Quantitative evidence for dimorphism suggests sexual selection in the maxillary caniniform process of Placerias hesternus.

Placerias hesternus, a Late Triassic dicynodont, is one of the last megafaunal synapsids of the Mesozoic. The species has a tusk-like projection on its maxillary bone, known as the caniniform process. This process has been hypothesized to be sexually dimorphic since the 1950s, however this claim has not been thoroughly investigated quantitatively. Here, we examined maxillae, premaxillae, quadrates, and fibulae from a single population from the Placerias Quarry in the Blue Mesa Member of the Chinle Formation, near St. Johns, Arizona, USA to determine if the caniniform process is dimorphic. We made a total of 25 measurements from the four bones and used a maximum likelihood framework to compare the fit of unimodal versus bimodal distributions for each set of measurements. Our results from complete maxillae reveal that the caniniform process has two distinct morphs, with a shorter and longer form. This interpretation is substantiated both by strong statistical support for bimodal distribution of caniniform lengths, and by clustering analysis that clearly distinguishes two morphs for the maxillae. Clustering analysis also shows support for potential dimorphism in the shape of the quadrate. However, no measurements from elements other than the maxilla have a strong likelihood of bimodal distribution. These results support the long-standing hypothesis that the caniniform in Placerias was dimorphic. Alternative explanations to sexual dimorphism that could account for the dimorphism among these fossils include the presence of juveniles in the sample or time-averaged sampling of a chronospecies, but both have been previously rejected for the Placerias Quarry population. The lack of strong dimorphism in non-maxilla elements and increased variation in caniniform length of the large-caniniform morph suggest that the caniniform is a secondary sexual trait, possibly used in intraspecific competition.

The osteology of Shuvosaurus inexpectatus, a shuvosaurid pseudosuchian from the Upper Triassic Post Quarry, Dockum Group of Texas, USA.

A vast array of pseudosuchian body plans evolved during the diversification of the group in the Triassic Period, but few can compare to the toothless, long-necked, and bipedal shuvosaurids. Members of this clade possess theropod-like character states mapped on top of more plesiomorphic pseudosuchian character states, complicating our understanding of the evolutionary history of the skeleton. One taxon in this clade, Shuvosaurus inexpectatus has been assigned to various theropod dinosaur groups based on a partial skull and referred material and its postcranium was assigned to a different taxon in Pseudosuchia. After the discovery of a skeleton of a shuvosaurid with a Shuvosaurus-like skull and a pseudosuchian postcranial skeleton, it became clear Shuvosaurus inexpectatus was a pseudosuchian. Nevertheless, a number of questions have arisen about what skeletal elements belonged to Shuvosaurus inexpectatus, the identification of skull bones, and the resulting implication for pseudosuchian evolution. Here, we detail the anatomy of the skeleton Shuvosaurus inexpectatus through a critical lens, parse out the bones that belong to the taxon or those that clearly do not or may not belong to the taxon, rediagnose the taxon based on these revisions, and compare the taxon to other archosaurs. We find that Shuvosaurus inexpectatus possesses similar anatomy to other shuvosaurids but parts of the skeleton of the taxon clarifies the anatomy of the group given that they are preserved in Shuvosaurus inexpectatus but not in others. Shuvosaurus inexpectatus is represented by at least 14 individuals from the West Texas Post Quarry (Adamanian holochronozone) and all Shuvosaurus inexpectatus skeletal material from the locality pertains to skeletally immature individuals. All of the skeletons are missing most of the neural arches, ribs, and most of the forelimb. We only recognize Shuvosaurus inexpectatus from the Post Quarry and all other material assigned to the taxon previously is better assigned to the broader group Shuvosauridae.

Braincase and neuroanatomy of the lagerpetid Dromomeron gregorii (Archosauria, Pterosauromorpha) with comments on the early evolution of the braincase and associated soft tissues in Avemetatarsalia.

The anatomy of the braincase and associated soft tissues of the lagerpetid Dromomeron gregorii (Archosauria: Avemetatarsalia) from the Late Triassic of the United States is here described. This corresponds to the first detailed description of cranial materials of Lagerpetidae, an enigmatic group of Late Triassic (c. 236-200 Million years ago) animals that are the closest known relatives of pterosaurs, the flying reptiles. The braincase of D. gregorii is characterized by the presence of an anteriorly elongated laterosphenoid and a postparietal, features observed in stem-archosaurs but that were still unknown in early members of the avian lineage of archosaurs. Using micro-computed tomography (CT-scan data), we present digital reconstructions of the brain and endosseous labyrinth of D. gregorii. The brain of D. gregorii exhibits a floccular lobe of the cerebellum that projects within the space of the semicircular canals. The semicircular canals are relatively large when compared to other archosauromorphs, with the anterior canal exhibiting a circular shape. These features of the sensory structures of D. gregorii are more similar to those of pterosaurs than to those of other early avemetatarsalians. In sum, the braincase anatomy of D. gregorii shows a combination of plesiomorphic and apomorphic features in the phylogenetic context of Archosauria and suggests that the still poorly understood early evolution of the braincase in avemetatarsalians is complex, with a scenario of independent acquisitions and losses of character states.

The osteology of the Late Triassic reptile Scleromochlus taylori from µCT data.

