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Predating Darwin's theory of evolution, the holotype of Saurodesmus robertsoni is a long-standing enigma. Found at the beginning of 1840s, the specimen is a damaged stylopodial bone over decades variably assigned to turtles, archosaurs, parareptiles, or synapsids, and currently nearly forgotten. We redescribe and re-assess that curious specimen as a femur and consider Saurodesmus robertsoni as a valid taxon of a derived cynodont (?Tritylodontidae). It shares with probainognathians more derived than Prozostrodon a mainly medially oriented lesser trochanter and with the clade reuniting tritylodontids, brasilodontids, and mammaliaforms (but excluding tritheledontids) the presence of a projected femoral head, offset from the long axis of the femoral shaft; a thin, plate-like greater trochanter; a distinct dorsal eminence proximal to the medial (tibial) condyle located close to the level of the long axis of the femoral shaft and almost in the middle of the width of the distal expansion; and a pocket-like fossa proximally to the medial (tibial) condyle. Saurodesmus robertsoni is most similar to tritylodontids, sharing at least with some forms: the relative mediolateral expansion of the proximal and distal regions of the femur, the general shape and development of the greater trochanter, the presence of a faint intertrochanteric crest separating the shallow intertrochanteric and adductor fossae, and the general outline of the distal region as observed dorsally and distally. This makes Saurodesmus robertsoni the first Triassic cynodont from Scotland and, possibly, one of the earliest representatives of tritylodontids and one of the latest non-mammaliaform cynodonts worldwide. Moreover, it highlights the need for revisiting historical problematic specimens, the identification of which could have been previously hampered by the lack of adequate comparative materials in the past. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Teleosauroid thalattosuchians were a clade of semi-aquatic crocodylomorphs that achieved a broad geographic distribution during the Mesozoic. While their fossils are well documented in Western European strata, our understanding of teleosauroids (and thalattosuchians in general) is notably poorer in Central-Eastern Europe, and from Poland in particular. Herein, we redescribe a teleosauroid rostrum (MZ VIII Vr-72) from middle Oxfordian strata of Załęcze Wielkie, in south-central Poland. Until now, the specimen has been largely encased in a block of limestone. After preparation, its rostral and dental morphology could be evaluated, showing the specimen to be a non-machimosaurin machimosaurid, similar in morphology to taxa Neosteneosaurus edwardsi and Proexochokefalos heberti. The well-preserved teeth enable us to study the specimen feeding ecology through the means of comparing its teeth to other teleosauroids through PCoA analysis. Comparisons with inferred closely related taxa suggest that the referred specimen was a macrophagous generalist. Notably, MZ VIII Vr-72 displays a prominent pathological distortion of the anterior rostrum, in the form of lateral bending. The pathology affects the nasal passage and tooth size and position, and is fully healed, indicating that, despite its macrophagous diet, it did not prevent the individual from food acquisition. [ABSTRACT FROM AUTHOR]

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]

Preservation of soft parts (collagen fibres, blood vessels and cells) in extinct vertebrates is rare and usually limited to fossilized bone and cartilage. Well-preserved coarse collagenous fibre bundles embedded in a mineralized matrix of tendons, as well as numerous hollow, tubular structures consistent morphologically with fibril bundles, blood vessels and associated cells, were identified in ossified tendons of Late Cretaceous ornithischians from North America and Central East Asia. Detailed, high-accuracy imaging, along with spectroscopic characterization of those fibrous structures and comparison with ossified tendons of modern-day turkeys, support the proposition that physiologically driven tendon ossification is common for avians and non-avian dinosaurs. The examined soft parts were preserved through the pathway of iron-induced crosslinking and alumino-silification, documenting a variety of pathways for the preservation of soft parts, depending on the burial environment. For the first time, the structure of dinosaur fossilized tendons is analysed in detail, revealing shared histogenetic principles with modern birds and the nature of preservation. [ABSTRACT FROM AUTHOR]

Despite nearly two centuries of intensive research of dicynodont diversity and distribution, the progress of the last two decades makes the early 21st century a dicynodont renaissance. Here we introduce Woznikella triradiata n. gen., n. sp., a Late Triassic European kannemeyeriiform with stahleckeriid affinities that may represent an early diverging lineage of that family, preceding its split into Placeriinae King, 1988 and Stahleckeriinae Lehman, 1961. Woznikella triradiata n. gen., n. sp. is distinguished from other dicynodonts by its autapomorphic, slender scapula with distinctly expanded dorsal and ventral parts, acromion directed anterodorsally, and inconspicuous scapular spine. Furthermore, we review reported global occurrences of Triassic dicynodonts, including fragmentary, indeterminate finds that may capture cryptic diversity, and analyze the biogeography of the Permian and Triassic Dicynodontia. According to our results, the region of southeastern Africa (Malawi, Mozambique, Namibia, South Africa, Tanzania, Zambia) throughout the Permian and Triassic served as a hotspot of dicynodont diversity and origin point for lineages migrating north and west. Multiple taxa independently migrated to the Americas and Eurasia, indicating open passages between the regions of Southern and Northern Hemisphere. Southern migrations from the Northern Hemisphere appear to be very rare, however. [ABSTRACT FROM AUTHOR]

