Srilankametrus indus ( DeGeer, 1778 ), 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 : 417-421

Srilankametrus indus ( DeGeer, 1778) ,

comb. nov.

Figures 7H, 9H, 8G, I, 10, 28A, B, 42A, B, 57A– D, 76D, 77D, 78D, 198, 238–242, table 1

Scorpio afer Linnaeus, 1758: 624 (part; synonymized by DeGeer, 1778: 341); 1767: 1038 (part); Fabricius, 1775: 399 (part); Linnaeus, 1775: 1091 (part); Fabricius, 1781: 550 (part); 1787: 348 (part); 1793: 434 (part); Herbst, 1800: 38–42 (part); Latreille, 1803: 267 (part); 1804: 120–122 (part); 1817: 106 (part); Dufour, 1856: 563, 571, 573, 574, 584, 590, 593, 614, 615, 622, 629, 631, 632, 641, 642, 644, 645, 648 (part).

Scorpio indus DeGeer, 1778: 341–343 ; 1783: 132–134; Kovařík, 2004: 2.

Scorpio ceilonicus: Herbst, 1800 : pl. V, fig. 1.

Scorpio ceylonicus Herbst, 1800: 83 , 84 (synonymized by Kraepelin, 1894: 46); Pocock, 1893: 308, 309; Kraepelin, 1894: 31, 46–51, 54, 55, 57, 240, 246, pl. I, fig. 29 (misidentification, part); Lönnberg, 1897b: 186 (misidentification); 1898a: 83; Couzijn, 1981: 129.

Buthus afer (nec Scorpio afer Linnaeus, 1758 ): C.L. Koch, 1836: 17–19 (misidentification, part); 1850: 87 (part).

Buthus megacephalus C.L. Koch, 1836: 73–75 , pl. XCVII, fig. 224 (synonymized by Thorell, 1876b: 164, 168, 203, 204); Gervais, 1844a: 60; C.L. Koch, 1850: 88.

Scorpio (Buthus) afer: Gervais, 1844a: 60 (part).

Scorpio (Buthus) ceylanicus: Gervais, 1844a: 62 .

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

Pandinus megacephalus: Thorell, 1876b: 164 , 168, 203–205, 209–211 (misidentification); Karsch, 1884: 69; Thorell, 1893: 381, 382 (misidentification, part).

Scorpio crassimanus Becker, 1880: 140 , 141, pl. III, fig. 1 (synonymized by Kraepelin, 1899: 113).

Pandinus ceylonicus: Karsch, 1884: 69 .

Scorpio megacephalus: Pocock, 1893: 307 , 309, 311.

Palamnaeus ceylonicus: Kraepelin, 1898: 439 , 441.

Heterometrus indus: Kraepelin, 1899: 109 , 113, 114 (part); Werner, 1902: 601; Lampe, 1918: 198; Roewer, 1929: 612 (part); Giltay, 1931: 8; Werner, 1934: 277 (part); Kästner, 1941: 233 (part), fig. 212; Roewer, 1943: 226 (misidentification, part); Takashima, 1945: 92; Bücherl, 1959: 269; Lourenço and Cloudsley-Thompson, 1996: 140, figs. 4, 5; Kovařík, 2004: 17, 20, 21, 32, 51, 52, table 2 (part); 2009: 35, 38, 47–49, 80, 100, 105, table 1 (part), figs. 50–56, 199, 200, 249–252; Javed et al., 2010a: 147; Tahir and Prendini, 2014: 11, 12; Kovařík et al., 2016: 1, 5, 10, 97–104, 106, 107, 110, 111, 115, 117–119 (part), figs. 15, 482–507, 573, 575, 577; 2019a: 12, fig. 53.

Palamnaeus indus: Pocock, 1900a: 96 , 97, fig. 20A.

Palamnaeus ndus: Pocock, 1900a: 86 .

Heterometrus (Heterometrus) indus: Couzijn, 1978: 330 , table 1.

Heterometrus (Srilankametrus) indus: Couzijn, 1981: 80 , 82, 94, 121, 123, 172, 181, figs. 21, 60, 66a (part); Tikader and Bastawade, 1983: 544; Kovařík, 1992: 185; 1998: 137, 138 (part); Fet, 2000: 446, 447 (part); Kovařík, 2002: 17 (part); 2004: 17.

