Terrestrial tetrapods originated during the Middle Devonian, and the group rapidly diversified throughout the subsequent Carboniferous and Permian periods. Feeding in air rather than water is expected to require changes to tetrapod mandibular form and function. Previous biomechanical studies on jaw evolution, however, found that the increase in functional disparity lagged behind terrestrialisation by approximately 70 Myr, coinciding with the origin of amniotes and herbivory. We expand on a previous dataset composed primarily of non-amniote tetrapods to identify the drivers of this diversification, including representatives of all major amniote clades from the Permo-Carboniferous. First, we measured nine biomechanical traits from 111 tetrapod jaws in medial view. Second, we performed an Elliptical Fourier Analysis on 198 jaws in lateral view and 73 jaws in occlusal view. The first peak in jaw disparity, during the Pennsylvanian, occurs in carnivorous non-amniote tetrapods. However, the jaws of amniotes, particularly those inferred as herbivorous, are consistently more disparate than non-amniote tetrapods from the early Permian, especially in terms of jaw depth, symphysial length and force transmission. Functional and shape disparity of Palaeozoic tetrapod jaws follow a similar pattern that is explained by large-scale faunal turnovers and ecosystem structures.

Ankylosaurs were a group of heavily armored non-avian dinosaurs (Dinosauria, Ankylosauria), represented by a relatively abundant fossil record from the Cretaceous of North and South America. Their dermal skeleton was characterized by large osteoderms whose development and functional role have been largely investigated. However, interstitial small ossicles, forming between these osteoderms, have been far more overlooked and it remains unknown whether they were formed through the ossification of a preexisting fibrous matrix of connective tissue (i.e., metaplasia) or by a cell-induced differentiation of new fiber bundles followed by mineralization (i.e., neoplasia sensu (Zeitschrift für Wissenschaftliche Zoologie, 1858, 9, 147)). Here, we propose a hypothesis on the developmental origin of these small ossicles in the ankylosaurian Antarctopelta oliveroi using light microcopy, scanning electron microscopy and three-dimensional virtual histology through propagation phase-contrast synchrotron radiation micro-computed tomography (PPC-SRμCT). Ossicles are located in the dermis. They are composed of two layers: (1) a thin external layer, and (2) a thick basal plate, composed of collagen fiber bundles, which forms the main part of the ossicle. The external layer is made of a smooth, vitreous mineralized tissue that does not look like bone. The basal plate, however, is of osseous origin. In this basal plate, the collagen fiber bundles are organized in two orthogonal systems: one horizontal-observable in cross-sections-and one vertical-observable in the primary plane of sections sensu (Journal of Vertebrate Paleontology, 2004, 24, 874). The horizontal system is itself composed of successive layers of collagen fiber bundles arranged into an orthogonal plywood-like structure. The bundles of the vertical system radiate from the center of the ossicle at the level of the transition between the external layer and the basal plate and run towards the periphery of the basal plate. Their thickness increases from the center of the ossicle towards its periphery. Numerous bundles of the vertical system form thin threads that interweave and penetrate within the thick bundles of the horizontal system. Our new data suggest that the ossicles were at least partially formed by metaplasia, that is, through the ossification of a preexisting fibrous matrix of connective tissue. This process was probably supplemented by a cell-induced differentiation of new fiber bundles laid down prior to their incorporation into the fibrous system and its mineralization. This process looks more akin to neoplasia sensu (Zeitschrift für Wissenschaftliche Zoologie, 1858, 9, 147) than to metaplasia. Consequently, metaplastic and neoplastic processes may coexist in these ossicles with a possible differential expression during ontogeny.

Turtles have high shape variation of their mandibles, likely reflecting adaptations to a broad variety of food items and ingestion strategies. Here, we compare functional disparity measured by biomechanical proxies and character disparity measured by discrete morphological characters. Functional and character disparities vary between clades and ecological groups and are thus decoupled. Comparisons with cranial disparity also indicate decoupled patterns within the turtle skull. Exploration of mandibular patterns reveals that several biomechanical configurations or character state combinations can lead to the same feeding type (i.e., convergence) or that high functional disparity can be achieved at a low exhaustion of character state combinations (e.g., cryptodires). Dietary specialists show larger functional disparity than generalists, but the phylogenetically widespread generalist ecology leads to high character disparity signals in the ecotype. Whereas character disparity generally shows high phylogenetic signal, functional disparity patterns correspond to dietary specializations, which may occur convergently across different groups. Despite this, individual functional measurements have overlapping ranges across ecogroups and do not always conform to biomechanical expectations. Jaw opening and closing biomechanical advantages model trade-offs between force transmission and opening/closing speeds, and turtles show a variety of combinations of values that we try to synthesize into several "jaw types". Closing mechanical advantage shows that turtles retain high levels of force transmission at the anterior jaw end compared with other groups (e.g., pseudosuchians). This can possibly be explained as an evolutionary adaptation to retain high bite forces at small head sizes.

