RESEARCH ARTICLE An Extremely Peramorphic Newt (Urodela: Salamandridae: Pleurodelini) from the Latest Oligocene of Germany, and a New Phylogenetic Analysis of Extant and Extinct

We describe an Oligocene newt specimen from western Germany that has gone practically unnoticed in the literature despite having been housed in the Museum für Naturkunde (Ber-lin) for a century. It is referable to the coeval Chelotriton, but is unusually peramorphic; for many characters it is more peramorphic than all other caudates or even all other lissamphi-bians. Most noticeable are the position of the jaw joints far caudal to the occiput, the honey-combed sculpture on the maxilla, and the possible presence of a septomaxilla (which would be unique among salamandrids). Referral to a species would require a revision of the genus, but the specimen likely does not belong to the type species. A phylogenetic analysis of nonmolecular characters of Salamandridae, far larger than all predecessors, confirms the referral to Chelotriton. It further loosely associates the Oligocene Archaeotriton and the Miocene Carpathotriton with the extant Lissotriton, though the former may alternatively lie outside Pleurodelinae altogether. The Miocene? I. randeckensismay not belong to the extant Ichthyosaura. The Miocene “Triturus ” roehrsi is found neither with the extant Omma-totriton nor with Lissotriton, but inside an Asian/aquatic clade or, when geographic distribu