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Impressions of vertebrate bodies or their parts, such as trace fossils and natural molds of bones, are a valuable source of information about ancient faunas which may supplement the standard fossil record based on skeletal elements. Whereas trace fossils of animal activity are relatively common and actively studied within the field of ichnology, and natural impressions of internal or external surfaces are a frequent preservation mode in fossil invertebrates, natural molds of bones are comparatively rare and less extensively documented and discussed. Among them, internal molds (steinkerns) of turtle shells are a relatively well-known form of preservation, but the mechanisms and taphonomic prerequisites leading to their formation are poorly studied. External shell molds are even less represented in the literature. Herein, we describe a historic specimen of a natural external turtle plastron mold from the Triassic (Norian) Löwenstein Formation of Germany-a formation which also yielded a number of turtle steinkerns. The specimen is significant not only because it represents an unusual form of preservation, but also due to its remarkably large size and the presence of a potential shell pathology. Although it was initially interpreted as Proterochersis sp., the recent progress in the knowledge of proterochersid turtles leading to an increase in the number of known taxa within that group allows us to verify that assessment. We confirm that the specimen is morphologically consistent with the genus and tentatively identify it as Proterochersis robusta, the only representative of that genus from the Löwenstein Formation. We note, however, that its size exceeds the size observed thus far in Proterochersis robusta and fits within the range of Proterochersis porebensis from the Grabowa Formation of Poland. The marks interpreted as shell pathology are morphologically consistent with Karethraichnus lakkos-an ichnotaxon interpreted as a trace of ectoparasites, such as leeches. This may support the previously proposed interpretation of Proterochersis spp. as a semiaquatic turtle. Moreover, if the identification is correct, the specimen may represent a very rare case of a negative preservation of a named ichnotaxon. Finally, we discuss the taphonomy of the Löwenstein Formation turtles in comparison with other Triassic turtle-yielding formations which show no potential for the preservation of internal or external shell molds and propose a taphonomic model for the formation of such fossils., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Szczygielski et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Data on turtle limb bone histology and microstructure are spotty, especially for Mesozoic taxa, despite significant progress made in recent years. Here we provide first detailed information on the stylopodia of the Late Triassic stem turtles Proganochelys quenstedtii Baur, 1887 from Switzerland and Proterochersis porebensis Szczygielski & Sulej, 2016 from Poland. In both taxa we observed large, internal medullary regions filled with endosteal trabeculae and poorly to moderately vascularized parallel-fibered (grading locally to lamellar) periosteal cortices. Primary vasculature is predominantly longitudinal, in Proterochersis porebensis locally with radial inclination. In large specimens, secondary remodeling is significant in the deeper cortex, but it neither completely obliterates the primary tissue nor reaches the external surface of the bone in either taxon. Comparison of histological data, limb morphology, shell and limb lengths as well as proportions reveal differences in growth patterns between the taxa: Proganochelys quenstedtii seems to grow faster during early life stages than Proterochersis porebensis and attained distinctly larger body sizes earlier in ontogeny, even though the asymptotic body size is roughly the same for both species. Overall, the histological and microstructural characteristics of stylopodial bones of Triassic turtles more closely resemble those of more recent representatives of that group than earlier stem turtles. [ABSTRACT FROM AUTHOR]

The turtle locomotor system is heavily modified owing to the severe impact of development of the shell on the body plan of these reptiles. Although limb and girdle osteology of the earliest, Triassic turtles is relatively well understood in general, the exact impacts of variability, ontogeny and preservation (e.g. deformation) on the observed morphologies have rarely been considered in detail. Here, we describe in detail and document the osteology and intraspecific variability of the limbs and girdles of Proterochersis spp. the basalmost true turtles (Testudinata). We also provide a synthesis of currently available data and detailed comparisons with other Triassic stem turtles to gain a better understanding of the diagnostic value of the early turtle appendicular skeleton and to pave the way for future biomechanical and functional studies. Our data suggest that Proterochersis spp. could be at least partly aquatic and could change their preferred habitat during ontogeny, with larger (and, presumably, older) specimens presenting more characters suggestive of a more terrestrial environment. [ABSTRACT FROM AUTHOR]

Shell suture obliteration (ankylosis) was exceptionally frequent in the earliest turtles, in contrast to post-Triassic taxa. Since modern turtles grow mostly along sutures, early ankylosis in Triassic taxa is intriguing. The Triassic turtle Proterochersis porebensis Szczygielski & Sulej, 2016 is known from numerous specimens, allowing observation of shell microstructure changes during ontogeny. Shell ankylosis occurred seemingly randomly in individuals of variable size, including small and morphologically juvenile, and completely obscured the initial bony composition. We propose that this phenomenon in the Triassic turtles can be an effect of early evolutionary stages of shell histogenesis and physiological mechanisms still used in shell regeneration in modern species. We also describe some parallels between the unusual peripheral microstructure of another Triassic turtle, Waluchelys cavitesta Sterli, Martínez, Cerda & Apaldetti, 2020, and Proterochersis porebensis. Microstructural changes imply that Proterochersis porebensis could change habitat during ontogeny, small individuals appearing more aquatic and larger more terrestrial. [ABSTRACT FROM AUTHOR]

The material historically referred to Chelytherium obscurum is exceptional for a number of reasons. It is the first described Triassic turtle, the oldest testudinate among the British palaeontological collections, and it is one of the oldest testudinates in the world. Nevertheless, after its establishment in 1863, the taxon remained in a taxonomic limbo and was eventually almost forgotten. However, the recent reconsideration of proterochersid turtles, with improved understanding of their anatomy, allows reassessment of Chelytherium obscurum as a member of the Proterochersidae, subjectively synonymous with Proterochersis robusta. Despite nomenclatural priority, in accordance with the suggestions of previous authors, the name Proterochersis robusta is endorsed to be upheld. [ABSTRACT FROM AUTHOR]

