♀ (ZMB 7244). “Amerique?,” 1 ♀ (MNHN RS 0120). “India?,” 1 ♂ (ZMB 2283), 1 ♀ (MNHN RS 0149 [Simon coll. 8212]). INDIA: 1 ♀ (BMNH). “Ind. Or.” [India Orientalis], “Ostindien” [East India], 2 ♂ (ZMB 2282). Puducherry: Puducherry Distr.: Pondichéry [Pondicherry /Puduchcheri/ Puducherry, 11°59′N 79°49′E], viii.1901, M. Maindron, 1 ♂, 1 ♀ (MNHN RS 0079), 1 ♀, 1 subad. ♂, 2 subad. ♀, 1 juv. ♀ (MNHN RS 0087), 5 ♂, 10 ♀, 3 subad. ♂, 7 subad. ♀, 3 juv. ♀ (MNHN RS 6168), 1902 [probably viii.1901], M. Maindron, 1 ♀ (MNHN RS 3246).

Figures

FIGURE 10. Phylogeny of Asian forest scorpions (Scorpionidae: Heterometrinae Simon, 1879, stat. nov.), obtained by simultaneous phylogenetic analysis of 186 morphological characters and 4188 aligned DNA nucleotides from three mitochondrial and two nuclear gene loci (Loria and Prendini, in press). Maximum likelihood tree with unambiguous morphological synapomorphies optimized. Black circles indicate uniquely derived apomorphic states, white circles parallel derivations of apomorphic states. Numbers above circles indicate characters, numbers below indicate states (appendix 1).

FIGURE 27. Srilankametrus Couzijn, 1981, stat. nov. et stat. rev., carapace, dorsal aspect. A, B. Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev., A. ♂, B. ♀ (MNHN RS 0079). C, D. Srilankametrus couzijni, sp. nov., C. holotype ♂ (BMNH 1896.7.30.121), D. paratype ♂ (SMF 5332/1). E, F. Srilankametrus gravimanus (Pocock, 1894), comb. nov., E. ♂, F. ♀ (AMNH), Kokmotte, Sri Lanka. Scale bars = 2 mm.

FIGURE 41. Srilankametrus Couzijn, 1981, stat. nov. et stat. rev., sternum and pectines, ventral aspect. A, B. Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev., A. ♂, B. ♀ (MNHN RS 0079). C, D. Srilankametrus couzijni, sp. nov., C. holotype ♂ (BMNH 1896.7.30.121), D. paratype ♂ (SMF 5332/1). E, F. Srilankametrus gravimanus (Pocock, 1894), comb. nov., E. ♂, F.♀ (AMNH), Kokmotte, Sri Lanka. Scale bars = 2 mm.

FIGURE 56. Srilankametrus Couzijn, 1981, stat. nov. et stat. rev., legs I–IV, tibiae, basitarsi and telotarsi, ventral aspect. A–D. Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev., ♂ (MNHN RS 0079). E–H. Srilankametrus couzijni, sp. nov., holotype ♂ (BMNH 1896.7.30.121). I–L. Srilankametrus gravimanus (Pocock, 1894), comb. nov., ♂ (AMNH), Kokmotte, Sri Lanka. Scale bars = 2 mm.

FIGURE 76. Srilankametrus Couzijn, 1978, stat. nov. et stat. rev., metasoma and telson, dorsal aspect. A. Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev., ♂ (MNHN RS 0079). B. Srilankametrus couzijni, sp. nov., holotype ♂ (BMNH 1896.7.30.121). C. Srilankametrus gravimanus (Pocock, 1894), comb. nov., ♂ (AMNH), Kokmotte, Sri Lanka. D. Srilankametrus indus (DeGeer, 1778), comb. nov., ♂ (AMNH), Kandy, Sri Lanka. E. Srilankametrus pococki, sp. nov., holotype ♀ (AMNH), Kodigala, Sri Lanka. F. Srilankametrus serratus (Pocock, 1900), comb. nov., ♂ (ZMH), Ceylon [Sri Lanka]. Scale bars = 5 mm

FIGURE 77. Srilankametrus Couzijn, 1978, stat. nov. et stat. rev., metasoma and telson, lateral aspect. A. Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev., ♂ (MNHN RS 0079). B. Srilankametrus couzijni, sp. nov., holotype ♂ (BMNH 1896.7.30.121). C. Srilankametrus gravimanus (Pocock, 1894), comb. nov., ♂ (AMNH), Kokmotte, Sri Lanka. D. Srilankametrus indus (DeGeer, 1778), comb. nov., ♂ (AMNH), Kandy, Sri Lanka. E. Srilankametrus pococki, sp. nov., holotype ♀ (AMNH), Kodigala, Sri Lanka. F. Srilankametrus serratus (Pocock, 1900), comb. nov., ♂ (ZMH), Ceylon [Sri Lanka]. Scale bars = 5 mm.

FIGURE 78. Srilankametrus Couzijn, 1978, stat. nov. et stat. rev., metasoma and telson, ventral aspect. A. Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev., ♂ (MNHN RS 0079). B. Srilankametrus couzijni, sp. nov., holotype ♂ (BMNH 1896.7.30.121). C. Srilankametrus gravimanus (Pocock, 1894), comb. nov., ♂ (AMNH), Kokmotte, Sri Lanka. D. Srilankametrus indus (DeGeer, 1778), comb. nov., ♂ (AMNH), Kandy, Sri Lanka. E. Srilankametrus pococki, sp. nov., holotype ♀ (AMNH), Kodigala, Sri Lanka. F. Srilankametrus serratus (Pocock, 1900), comb. nov., ♂ (ZMH), Ceylon [Sri Lanka]. Scale bars = 5 mm.

FIGURE 199. Map of South India and Sri Lanka, plotting known locality records of two species of Sahyadrimetrus, gen. nov., and four species of Srilankametrus Couzijn, 1981, stat. nov., based on material examined, and verified records from virtual museums and the literature: Sahyadrimetrus mathewi, sp. nov.; Sahyadrimetrus scaber (Thorell, 1876), comb. nov.; Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev.; Srilankametrus couzijni, sp. nov.; Srilankametrus gravimanus (Pocock, 1894), comb. nov.; Srilankametrus yaleensis (Kovařík et al., 2019), comb. nov.