Aging the oldest turtles: the placodont affinities of Priscochelys hegnabrunnensis

Priscochelys hegnabrunnensis, a fragmentary piece of armour shell from the Muschelkalk of Germany (Upper Triassic) with few diagnostic morphological features, was recently proposed to represent the oldest known stem turtle. As such, the specimen is of high importance because it shifts the date of the first appearance of turtles back about 20 Ma, which equals about 10% of the total stratigraphic range of the group. In this paper, I present new morphologic, histologic and neutron tomographic (NT) data that relate to the microstructure of the bone of the specimen itself. In opposition to the previous morphologic descriptions, P. hegnabrunnensis was found to share several distinctive features (i.e. bone sutures congruent with scute sulci, absence of a diploe structure with interior cancellous bone, thin vascular canals radiating outwards from distinct centres in each field and rugose ventral bone surface texture consisting of mineralised fibre bundles) with cyamodontoid placodonts (Diapsida: Sauropterygia) and fewer with stem turtles (i.e. depth of sulci). Two aspects that were previously thought to be relevant for the assignment to the turtle stem (conical scutes and presence of foramina) are argued to be of dubious value. P. hegnabrunnensis is proposed to represent a fragmentary piece of cyamodontoid armour consisting of fused conical plates herein. The specimen is not a part of the turtle stem and thus does not represent the oldest turtle. Accordingly, P. hegnabrunnensis does not shorten the ghost lineage to the potential sister group of turtles.