Among terrestrial tetrapods, the origin of herbivory marked a key evolutionary event that allowed for the evolution of modern terrestrial ecosystems. A 100 Ma gap separates the oldest terrestrial tetrapods and the first undisputed herbivorous tetrapods. While four clades of early tetrapod herbivores are undisputed amniotes, the phylogenetic position of Diadectomorpha with respect to Amniota has long been controversial. Given that the origin of herbivory coincides with the oldest amniotes, and obligate herbivory is unknown within amphibians, this suggests that a key adaptation necessary to evolve obligate herbivory is unique to amniotes. Historically, phylogenetic analyses have found Diadectomorpha as the sister-group to amniotes, but recent analyses recover Diadectomorpha as sister-group to Synapsida, within Amniota. We tested whether diadectomorphs are amniotes by updating the most recent character–taxon matrix. Specifically, we added new characters from the lower jaw and added diadectomorph taxa, resulting in a dataset of 341 characters and 61 operational taxonomic units. We updated the description of five diadectomorph jaws using microcomputed tomography data. Our majority-rule consensus places Diadectomorpha as sister-group to Synapsida; other methods do not recover this relationship. We revise diadectomorph taxonomy, erecting a new species from the early Permian Bromacker locality, Germany, and a new genus to accommodate ‘Diadectes’ sanmiguelensis.

We report the first trionychid turtle skull from the Lance Formation (Maastrichtian) of Wyoming, USA, a relatively complete cranium of Axestemys infernalis. Included is an osteological and preliminary neuroanatomical description aided by a medical CTscan of the specimen. Comparison with other published crania of A. infernalis yielded considerable intraspecific variation. Notably, our specimen does not bear an ossified premaxilla, the postorbital extends ventrally and the maxilla-jugal suture is positioned posterior to the orbit. Based on this new anatomical data, both a parsimony and a Bayesian analysis of an updated character-taxon matrix of Trionychia have been run, which found A. infernalis as either outside Plastomenidae+Cyclanorbinae or as an early diverging stem-cyclanorbine. Moreover, for the first time a fossil trionychid braincase is described in detail.

A mandible of a Late Cretaceous sea turtle with affinities to Ctenochelys is reported from the Maastrichtian type area of the Netherlands. The triangular mandible has a well-developed symphyseal ridge surrounded on both sides by large, concave areas on the triturating surface. It represents the first potential occurrence of Ctenochelys from the Maastrichtian type area. This finding increases the diversity of the turtle fauna known from the Maastrichtian type area.

Examples of photoluminescence (PL) are being reported with increasing frequency in a wide range of organisms from diverse ecosystems. However, the chemical basis of this PL remains poorly defined, and our understanding of its potential ecological function is still superficial. Among mammals, recent analyses have identified free-base porphyrins as the compounds responsible for the reddish ultraviolet-induced photoluminescence (UV-PL) observed in the pelage of springhares and hedgehogs. However, the localization of the pigments within the hair largely remains to be determined. Here, we use photoluminescence multispectral imaging emission and excitation spectroscopy to detect, map, and characterize porphyrinic compounds in skin appendages in situ. We also document new cases of mammalian UV-PL caused by free-base porphyrins in distantly related species. Spatial distribution of the UV-PL is strongly suggestive of an endogenous origin of the porphyrinic compounds. We argue that reddish UV-PL is predominantly observed in crepuscular and nocturnal mammals because porphyrins are photodegradable. Consequently, this phenomenon may not have a specific function in intra- or interspecific communication but rather represents a byproduct of potentially widespread physiological processes.