The Middle Triassic remains a poorly understood time in the evolution of land vertebrates. Here, we report a new Ladinian-age vertebrate assemblage from Miedary (southern Poland). It consists of more than 20 taxa including fish (four species of Hybodontiformes, cf. Gyrolepis, Redfieldiiformes, 'Thelodus', Saurichthys, Serrolepis, Prohalecites, Ptychoceratodus), amphibians (Mastodonsaurus, Gerrothorax, Plagiosternum, chroniosuchian Bystrowiella), and reptiles (Owenettidae, Blezingeria, Nothosaurus, Tanystropheus, an additional, yet unidentified tanystropheid, the doswelliid Jaxtasuchus, and another archosauromorph, as well as eight archosauriform tooth morphotypes). Preliminary comparisons suggest biogeographic and environmental similarities with roughly contemporaneous localities known from the southwestern part of the Germanic Basin. Among differences in these two areas are the presence of a new armored archosauromorph and a surprising abundance of Tanystropheus remains in the new Polish site. Miedary is currently the richest source of three-dimensionally preserved Tanystropheus material in the world, which will be crucial for a better understanding of the preferred environment and lifestyle of this highly specialized reptile. [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]

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]

Duck‐billed dinosaurs (Hadrosauridae) were the most common ornithopods of the Late Cretaceous. Second only to sauropods and in many cases exceeding the sizes of the largest land mammals (such as indricotheres or proboscideans), they are among the largest terrestrial herbivores to have walked the Earth. Despite their gigantic size, diversity and abundance, their growth strategies remain poorly understood. Here, we examine the bone microstructure of several Mongolian hadrosauroids of varied adult sizes. The small and middle‐sized species have lines of arrested growth (LAGs). On the other hand, one of the largest duck‐billed dinosaurs, Saurolophus angustirostris, shows uninterrupted growth, comparable with other big hadrosaurs for which the lack of cyclical growth arrests was interpreted as a result of living in the polar region. Since both of the studied taxa inhabited warmer, continental, monsoon‐influenced environments of the Late Cretaceous Mongolia, we propose that the absence of LAGs is not a climatic‐driven condition but rather connected with the animal's size (i.e. ontogeny). Our results show that, like sauropods, hadrosaurs changed their growth dynamics from cyclical to continuous during their evolution, which made it possible for them to achieve comparable body sizes. [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]

The presence of a pathology in the vertebral column of the early Permian mesosaurid specimen ZPAL R VII/1, being one of the oldest amniotic occurrences of congenital scoliosis caused by a hemivertebra, was recently recognized. Here we provide CT data to further characterize the phenomenon. The affected hemivertebra is wedged (incarcerated) between the preceding and succeeding vertebrae. The neural canal is misshapen but continuous and the number of dorsal ribs on each side of the specimen corresponds with the number of the vertebrae, documenting its congenital (homeobox-related) derivation. [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]

Although Western Europe has yielded numerous Jurassic turtle taxa, several represented by cranial material or complete skeletons, the fossil record of the Jurassic turtles remains scarce to the north and east from Germany. Although some Late Jurassic testudinates were historically described from Poland, they were, thus far, represented by fragmentary remains that never were properly figured or described in detail. Therefore, very little is known about the mid‐Mesozoic diversity of turtles in that region of the continent. A new pancryptodiran turtle genus and species, Owadowia borsukbialynickae, is described from the uppermost Jurassic (Tithonian, ca. 148 Ma) carbonate sediments of the Kcynia Formation in Owadów‐Brzezinki Quarry, near Tomaszów Mazowiecki in central Poland. The lower jaw morphology and palaeoecological setting inhabited by the new genus and species, together with the trophic relationships of the Jurassic pancryptodiran turtles, are discussed in an attempt to determine the potential range of mode of life of O. borsukbialynickae. We propose that the new specimen belongs to a new durophagous pancryptodiran turtle taxon. O. borsukbialynickae might have spent considerable time in the marine environment and specialized on eating hard‐shelled invertebrates like bivalves and decapod crustaceans, common to that setting. [ABSTRACT FROM AUTHOR]

