Srilankametrus indus ( DeGeer, 1778 ), Prendini & Loria, 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

publication ID

http://doi.org/ 10.1206/0003-0090.442.1.1

persistent identifier

http://treatment.plazi.org/id/5E6CB374-FE51-6CAC-FF08-60DAFF3CDC2E

treatment provided by

Felipe

scientific name

Srilankametrus indus ( DeGeer, 1778 )
status

 

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   View Cited Treatment (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) GoogleMaps   ; 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]) GoogleMaps   ; 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]) GoogleMaps   ; Paradeniya [Peradeniya, 07°16′N 80°36′E], vii.1914, B.H. Buxton, 2 ♂, 1 subad. ♂, 1 subad. ♀, 1 juv. ♀ (MNHN RS 0081) GoogleMaps   .

Kingdom

Animalia

Phylum

Arthropoda

Class

Arachnida

Order

Scorpiones

Family

Scorpionidae

Genus

Srilankametrus

Loc

Srilankametrus indus ( DeGeer, 1778 )

Prendini, Lorenzo & Loria, Stephanie F. 2020
2020
Loc

Heterometrus gravimanus: Kovařík, 2004: 1

Kovarik, F. 2004: 1
2004
Loc

Heterometrus (Srilankametrus) indus: Couzijn, 1981: 80

Kovarik, F. 2002: 17
Fet, V. 2000: 446
Kovarik, F. 1998: 137
Kovarik, F. 1992: 185
Tikader, B. K. & D. B. Bastawade 1983: 544
Couzijn, H. W. C. 1981: 80
1981
Loc

Heterometrus (Srilankametrus) indus indus: Couzijn, 1981: 39

Fet, V. 2000: 447
Vachon, M. 1982: 78
Couzijn, H. W. C. 1981: 39
1981
Loc

Heterometrus (Srilankametrus) indus laevitensus Couzijn, 1981: 39

Fet, V. 2000: 447
Kovarik, F. 1998: 137
Couzijn, H. W. C. 1981: 39
1981
Loc

Heterometrus (Heterometrus) indus:

Couzijn, H. W. C. 1978: 330
1978
Loc

Palamnaeus indus:

Pocock, R. I. 1900: 96
1900
Loc

Palamnaeus ndus:

Pocock, R. I. 1900: 86
1900
Loc

Heterometrus indus:

Pliskova, J. & F. Kovarik & O. Kosulic & F. St'ahlavsky 2016: 1
Tahir, H. M. & L. Prendini 2014: 11
Javed, S. M. M. & Z. A. Mirza & F. Tampal & W. R. Lourenco 2010: 147
Kovarik, F. 2004: 17
Lourenco, W. R. & J. L. Cloudsley-Thompson 1996: 140
Bucherl, W. 1959: 269
Takashima, H. 1945: 92
Roewer, C. F. 1943: 226
Kastner, A. 1941: 233
Giltay, M. L. 1931: 8
Roewer, C. F. 1929: 612
Lampe, E. 1918: 198
Kraepelin, K. 1899: 109
1899
Loc

Palamnaeus ceylonicus:

Kraepelin, K. 1898: 439
1898
Loc

Scorpio megacephalus:

Pocock, R. I. 1893: 307
1893
Loc

Pandinus ceylonicus:

Karsch, F. 1884: 69
1884
Loc

Scorpio crassimanus

Kraepelin, K. 1899: 113
Becker, L. 1880: 140
1880
Loc

Pandinus megacephalus: Thorell, 1876b: 164

Thorell, T. 1893: 381
Karsch, F. 1884: 69
Thorell, T. 1876: 164
1876
Loc

Heterometrus afer: Simon, 1872b: 53

Simon, E. 1872: 53
1872
Loc

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

Gervais, P. M. 1844: 60
1844
Loc

Scorpio (Buthus) ceylanicus:

Gervais, P. M. 1844: 62
1844
Loc

Buthus afer

Koch, C. L. 1836: 17
1836
Loc

Buthus megacephalus C.L. Koch, 1836: 73–75

Thorell, T. 1876: 164
Koch, C. L. 1850: 88
Gervais, P. M. 1844: 60
Koch, C. L. 1836: 75
1836
Loc

Scorpio ceylonicus

Couzijn, H. W. C. 1981: 129
Lonnberg, E. 1897: 186
Kraepelin, K. 1894: 46
Kraepelin, K. 1894: 31
Pocock, R. I. 1893: 308
Herbst, J. F. W. 1800: 83
1800
Loc

Scorpio indus

Kovarik, F. 2004: 2
DeGeer, C. & K. DeGeer 1783: 132
DeGeer, C. & K. DeGeer 1778: 343
1778
Loc

Scorpio afer

Dufour, L. 1856: 563
Latreille, P. A. 1803: 267
Herbst, J. F. W. 1800: 38
Fabricius, J. C. 1781: 550
DeGeer, C. & K. DeGeer 1778: 341
Fabricius, J. C. 1775: 399
Linnaeus, C. & C. von Linne 1775: 1091
Linnaeus, C. & C. von Linne 1758: 624
1758