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

The genera and species of the Asian forest scorpions (Scorpionidae Latreille, 1802) are revised based on a phylogenetic analysis of 186 morphological characters and 4188 base pairs of concatenated DNA sequence from three mitochondrial loci and two nuclear loci. Revision of the Asian scorpionids required a critical reappraisal of the suprageneric classification of Scorpionidae, on the basis of which the monotypic Indian scorpionoid genus, RugodentusBastawade et al., 2005, stat. rev., and its type species, Rugodentus keralaensisBastawade et al., 2005, stat. rev., are revalidated, and subfamily Rugodentinae Bastawade et al., 2005, revalidated and elevated to the rank of family, Rugodentidae Bastawade et al., 2005, stat. nov. et stat. rev.; Heterometrinae Simon, 1879, stat. nov., and Opistophthalminae Rossi, 2016, stat. nov., are elevated to the rank of subfamily; PandinopsisVachon, 1974, stat. nov., and Pandipalpus Rossi, 2015, stat. nov., are elevated to the rank of genus, resulting in two new combinations: Pandinopsis dictator (Pocock, 1888), comb. nov., and Pandipalpus viatoris (Pocock, 1890), comb. nov.; and 10 new synonyms are presented: Pandinopsini Rossi, 2016 = Pandininae Thorell, 1876, syn. nov.; Protophthalmini Rossi, 2016 = Opistophthalminae Rossi, 2016, syn. nov.; ProtophthalmusLawrence, 1969 = Opistophthalmus C.L. Koch, 1837, syn. nov.; Pandinoides (Dunlopandinoides) Rossi, 2016 = PandinoidesFet, 2000, syn. nov.; Pandinurus (Pandicaporiaccous) Rossi, 2015 = Pandiborellius Rossi, 2015, syn. nov.; Buthus defensor C.L. Koch, 1837 = Pandinurus gregoryi (Pocock, 1896), syn. nov.; Buthus heros C.L. Koch, 1837 = Pandinurus exitialis (Pocock, 1888), syn. nov.; Pandinus loweiKovařik, 2012 = Pandipalpus viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) pygmaeus Rossi, 2015 = Pandipalpus viatoris (Pocock, 1890), syn. nov.; Pandinus intermedius Borelli, 1919 = Pandinurus citernii (Borelli, 1919), syn. nov. The following revisions are implemented to the classification of the Asian forest scorpions (Heterometrinae). Three former subgenera of Heterometrus Ehrenberg, 1828 are revalidated and elevated to the rank of genus: ChersonesometrusCouzijn, 1978, stat. nov. et stat. rev.; JavanimetrusCouzijn, 1981, stat. nov. et stat. rev.; and SrilankametrusCouzijn, 1981, stat. nov. et stat. rev. One subgenus is elevated to the rank of genus: GigantometrusCouzijn, 1978, stat. nov. Two new genera and eight new species are described: Deccanometrus, gen. nov.; Sahyadrimetrus, gen. nov.; Chersonesometrus bastawadei, sp. nov.; Chersonesometrus hendersoni, sp. nov.; Chersonesometrus nathanorum, sp. nov.; Chersonesometrus shivashankari, sp. nov.; Sahyadrimetrus mathewi, gen. et sp. nov.; Sahyadrimetrus tikaderi, gen. et sp. nov.; Srilankametrus couzijni, sp. nov.; Srilankametrus pococki, sp. nov. Heterometrus sensu stricto is restricted to eight species of the nominotypical subgenus, all other species, formerly placed in Heterometrus, are transferred to appropriate genera, five species are revalidated, and two subspecies elevated to the rank of species, resulting in 28 new combinations: Chersonesometrus beccaloniae (Kovařik, 2004), comb. nov.; Chersonesometrus fulvipes (C.L. Koch, 1837), comb. nov.; Chersonesometrus madraspatensis (Pocock, 1900), comb. nov.; Chersonesometrus pelekomanus (Couzijn, 1981), comb. nov. et stat. rev.; Chersonesometrus tristis (Henderson, 1919), comb. nov.; Chersonesometrus wroughtoni (Pocock, 1899), comb. nov.; 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 (Kovařik, 2004), comb. nov.; Deccanometrus xanthopus (Pocock, 1897), comb. nov.; Gigantometrus swammerdami (Simon, 1872), comb. nov.; Gigantometrus titanicus (Couzijn, 1981), comb. nov. et