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BackgroundTurtle 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 Proterochersis porebensis (Proterochersidae, the sister group to all other known testudinatans) from the Late Triassic (Norian) of Germany and Poland.MethodsAll the available shell remains of Proterochersis robusta (13 specimens) and Proterochersis 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.ResultsProterochersis 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 Proterochersis porebensis allow estimation of its maximal carapace length at approximately 80 cm, while Proterochersis robusta appears to have reached lower maximal sizes.DiscussionThis 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.

Aetosaurs are armored basal archosaurs that played a significant role in land ecosystems during the Late Triassic (237–201 Ma). The polish species Stagonolepis olenkae Sulej, 2010 described from the Krasiejów locality (southern Poland) is one of the oldest known representatives of the group. Abundant and well-preserved material, including partially articulated specimens, allows a detailed description of the forelimbs in this species. The forelimbs of S. olenkae are the most similar to that of large aetosaurs like Desmatosuchus smalli, Desmatosuchus spurensis, Longosuchus meadei, Typothorax coccinarum or Stagonolepis robertsoni. Several characters recognized in the forelimbs of S. olenkae suggest its adaptation for scratch-digging. The most salient of these features are: short forearm, carpus, and hands, with the radius shorter than the humerus, carpus and manus shorter than the radius (excluding terminal phalanges); a prominent deltopectoral crest that extends distally on the humerus and a wide prominent entepicondyle, a long olecranon process with well-marked attachment of triceps muscle; hooked, laterally compressed, claw-like terminal phalanges with ornamentation of small pits (indicative of well-developed keratin sheaths). S. olenkae might have used its robust forelimbs to break through the compacted soil with its claws and proceed to dig in search of food in softened substrate with the shovel-like expansion at the tip of its snout. The entire forelimb of S. olenkae is covered by osteoderms, including the dorsal surface of the hand, which is unusual among aetosaurs and have not been noted for any species up to date.

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