We report the first occurrence of congenital scoliosis in an early Permian aquatic parareptile, Stereosternum tumidum from Paraná state, Brazil. The spine malformation is caused by a congenital hemivertebra. These observations give insight into the biomechanical aspects of underwater locomotion in an axial skeleton-compromised aquatic amniote. This is the oldest record of a hemivertebra in an aquatic animal. [ABSTRACT FROM AUTHOR]

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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]

An almost complete dinosaur femur found in the Baynshire Formation (late Cenomanian to Santonian; Mongolia) in 1963 during the Polish–Mongolian Paleontological Expeditions is described here for the first time. The morphology of the femur and bone histology suggest that the specimen was a representative of the Titanosauriformes, the predominant sauropod clade during the Cretaceous. Bone tissue exclusive to this clade, modified laminar bone that indicates a reduced growth rate, is identified in the thin sections taken from the femur. Based on its bone histology, the already ca. 20-meter-long specimen appears to be a subadult. This specimen achieved a significantly larger size compared with other Titanosauriformes with modified lamellar bone at a similar growth stage. [ABSTRACT FROM AUTHOR]

The humerus is central for locomotion in turtles as quadrupedal animals. Osteological variation across testudine clades remains poorly documented. Here, we systematically describe the humerus anatomy for all major extant turtle clades based on 38 species representing the phylogenetic and ecological diversity of crown turtles. Three Late Triassic species of shelled stem turtles (Testudindata) are included to establish the plesiomorphic humerus morphology. Our work is based on 3D models, establishing a publicly available digital database. Previously defined terms for anatomical sides of the humerus (e.g., dorsal, ventral) are often not aligned with the respective body sides in turtles and other quadrupedal animals with sprawling gait. We propose alternative anatomical directional terms to simplify communication: radial and ulnar (the sides articulating with the radius/ulna), capitular (the side bearing the humeral head), and intertubercular (opposite to capitular surface). Turtle humeri show low morphological variation with exceptions concentrated in locomotory specialists. We propose 15 discrete characters to summarize osteological variation for future phylogenetic studies. Disparity analyses comparing non-shelled and shelled turtles indicate that the presence of the shell constrains humerus variation. Flippered aquatic turtles are released from this constraint and significantly increase overall disparity. Ontogenetic changes of turtle humeri are related to increased ossification and pronunciation of the proximal processes, the distal articulation areas, and the closure of the ectepicondylar groove to a foramen. Some turtle species retain juvenile features into adulthood and provide evidence for paedomorphic evolution. We review major changes of turtle humerus morphology throughout the evolution of its stem group., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

An absence of ancient archaeological and palaeontological evidence of pneumonia contrasts with its recognition in the more recent archaeological record. We document an apparent infection-mediated periosteal reaction affecting the dorsal ribs in a Middle Triassic eosauropterygian historically referred to as ' Proneusticosaurus ' silesiacus . High-resolution X-ray microtomography and histological studies of the pathologically altered ribs revealed the presence of a continuous solid periosteal reaction with multiple superficial blebs (protrusions) on the visceral surfaces of several ribs. Increased vascularization and uneven lines of arrested growth document that the pathology was the result of a multi-seasonal disease. While visceral surface localization of this periosteal reaction represents the earliest identified evidence for pneumonia, the blebs may have an additional implication: they have only been previously recognized in humans with tuberculosis (TB). Along with this diagnosis is the presence of focal vertebral erosions, parsimoniously compared to vertebral manifestation of TB in humans., Competing Interests: We declare we have no competing interests.

All derived turtles are characterized by one of the strongest reductions of the dorsal elements among Amniota, and have only 10 dorsal and eight cervical vertebrae. I demonstrate that the Late Triassic turtles, which represent successive stages of the shell evolution, indicate that the shift of the boundary between the cervical and dorsal sections of the vertebral column occurred over the course of several million years after the formation of complete carapace. The more generalized reptilian formula of at most seven cervicals and at least 11 dorsals is thus plesiomorphic for Testudinata. The morphological modifications associated with an anterior homeotic change of the first dorsal vertebra towards the last cervical vertebra in the Triassic turtles are partially recapitulated by the reduction of the first dorsal vertebra in crown-group Testudines, and they resemble the morphologies observed under laboratory conditions resulting from the experimental changes of Hox gene expression patterns. This homeotic shift hypothesis is supported by the, unique to turtles, restriction of Hox-5 expression domains, somitic precursors of scapula, and brachial plexus branches to the cervical region, by the number of the marginal scute-forming placodes, which was larger in the Triassic than in modern turtles, and by phylogenetic analyses., Competing Interests: I declare I have no competing interests.