Heterometrus (Srilankametrus) indus indus: Couzijn, 1981: 39 , 82, 121–124, 129, 172, 191, table 7 (part), fig. 60; Vachon, 1982: 78, 79, 96, 101, 103–105, 110, figs. 50, 70–72, 75, 80, 81; Tikader and Bastwade, 1983: 550, 555, 561; Fet, 2000: 447.

Heterometrus (Srilankametrus) indus laevitensus Couzijn, 1981: 39 , 83, 124, 125, 172, 191, table 7, figs. 36, 60; Kovařík, 1998: 137; Fet, 2000: 447, 448; syn. nov.

Heterometrus gravimanus: Kovařík, 2004: 1 , 2, 15, 17, 44, 51, table 2 (misidentification, part); 2009: 35, 38, 47, table 1 (misidentification, part).

TYPE MATERIAL: Buthus megacephalus : 2 syntypes, probably East Indies [lost]. Scorpio afer : Syntypes [sex?], “Africa,” [lost]. Scorpio indus : Holotype: 1 ♀ (NRS), “India” [examined]. Heterometrus (Srilankametrus) indus laevitensus : Holotype ♂ (MNHN RS 0089), Madras [examined]. SRI LANKA: Scorpio ceylonicus : Syntypes [sex?], Ceylon [lost]. Scorpio crassimanus : Holotype: 1 subad. ♀ (MRHNB [S.G. 38]), Netherlands Indies [examined; label in bottle: “Ceylan”].

According to the original description ( Becker, 1880: 140), the holotype of Scorpio crassimanus , from “Indus néerlandaises,” was deposited in the “Musée de Bruxelles.” Kraepelin (1894) synonymized S. crassimanus with H. indus . A specimen from the MRHNB, previously examined by K. Kraepelin, and matching Becker’s (1880) description and habitus illustration (pl. III, fig. 1) of S. crassimanus , e.g., in total length, carapace shape (including deep notch in anterior margin), granular pedipalp chela manus, and pectinal tooth counts, was examined during the present investigation. It was concluded to be the holotype of S. crassimanus , despite the absence of a label indicating the same, and the type locality given as “ Ceylan ” rather than “Indus néerlandaises.” The Netherlands Indies is modern Indonesia. However, Ceylon (i.e., Sri Lanka) was also a Dutch colony until approximately 1800.

