Morphological and mitochondrial DNA data reshuffle the taxonomy of the genera Atopochetus Attems, Litostrophus Chamberlin and Tonkinbolus Verhoeff (Diplopoda: Spirobolida: Pachybolidae), with descriptions of nine new species

Species-level taxonomy and phylogeny of two genera of South-East (SE) Asian pachybolid millipedes are analysed with a combination of morphological characters and DNA sequences (two mitochondrial gene fragments: COI and 16S rRNA). Strong support is found for the genera Litostrophus Chamberlin, 1921 and Atopochetus Attems, 1953 and for a clade consisting of Litostrophus + Atopochetus. Four species of Litostrophus are recognised and (re)described: L. segregatus Chamberlin, 1921 (type species of the genus), L. scaber (Verhoeff, 1938), comb. nov., L. chamaeleon, sp. nov. and L. saraburensis, sp. nov. The genus Tonkinbolus Verhoeff, 1938 (type species T. scaber Verhoeff, 1938) is synonymised under Litostrophus. Atopochetus (type species A. rubropunctatus Attems, 1953), hitherto considered a dubious synonym of Aulacobolus Pocock, 1903, is re-instated for several species until now placed in Tonkinbolus and seven new species. All in all, 10 species of Atopochetus are (re)described: A. dollfusii (Pocock, 1893) (= Aulacobolus rubropunctatus Attems, 1938, syn. nov.), A. moulmeinensis (Pocock, 1893), A. sumatranus (Carl, 1906) (= Trachelomegalus laciniatus Attems, 1937, syn. nov.), A. anaticeps, sp. nov., A. helix, sp. nov., A. setiferus, sp. nov., A. spinimargo, sp. nov., A. truncatus, sp. nov., A. uncinatus, sp. nov. and A. weseneri, sp. nov. Three species recently included in Tonkinbolus are left incertae sedis because they are based exclusively on female type specimens: Spirobolus capucinus Porat, 1896, S. caudulanus Karsch, 1881 and S. macrurus Pocock, 1893. The results of the DNA analysis (COI and 16S rRNA) are congruent with morphological (gonopodal) characters in terms of delimitation of species of Litostrophus and Atopochetus. This is the first DNA study of SE Asian Pachybolidae and as such it provides a basis for further evolutionary and biogeographic studies of SE Asian millipedes.