New data on the crown proteid Bishara backa from the Upper Cretaceous (Bostobe Formation) of Kazakhstan: implications for early evolution and palaeobiogeography of proteidae

Abstract

ABSTRACT We provide a detailed redescription of all available material for the geologically oldest proteid salamander Bishara backa from the Upper Cretaceous (Santonian – lower Campanian) Bostobe Formation in Kazakhstan, consisting an atlantal centrum (neotype of Bishara backa) and a trunk vertebra. Bishara backa is characterised by the following combination of characters: large, dorsoventrally compressed and almost flat anterior cotyles of atlas enclosing an angle about 230°; anterior cotylar surfaces are continuous medially across lateral surfaces of small, dorsally placed intercotylar tubercle; ventrolateral ridges are alar-like; atlantal transverse processes are bicipitate; oblique anterior alar processes and lateral fossae on atlas present; transverse processes on trunk vertebra are bicipitate and divergent; alar processes on trunk vertebra are prominent; anterodorsal ridge on trunk vertebra is present; neural arch of trunk vertebra with high neural spine and grooves on its posterior face; two large subcentral foramina on ventral surface of the centrum. Our phylogenetic analyses finds Bishara backa as a crown proteid and a basal member of Eurasian clade of crown proteids (Bishara+Proteus+Mioproteus). We propose a model of proteid evolution that includes three main episodes of radiations with the last two taking place during the Late Cretaceous and the Palaeocene – Eocene thermal maximum.