stat. rev.; Heterometrus glaucus (Thorell, 1876), comb. nov. et stat. rev.; Heterometrus laevigatus (Thorell, 1876), comb. nov. et stat. rev.; Heterometrus silenus (Simon, 1884), comb. nov. et stat. rev.; Javanimetrus cyaneus (C.L. Koch, 1836), comb. nov.; Sahyadrimetrus barberi (Pocock, 1900), comb. nov.; Sahyadrimetrus kanarensis (Pocock, 1900), comb. nov.; Sahyadrimetrus rugosus (Couzijn, 1981), comb. et stat. nov.; Sahyadrimetrus scaber (Thorell, 1876), comb. nov.; Srilankametrus caesar (C.L. Koch, 1841), comb. nov. et stat. rev.; Srilankametrus gravimanus (Pocock, 1894), comb. nov.; Srilankametrus indus (DeGeer, 1778), comb. nov.; Srilankametrus serratus (Pocock, 1900), comb. nov; Srilankametrus yaleensis (Kovařik et al., 2019), comb. nov. Twenty-seven new synonyms are presented: Scorpio leiodermaDufour, 1856 = Sahyadrimetrus scaber (Thorell, 1876), syn. nov.; Palamnaeus costimanus var. β borneensis Thorell, 1876 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Palamnaeus liophysaThorell, 1888 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Palamnaeus oatesii Pocock, 1900 = Heterometrus petersii (Thorell, 1876), syn. nov.; Palamnaeus swammerdami flavimanus Pocock, 1900 = Gigantometrus swammerdami (Simon, 1872), syn. nov.; Heterometrus liophysa var. madoerensisKopstein, 1921 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus laevifronsRoewer, 1943 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus (Chersonesometrus) granulomanusCouzijn, 1981 = Srilankametrus caesar (C.L. Koch, 1841), syn. nov.; Heterometrus (Heterometrus) liophysa separatusCouzijn, 1981 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus (Heterometrus) liophysa spartanicusCouzijn, 1981 = Heterometrus glaucus (Thorell, 1876), syn. nov.; Heterometrus (Heterometrus) longimanus bengkalitensisCouzijn, 1981 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Heterometrus (Heterometrus) longimanus marmoratusCouzijn, 1981 = Heterometrus longimanus (Herbst, 1800), syn. nov.; Heterometrus (Heterometrus) petersii mindanaensisCouzijn, 1981 = Heterometrus silenus (Simon, 1884), syn. nov.; Heterometrus (Heterometrus) spinifer solitariusCouzijn, 1981 = Heterometrus spinifer (Ehrenberg, 1828), syn. nov.; Heterometrus (Srilankametrus) indus laevitensusCouzijn, 1981 = Srilankametrus indus (DeGeer, 1778), syn. nov.; Heterometrus (Heterometrus) keralaensisTikader and Bastawade, 1983 = Sahyadrimetrus rugosus (Couzijn, 1981), syn. nov.; Heterometrus cimrmaniKovařik, 2004 = Heterometrus laevigatus (Thorell, 1876), syn. nov.; Heterometrus mysorensisKovařik, 2004 = Chersonesometrus tristis (Henderson, 1919), syn. nov.; Heterometrus nepalensisKovařik, 2004 = Deccanometrus bengalensis (Pocock, 1900), syn. nov.; Heterometrus rolcikiKovařik, 2004 = Sahyadrimetrus scaber (Thorell, 1876), syn. nov.; Heterometrus sejnaiKovařik, 2004 = Javanimetrus cyaneus (C.L. Koch, 1836), syn. nov.; Heterometrus tibetanusLourenco et al., 2005 = Deccanometrus bengalensis (Pocock, 1900), syn. nov.; Heterometrus liangiZhu and Yang, 2007 = Heterometrus silenus (Simon, 1884), syn. nov.; Heterometrus telanganaensis Javed et al., 2010 = Deccanometrus xanthopus (Pocock, 1897), syn. nov.; Heterometrus atrascorpiusMirza et al., 2012 = Chersonesometrus beccaloniae (Kovařik, 2004), syn. nov.; Heterometrus minotaurusPliskova et al., 2016 = Heterometrus laevigatus (Thorell, 1876), syn. nov.; Heterometrus bastawadei Rossi, 2016 = Rugodentus keralaensisBastawade et al., 2005, syn. nov. Another 25 synonyms by previous authors are confirmed, for a total of 51 synonyms in subfamily Heterometrinae. Revised diagnoses with comparative images, and a key and distributional atlas of the genera and species are provided, along with a summary of available data for their ecology and conservation status, where applicable.