DIAGNOSIS: Srilankametrus indus may be separated from other species of Srilankametrus as follows. The carapace is markedly dorsoventrally compressed, the lateral surfaces sloping gently (fig. 28A, B), in S. indus but slightly dorsoventrally compressed, the lateral surfaces sloping moderately, in S. serratus , and vaulted, the lateral surfaces sloping steeply, in S. caesar , S. couzijni , S. gravimanus , and S. yaleensis . The interocular and posterior sutures are absent in S. indus but present, the posterior sutures extending past the median ocular tubercle and connected anteriorly to the posterior bifurcations of the interocular suture in all other species except S. pococki . The carapace interocular surface is entirely smooth (fig. 28A, B) in S. indus whereas the frontal lobes and medial region of the interocular surface are granular with smooth areas in S. caesar . The carapace anterolateral and mediolateral surfaces are smooth or nearly so in S. indus but granular in all other species. The carapace posterolateral surfaces of the female are granular in S. indus but smooth or nearly so in S. gravimanus and S. yaleensis . The pedipalp patella dorsomedian carina of the female is absent or obsolete in S. indus but entirely to predominantly costate in S. caesar . The patella retro- dorsal carina of the female is absent or obsolete in S. indus 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. indus but absent or obsolete in S. gravimanus and S. yaleensis . The pedipalp chela of the adult male (fig. 241) is moderately to densely setose in S. indus but sparsely setose in all other species except S. pococki . The chela manus dorsal surface (between the dorsomedian and digital carinae) is curved and slightly to markedly convex in S. indus but flat in S. pococki . The proximal margin (lobe) of the dorsal surface is moderately curved and aligned with the proximal margin of the condyle (articulation with patella) in S. indus but markedly curved and proximal to the proximal margin of the condyle in S. couzijni . The dorsomedian carina is obsolete on the chela fixed finger and manus in S. indus , pronounced and costate on the fixed finger and distally on the manus in S. caesar , S. couzijni , and S. gravimanus , and pronounced and costate on the fixed finger and distal three-quarters of the manus in S. yaleensis . 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. 241) of S. indus but similar to the DSC–DC in the male of S. gravimanus . The dorsal secondary and subdigital carinae of the male (fig. 241) are absent or obsolete in S. indus but entirely to predominantly granular in S. caesar , S. couzijni , and S. serratus , and entirely to predominantly costate in S. gravimanus and S. yaleensis . The digital carina is absent or obsolete (figs. 241, 242) in S. indus but entirely to predominantly granular in S. caesar and S. serratus , and entirely to predominantly costate in S. couzijni , S. gravimanus , and S. yaleensis . The retromedian carina of the male is absent or obsolete in S. indus but entirely to predominantly granular in S. caesar and S. serratus , and entirely to predominantly costate in S. couzijni , S. gravimanus , and S. yaleensis . The manus ventral surface is angled, with the axis of the retroventral carina ventral to the axis of the ventro- median carina in S. indus , but flat, with the axes of the retroventral and ventromedian carinae in approximately the same plane in all other species except S. pococki . Macroseta st on the retroventral surfaces of the basitarsi of legs I and II is setiform (fig. 57A–D) in S. indus but spiniform in S. caesar , S. gravimanus , and S. serratus . The lateral surfaces of mesosomal tergites I–VI are smooth in S. indus but granular in all other species except S. pococki and S. yaleensis . The dorsosubmedian carinae are costate on metasomal segments I–IV (fig. 76D) in S. indus , costate on I and II or I–III and granular or costate-granular on III and IV or IV, in S. yaleensis , and granular or costate-granular on I– IV in all other species. The ventral intercarinal surfaces of metasomal segment IV are smooth in the male and female (fig. 78D) of S. indus but granular in the male of S. couzijni and S. gravimanus and the male and female of S. caesar . The dorsolateral carinae of metasomal segment V are obsolete and discontinuous to absent (fig. 77D) in S. indus but distinct and continuous in all other species except S. pococki . The dorsal intercarinal surface of metasomal segment V is smooth in the male and female (fig. 76D) of S. indus , but granular in the male and female of S. caesar and S. gravimanus , and the male of S. couzijni , S. serratus and S. yaleensis . The telson is blackish, as dark as metasomal segment V, in S. indus but dark reddish brown, paler than segment V in S. serratus and S. yaleensis . The telson vesicle is globose in S. indus but elongate in S. caesar , S. couzijni , S. gravimanus , and S. yaleensis .

DISTRIBUTION: This species is endemic to Sri Lanka (fig. 198, table 1) and has been recorded only in the Central Province. The distribution of S. indus is allopatric with those of other species of Srilankametrus .

ECOLOGY: This species inhabits primary rainforest in the Knuckles Mountain Range in the central part of the island (fig. 7H). The locality records for which data are available vary from 220 m to 560 m above sea level. This species is fossorial and pelophilous, excavating moderately deep single-occupant burrows in hard, clayey soils, in earthen banks, under logs or stones ( Kovařík et al., 2016). Burrow entrances are characteristically broad and flat (fig. 8G, I). The following scorpions have been recorded in sympatry: the buthids Buthoscorpio sarasinorum and Reddyanus ranawanai Kovařík et al., 2016 ; and the scorpionid G. titanicus ( Kovařík et al., 2016) .

REMARKS: The following synonyms are recognized: Scorpio afer Linneaus, 1758 (part) = Srilankametrus indus ( DeGeer, 1778) , comb. nov., synonymized by DeGeer (1778); Scorpio ceylonicus Herbst, 1800 = S. indus , synonymized by Kraepelin (1894); Buthus megacephalus C.L. Koch, 1836 = S. indus , first synonymized by Thorell (1876) not Kraepelin (1894) as stated by Fet (2000: 447); Scorpio crassimanus Becker, 1880 = S. indus , first synonymized by Kraepelin (1899) not Kraepelin (1894) as stated by Fet (2000: 447).

