Systematics and phylogenetic relationships of Whip snakes (Hierophis Fitzinger) and Zamenis andreana Werner, 1917 (Reptilia: Squamata: Colubrinae)

Abstract

Morphological and molecular data of Coluber (sensu lato) andreanus and all recognised species of the Palaearctic whip snake genus Hierophis Fitzinger are presented. Morphologically, Andreas' racer shows remarkable similarities to dwarf snakes (Eirenis spp., Pseudocyclophis persicus). Derived conditions of head and body pholidosis including dorsal scale reduction pattern and the number of apical pits separate Eirenis Jan and C. (s.l.) andreanus from Hierophis spp. Character states strongly support sister group relationship of dwarf snakes to Andreas' racer. Molecular analyses confirm monophyly of Hierophis spp., C. (s.l.) andreanus, and dwarf snakes of the genus Eirenis, and paraphyly of Hierophis auct. MtDNA sequences indicate a basal position of the eastern Palaearctic H. spinalis vis-à-vis the western species group (H. cypriensis, H. gemonensis, H. viridiflavus). These taxa belong to an early radiation within whip and dwarf snakes. The eastern Mediterranean H. caspius, H. jugularis, and H. schmidti appear to represent a paraphyletic grouping including C. (s.l.) andreanus (12S rDNA sequence data). Conflicting molecular and morphological results are discussed and the character phylogeny of external morphological features is re-assessed.