The out-of-India hypothesis: evidence from an ancient centipede genus, Rhysida (Chilopoda: Scolopendromorpha) from the Oriental Region, and systematics of Indian species

The Oriental Region has been a focus of biogeographical research for more than two centuries. We examined systematics and biogeography of the centipede genus Rhysida in this region. A robust species hypothesis for the Indian subcontinental and Southeast Asian Rhysida clade uses molecular, morphological and distribution data. Twelve species are recognized in two monophyletic species complexes, eight belonging to the Rhysida immarginata and four to the Rhysida longipes species complex. They include Rhysida aspinosa, Rhysida crassispina, R. immarginata, R. longipes and seven new species, five of which are formally named in this paper: Rhysida ikhalama, Rhysida konda, Rhysida lewisi, Rhysida pazhuthara and Rhysida sada. The nine Rhysida species are documented taxonomically and their morphological variation is reviewed. An integrative systematic approach reveals that diversity of Rhysida in the Indian subcontinent has been underestimated. Both species complexes started to diversify in the Early to Late Cretaceous in the Indian subcontinent. The out-of-India hypothesis is supported in both clades, because Southeast Asian species are nested in Indian subcontinental clades. Historical biogeographical analyses suggest two independent post-collision dispersal events, one in the immarginata clade and another where R. longipes expanded its range into Southeast Asia.