It is doubtful that Buthus megacephalus is synonymous with S. indus , based on characters in the original description and illustration, e.g., the narrower shape of the male pedipalp chela and reddish-brown coloration of the telson, compared with the black metasoma, consistent with several Indian species, but not S. indus . For example, Thorell (1893: 381) referred a specimen from Matheran, the type locality of Deccanometrus obscurus , to Pandinus megacephalus . In the absence of a type and a type locality, however, the true identity of B. megacephalus will never be verified. Additionally, Buthus caesar is not a synonym of S. indus , as proposed by Kraepelin (1894: 46) (as Scorpio ceylonicus ) and Couzijn (1981: 123), nor is Heterometrus (H.) spinifer solitarius a synonym of S. indus , as proposed by Kovařík et al. (2016) (see above).

Fet (2000: 447) stated that Thorell (1893: 381) synonymized Scorpio afer with S. indus , but, as noted by Kovařík (2004), DeGeer (1778: 341) was the first to do so, and Thorell (1893) was followed by Kraepelin (1894: 53). Fet (2000: 447) also stated that Kraepelin (1894: 46) synonymized B. megacephalus and S. crassimanus with S. indus . However, Kraepelin (1894: 46) synonymized B. megacephalus and S. crassimanus with S. ceylonicus , not S. indus . As noted by Kovařík (2004, 2009), Thorell (1876b: 168) was the first to synonymize B. megacephalus (as Pandinus megacelaphus instead of Buthus megacephalus ) with S. indus and Kraepelin (1899: 113) was the first to synonymize S. crassimanus with S. indus , followed by Couzijn (1981: 121). Kovařík (2004), also credited Lönnberg (1898a: 83) for synonymizing B. megacephalus with S. indus but, like Kraepelin (1894), Lönnberg (1898a) synonymized B. megacephalus with S. ceylonicus .

Kovařík (2004, 2009) stated that Kraepelin (1899: 113) and Pocock (1900a: 96) synonymized S. ceylonicus with S. indus , but Kraepelin (1894: 46) was the first to do so, as noted by Fet (2000: 447). Kraepelin (1894) recognized S. ceylonicus as a valid species and listed S. indus as a synonym thereof, despite being the older name.

As noted above, Kovařík (2004: 17) erroneously synonymized H. indus laevitensus with H. gravimanus based in part on the misidentification of a specimen from Tanjore, India, as H. gravimanus . Examination of the holotype of H. indus laevitensus , allegedly from Madras, revealed it to be conspecific with S. indus , rather than with the male from Tanjore, described above as the holotype of S. couzijni , or with Sri Lankan material of S. gravimanus . Kovařík’s (2004) synonym is therefore rejected and the following new synonym presented: Heterometrus (Srilankametrus) indus laevitensus Couzijn, 1981 = Srilankametrus indus ( DeGeer, 1778) , syn. nov.

MATERIAL EXAMINED: SRI LANKA: i.1965, F. Layard, 2 ♂ (MCZ) ; Ceylan, 1 ♂ (MNHN RS 3245 [Simon coll. 5770]); Ceylan ?, Dr. H. Mutvei, 1 ♀ (MNHN RS 8117). Central Prov. : Kandy Distr.: Kandy [07°17′N 80°38′E], 1902, Dr. Bedeker, 1 ♂ (MNHN RS 3263) ; Kandy , outskirts of Udawattakele, 07°18′N 80°38′E, 217 m, 30.i.2014, L. Prendini and P. Horsley, degraded tropical forest along roadcuts, UV light detection on warm, humid, moonless night, doorkeeping at burrow entrances in roadcuts and earthen banks, 3 ♀, 2 juv. ♂, 3 juv. ♀ (AMNH), 1 juv. ♀ (AMCC [LP 12282]) ; Kandy , Udawattakele, 07°18′N 80°38′E, 560 m, 31.i–4.ii.2014, L. Prendini and P. Horsley, primary tropical rainforest on steep hills, large trees with dense canopy and sparse understorey, moderate to thick leaflitter layer, many rotten logs with rock outcrops and scattered stones in places, quartzite geology, coarse sandy-loam soil, in burrows in earthen banks, often at base of stones or under logs, some under large logs or stones, 6 ♂, 9 ♀, 4 subad. ♂, 4 subad. ♀, 5 juv. ♂, 10 juv.♀ (AMNH), 1 juv. ♀ (AMCC [LP 12283]) ; Paradeniya [Peradeniya, 07°16′N 80°36′E], vii.1914, B.H. Buxton, 2 ♂, 1 subad. ♂, 1 subad. ♀, 1 juv. ♀ (MNHN RS 0081) .