The biogeographical history of Asian keelback snakes of the genusHebius(Squamata: Colubridae: Natricinae) in the Ryukyu Archipelago, Japan

Abstract

The islands of the central and southern parts of the Ryukyu Archipelago and Taiwan accommodate suites of allopatric endemics for many terrestrial vertebrates. These subregions, which were isolated as three independent islands since the Late Miocene period, became connected to the continent by the Early Pleistocene land bridge (EPL). Many endemics are considered to have emerged through vicariant events accompanied with the collapse of EPL, but this is not well corroborated. We conducted divergence dating analyses for samples of four allopatric keelback snakes from 20 islands using a published timeframe for natricine snakes. Phylogenetic analyses recognised the existence of eight subclades for the four species. Four of the five subclades in the Central Ryukyus have emerged after the initial isolation of three subregions but before the EPL collapse, indicating iterative divergences within the super-island and stagnation of diverged entities. In contrast, the Southern Ryukyus was subjected to multiple incursions from both Central Ryukyus and Taiwan, irrespective of the EPL connection. Consequently, among all seven nodes, only two were consistent with the assumption of vicariance. These findings suggest that dispersals and subsequent nonvicariant divergences have contributed to the formation of the current phylogeographical pattern in this continental island chain.