Background. Turtle shells tend to exhibit frequent and substantial variability, both in bone and scute layout. Aside from secondary changes, caused by diseases, parasites, and trauma, this variability appears to be inherent and result from stochastic or externally-induced flaws of developmental programs. It is, thus, expected to be present in fossil turtle species at least as prominently, as in modern populations. Descriptions of variability and ontogeny are, however, rare for fossil turtles, mainly due to rarity, incompleteness, damage, and post-mortem deformation of their remains. This paper is an attempt at description and interpretation of external shell variability in representatives of the oldest true turtles, Proterochersis robusta and P. porebensis (Proterochersidae, the sister group to all other known testudinatans) from the Late Triassic (Norian) of Germany and Poland. Methods. All the available shell remains of Proterochersis robusta (13 specimens) and P. porebensis (275 specimens) were studied morphologically in order to identify any ontogenetic changes, intraspecific variability, sexual dimorphism, and shell abnormalities. To test the inferred sexual dimorphism, shape analyses were performed for two regions (gular and anal) of the plastron. Results. Proterochersis spp. exhibits large shell variability, and at least some of the observed changes seem to be correlated with ontogeny (growth of gulars, extragulars, caudals, and marginals, disappearance of middorsal keel on the carapace). Several specimens show abnormal layout of scute sulci, several others unusual morphologies of vertebral scute areas, one has an additional pair of plastral scutes, and one extraordinarily pronounced, likely pathological, growth rings on the carapace. Both species are represented in a wide spectrum of sizes, from hatchlings to old, mature individuals. The largest fragmentary specimens of P. porebensis allow estimation of its maximal carapace length at approximately 80 cm, while P. robusta appears to have reached lower maximal sizes. Discussion. This is the second contribution describing variability among numerous specimens of Triassic turtles, and the first to show evidence of unambiguous shell abnormalities. Presented data supplement the sparse knowledge of shell scute development in the earliest turtles and suggest that at least some aspects of the developmental programs governing scute development were already similar in the Late Triassic to these of modern forms.

Pleurosternon bullockii is a turtle from the Early Cretaceous of Europe known from numerous postcranial remains. Only one skull has so far been referred to the species. Pleurosternon bullockii belongs to a group of turtles called pleurosternids, which is thought to include several poorly known taxa from the Late Jurassic and Early Cretaceous of Europe and North America. Pleurosternids and baenids, a group of North American turtles that lived from the Late Cretaceous to the Eocene, define a clade called Paracryptodira. Additionally, Paracryptodira likely includes compsemydids, and, potentially, helochelydrids. Character support for Paracryptodira is relatively weak, and many global phylogenetic studies fail to support paracryptodiran monophyly altogether. Proposed paracryptodiran synapomorphies are largely cranial, despite the poor characterization of pleurosternid cranial material. In addition to their questionable monophyly, the global position of paracryptodires is debated. Early studies suggest crown-turtle affinities, but most phylogenies find them as stem-turtles, irrespective of their monophyly. Here, we document the cranial osteology of Pleurosternon bullockii with the use of three-dimensional models derived from segmenting high-resolution X-ray micro-computed tomography (CT) scans. Pleurosternon bullockii has a primitive basipterygoid region of the skull, but a cryptodirelike acustico-jugular region. A surprising number of similarities with pleurodires exist, particularly in the laterally expanded external process of the pterygoid and in the posterior orbital wall. Our observations constitute an important step toward a phylogenetic reevaluation of Paracryptodira.

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