Marine reptiles from the Upper Jurassic of Central Europe are rare and often fragmentary, which hinders their precise taxonomic identification and their placement in a palaeobiogeographic context. Recent fieldwork in the Kimmeridgian of Krzyżanowice, Poland, a locality known from turtle remains originally discovered in the 1960s, has reportedly provided additional fossils thought to indicate the presence of a more diverse marine reptile assemblage, including giant pliosaurids, plesiosauroids, and thalattosuchians. Based on its taxonomic composition, the marine tetrapod fauna from Krzyżanowice was argued to represent part of the "Matyja-Wierzbowski Line"--a newly proposed palaeobiogeographic belt comprising faunal components transitional between those of the Boreal and Mediterranean marine provinces. Here, we provide a detailed re-description of the marine reptile material from Krzyżanowice and reassess its taxonomy. The turtle remains are proposed to represent a "plesiochelyid" thalassochelydian (Craspedochelys? sp.) and the plesiosauroid vertebral centrum likely belongs to a cryptoclidid. However, qualitative assessment and quantitative analysis of the jaws originally referred to the colossal pliosaurid Pliosaurus clearly demonstrate a metriorhynchid thalattosuchian affinity. Furthermore, these metriorhynchid jaws were likely found at a different, currently indeterminate, locality. A tooth crown previously identified as belonging to the thalattosuchian Machimosaurus is here considered to represent an indeterminate vertebrate. The revised taxonomy of the marine reptiles from Krzyżanowice, as well as the uncertain provenance of the metriorhynchid specimen reported from the locality, cast doubt on the palaeobiogeographic significance of the assemblage. [ABSTRACT FROM AUTHOR]

The shell of the oldest true turtle (Testudinata) branch (Proterochersidae) from the Late Triassic (Norian) of Poland and Germany was built in its anterior and posterior part from an osteodermal mosaic which developed several million years after the plastron, neurals and costal bones. We provide the most detailed description of the shell composition in proterochersids to date, together with a review of the shell composition in other Triassic pantestudinates. A scenario of early evolution of the turtle shell is proposed based on new data, and the possible adaptive meaning of the observed evolutionary changes is discussed. These observations are consistent with the trend of shell simplification previously reported in turtles. Several aspects of proterochersid shell anatomy are intermediate between Odontochelys semitestacea and more derived turtles, supporting their stem phylogenetic position. Three additional ossifications were sutured to xiphiplastra and pelvis in Proterochersis spp. and at least in some individuals the nuchal bone was paired. The peripherals, suprapygals, and pygal bone are most likely to be of osteodermal origin and homologous to the proterochersid shell mosaic. [ABSTRACT FROM AUTHOR]

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 Proterochersis 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 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. 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 Proterochersis porebensis allow estimation of its maximal carapace length at approximately 80 cm, while Proterochersis 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. [ABSTRACT FROM AUTHOR]

A recently discovered Norian outcrop in Poręba, Poland, has yielded numerous well-preserved turtle remains. These, together with historical materials from Germany, enabled the identification of two new proterochersid taxa: Proterochersis porebensis sp. nov. from Poland and Keuperotesta limendorsa gen. et sp. nov. from Germany. Moreover, two problematic taxa, Proterochersis intermedia Fraas, 1913; and Murrhardtia staeschei Karl & Tichy, 2000; are shown to be conspecific with Proterochersis robusta Fraas, 1913. New diagnoses for the family Proterochersidae Nopcsa, 1923 and all included taxa are provided. Proterochersids are of great importance to turtle phylogeny because of their age (they comprise the oldest fully shelled turtle species known to date) and their still-debated phylogenetic position (classically they are considered the basalmost Pleurodira, but in some analyses they are placed on the stem of Testudinata). Newly discovered plesiomorphic aspects of the anatomy of these three species included in the phylogenetic analysis demonstrate that they are not only the oldest, but are also the most basal fully shelled turtles. The unique features of their shell (e.g. the first thoracic rib unreduced and costal-bearing, and the osseous contact between the carapace and sacral vertebra) are of special significance for future research of the earliest stages of turtle shell evolution. This is the first contribution focused solely on the new, phylogenetically informative, and important characters of this group since the initial description of P. robusta more than 100 years ago. [ABSTRACT FROM AUTHOR]

Since 1990, several localities within the Keuper (upper Middle to Upper Triassic) strata in southern Poland have yielded remains of numerous terrestrial vertebrate species. Here we report a new Upper Triassic vertebrate assemblage from the rediscovered Kocury locality. An incomplete theropod dinosaur fibula named Velocipes guerichi described in 1932 was found there. The site was then forgotten and not explored until our excavations began in 2012, that yielded material of a lungfish, a proterochersid turtle, and a new typothoracin aetosaur Kocurypelta silvestris gen. et sp. nov. The new taxon is characterized by autapomorphies of the maxilla: an elongated edentulous posterior portion longer than 80% of the posterior maxillary process, a short medial shelf restricted to the posterior portion of the bone, an anteriorly unroofed maxillary accessory cavity, and lack of a distinct groove for choanal recess on the anteromedial surface of the bone. These new finds improve our knowledge on the vertebrate diversity of the Germanic Basin in the Late Triassic, evidencing the presence of yet unrecognized taxa. Additionally, the partial cranial aetosaur material emphasizes the issues with the aetosaurian taxonomy that is focused mostly on the osteoderm morphology. [ABSTRACT FROM AUTHOR]