A systematic comparative description of extant turtle humeri, with comments on humerus disparity and evolution based on fossil comparisons.

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.

Saurodesmus robertsoni Seeley 1891-The oldest Scottish cynodont.

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.

The first occurrence of machimosaurid crocodylomorphs from the Oxfordian of south-central Poland provides new insights into the distribution of macrophagous teleosauroids.

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 Zalecze 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.

Natural external plastron mold of the Triassic turtle Proterochersis: An unusual mode of preservation.

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.

Bone abnormalities in the middle Anisian marine sauropsids from Winterswijk.

While the occurrence of skeletal pathologies in Middle Triassic marine reptiles has been poorly documented until now, massive accumulations of bone remains from the Germanic Basin provide the opportunities for documentation. Herein, we describe skeletal abnormalities in the Middle Triassic bone material from the Vossenveld Formation of Winterswijk, the Netherlands. The aim of the study is to distinguish in the studied bones pathologies resulting from malady or trauma and taphonomic alterations. Furthermore, an attempt was made to assess on how the pathologies also represent paleoecological data. Our survey led to the identification of one broken and healed bone, one case of abnormal coossification, and one case of posttraumatic fibro-osseous dysplasia. While the latter two pathologies give little insight into the ecology and function of the affected animals, the fractured dentary is attributed to Nothosaurus marchicus, a common sauropterygian macropredator. It proves that the individual survived long enough to heal, despite the injury hampering its hunting potential. One abnormally shaped humerus is interpreted as postmortem taphonomic deformation, emphasizing the necessity of utilization of detailed diagnostics to distinguish actual paleopathologies from nonbiological distortion.

An insight into cancer palaeobiology: does the Mesozoic neoplasm support tissue organization field theory of tumorigenesis?

BACKGROUND: Neoplasms are common across the animal kingdom and seem to be a feature plesiomorphic for metazoans, related with an increase in somatic complexity. The fossil record of cancer complements our knowledge of the origin of neoplasms and vulnerability of various vertebrate taxa. Here, we document the first undoubted record of primary malignant bone tumour in a Mesozoic non-amniote. The diagnosed osteosarcoma developed in the vertebral intercentrum of a temnospondyl amphibian, Metoposaurus krasiejowensis from the Krasiejów locality, southern Poland. RESULTS: A wide array of data collected from gross anatomy, histology, and microstructure of the affected intercentrum reveals the tumour growth dynamics and pathophysiological aspects of the neoplasm formation on the histological level. The pathological process almost exclusively pertains to the periosteal part of the bone composed from a highly vascularised tissue with lamellar matrix. The unorganised arrangement of osteocyte lacunae observed in the tissue is characteristic for bone tissue types connected with static osteogenesis, and not for lamellar bone. The neoplastic bone mimics on the structural level the fast growing fibrolamellar bone, but on the histological level develops through a novel ossification type. The physiological process of bone remodelling inside the endochondral domain continued uninterrupted across the pathology of the periosteal part. CONCLUSIONS: Based on the results, we discuss our case study's consistence with the Tissue Organization Field Theory of tumorigenesis, which locates the causes of neoplastic transformations in disorders of tissue architecture.

Dinosaur senescence: a hadrosauroid with age-related diseases brings a new perspective of "old" dinosaurs.

Senile vertebrates are extremely rare in the fossil record, making their recognition difficult. Here we present the largest known representative of the Late Cretaceous hadrosauriform Gobihadros mongoliensis showing features of cessation of growth indicating attainment of the terminal size. Moreover, this is the first non-avian dinosaur with an age-related pathology recognized as primary calcium pyrophosphate deposition disease indicating its advanced age. Because senile dinosaurs are so rare and thus "senescence" in dinosaurs is unclear, we also propose a new unified definition of a senile dinosaur: an individual which achieved the terminal size as revealed by the presence of the external fundamental system and closed transcortical channels, has completely secondary remodeled weight-bearing bones and possesses non-traumatic, non-contagious bone pathologies correlated with advanced age.

New tomographic contribution to characterizing mesosaurid congenital scoliosis.

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.

Tuberculosis-like respiratory infection in 245-million-year-old marine reptile suggested by bone pathologies.

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.

Shell variability in the stem turtles Proterochersis spp.

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.

The oldest record of aquatic amniote congenital scoliosis.

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.

Homeotic shift at the dawn of the turtle evolution.

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.