Turtles are a charismatic reptile group with a peculiar body plan, which most notably includes the shell. Anatomists have often focused descriptive efforts on the shell and other strongly derived body parts, such as the akinetic skull, or the cervical vertebrae. Other parts of turtle osteology, like the girdles, limbs, and mandibles, are documented with less rigor and detail. The mandible is the primary skeletal element involved in food acquisition and initial food processing of turtles, and its features are thus likely linked to feeding ecology. In addition, the mandible of turtles is composed of up to seven bones (sometimes fused to as little as three) and has thus anatomical complexity that may be insightful for systematic purposes and phylogenetic research. Despite apparent complexity and diversity to the mandible of turtles, this anatomical system has not been systematically studied, not even in search of characters that might improve phylogenetic resolution. Here, we describe the mandibular osteology for all major subclades of extant turtles with the help of digitally dissected 3D models derived from high-resolution computed tomography (μCT) scans of 70 extant species. We provide 31 fully segmented mandibles, as well as 3D models of the mandibular musculature, innervation, and arterial circulation of the cryptodire Dermatemys mawii. We synthesize observed variation into 51 morphological characters, which we optimize onto a molecular phylogeny. This analysis shows some mandibular characters to have high systematic value, whereas others are highly homoplastic and may underlie ecological influences or other factors invoking variation.

The early Permian Richards Spur locality of Oklahoma has produced abundant material of numerous terrestrial fossil tetrapods, including various “microsaurs,” several of which are considered to belong to the clade Recumbirostra. We present a new partial skull of the recumbirostran “microsaur” Nannaroter mckinziei; through computed tomography (CT) analysis of both this new specimen and the holotype, we provide an updated description of the taxon. This new description provides novel information regarding several regions that could not be examined previously due to either being absent in the holotype or difficult to access. This includes missing and obscured aspects of the skull roof, braincase, lower jaw, and the palatal region. Furthermore, the new information obtained from this description was used to update phylogenetic character codings of Nannaroter, and a revised phylogenetic analysis was conducted. The results of this updated analysis are congruent with those of other recent phylogenetic analyses of recumbirostran “microsaurs.” This new information adds to the ever-growing body of early tetrapod CT data, which has been, and will continue to be, important in revealing details regarding early tetrapod anatomy, interrelationships, paleoecology, and evolution.

Therocephalia is one of the major therapsid clades and ranges from the middle Permian to Middle Triassic. The earliest therocephalians were large-bodied predators whose fossils are common in middle Permian rocks of South Africa, but have received little study. Here we present a redescription of the skull of the early therocephalian Lycosuchus based on a specimen from the middle Permian Tapinocephalus Assemblage Zone of the South African Karoo Basin. By using a computed tomographic (CT) reconstruction of this specimen, we describe for the first time several endocranial characters of this taxon including a highly ramified maxillary canal and the inner ear, which is characterized by a lengthened lateral semicircular canal, a feature previously only known from the anomodont Kawingasaurus among non-mammalian therapsids, and the presence of a cochlear recess, so far only known within Therocephalia from the highly specialized Triassic taxon Microgomphodon. We also provide new insights into patterns of tooth replacement in lycosuchids, which have proven controversial for this taxon. Craniodental characters generally support the placement of Lycosuchus as the most basal taxon in therocephalian phylogeny. The morphology of the maxillary canal and inner ear reveal a mosaic of features indicating a complex history of character acquisition and loss in Therocephalia, comparable to that of cynodonts.

The cave deposits of the Lower Permian Richards Spur locality in Oklahoma, USA, have produced an incredible number of terrestrial tetrapod taxa, many of which are currently only known from this locality. One of the many recent taxa to be described from the locality was the small lanthanosuchoid parareptile Feeserpeton oklahomensis. Represented by a well-preserved, near complete skull, F. oklahomensis would have been a small predatory reptile, likely preying upon arthropods, and contributes to the extensive tetrapod fauna that was present at Richards Spur. New computed tomography data of the holotype and only specimen has allowed us to visualize and describe previously obscured and inaccessible anatomy of this taxon. These areas include the mandibular ramus, the palate, the sphenethmoid, the epipterygoids, and the braincase. Furthermore, this new anatomical information allowed formerly unknown character codings to be updated, thus we also performed new phylogenetic analyses that incorporated this new information. The results of these updated phylogenetic analyses are very similar to those of past studies, with F. oklahomensis being found as the sister taxon to all other lanthanosuchoids.

The morphology and histology of long bones are important tools for understanding tetrapod locomotion and growth. In mammals, the epiphyses of long bones ossify from a secondary centre of ossificati ...