Scleromochlus taylori is one of the most enigmatic members of the herpetofauna from the Lossiemouth Sandstone Formation (Upper Triassic) of Elgin (Moray, Scotland). For many years it was thought to be closely related to pterosaurs and dinosaurs, but the anatomy of this animal is difficult to interpret because of the notoriously poor preservation of the six available specimens, which comprise void space in the sandstone after the bones were destroyed by diagenesis. Historically, these fossils have been studied using physical molds, which provide only incomplete, and potentially distorted, information. Due to these uncertainties, interpretations of the anatomy, phylogenetic relationships, and paleobiology of Scleromochlus taylori have remained contentious. Here, we use microcomputed tomographic (µCT) techniques to redescribe and illustrate the osteology of Scleromochlus in detail, building upon a short redescription of keystone features of the anatomy that we recently published. We digitally visualize, describe, and figure previously inaccessible-and thus unaltered-portions of its skeleton, as well as providing new observations on the exposed parts of each specimen. This work reveals many novel features of the skull, mandible, trunk, tail, girdles, forelimb, and hindlimb (particularly of the manus, femur, and pes), demonstrating that historic molding techniques failed, in some cases, to accurately capture the anatomy of Scleromochlus. Our review sheds light on some of the most controversial aspects of Scleromochlus morphology showing that this taxon retains plesiomorphic features of Avemetatarsalia in the postcranial skeleton, alongside a suite of synapomorphies diagnostic of pterosauromorphs (the broad clade of pterosaurs and taxa more closely related to them than dinosaurs), particularly one subgroup, the lagerpetids. Consistent with recent work, our updated phylogenetic analyses (Maximum Parsimony and Bayesian Inference) demonstrate that Scleromochlus taylori is an avemetatarsalian archosaur that is recovered firmly in an early diverging position within Pterosauromorpha, as a member of Lagerpetidae, thus shedding important information on the origin of pterosaurs, the first group of vertebrates to evolve powered flight.

A new archosauriform species from the Panchet Formation of India and the diversification of Proterosuchidae after the end-Permian mass extinction.

Proterosuchidae represents the oldest substantial diversification of Archosauromorpha and plays a key role in understanding the biotic recovery after the end-Permian mass extinction. Proterosuchidae was long treated as a wastebasket taxon, but recent revisions have reduced its taxonomic content to five valid species from the latest Permian of Russia and the earliest Triassic (Induan) of South Africa and China. In addition to these occurrences, several isolated proterosuchid bones have been reported from the Induan Panchet Formation of India for over 150 years. Following the re-study of historical specimens and newly collected material from this unit, we erect the new proterosuchid species Samsarasuchus pamelae, which is represented by most of the presacral vertebral column. We also describe cf. proterosuchid and proterosuchid cranial, girdle and limb bones that are not referred to Samsarasuchus pamelae. Phylogenetic analyses recovered Samsarasuchus pamelae within the new proterosuchid clade Chasmatosuchinae. The taxonomic diversity of Proterosuchidae is substantially expanded here, with at least 11 nominal species and several currently unnamed specimens, and a biogeographical range encompassing present-day South Africa, China, Russia, India, Brazil, Uruguay and Australia. This indicates a broader taxonomic, phylogenetic and biogeographic diversification of Proterosuchidae than previously thought in the aftermath of the end-Permian mass extinction.

New reptile shows dinosaurs and pterosaurs evolved among diverse precursors.

Dinosaurs and pterosaurs have remarkable diversity and disparity through most of the Mesozoic Era1-3. Soon after their origins, these reptiles diversified into a number of long-lived lineages, evolved unprecedented ecologies (for example, flying, large herbivorous forms) and spread across Pangaea4,5. Recent discoveries of dinosaur and pterosaur precursors6-10 demonstrated that these animals were also speciose and widespread, but those precursors have few if any well-preserved skulls, hands and associated skeletons11,12. Here we present a well-preserved partial skeleton (Upper Triassic, Brazil) of the new lagerpetid Venetoraptor gassenae gen. et sp. nov. that offers a more comprehensive look into the skull and ecology of one of these precursors. Its skull has a sharp, raptorial-like beak, preceding that of dinosaurs by around 80 million years, and a large hand with long, trenchant claws that firmly establishes the loss of obligatory quadrupedalism in these precursor lineages. Combining anatomical information of the new species with other dinosaur and pterosaur precursors shows that morphological disparity of precursors resembles that of Triassic pterosaurs and exceeds that of Triassic dinosaurs. Thus, the 'success' of pterosaurs and dinosaurs was a result of differential survival among a broader pool of ecomorphological variation. Our results show that the morphological diversity of ornithodirans started to flourish among early-diverging lineages and not only after the origins of dinosaurs and pterosaurs.

A diverse diapsid tooth assemblage from the Early Triassic (Driefontein locality, South Africa) records the recovery of diapsids following the end-Permian mass extinction.

Mass extinctions change the trajectory of evolution and restructure ecosystems. The largest mass extinction, the end-Permian, is a particularly interesting case due to the hypothesized delay in the recovery of global ecosystems, where total trophic level recovery is not thought to have occurred until 5-9 million years after the extinction event. Diapsids, especially archosauromorphs, play an important role in this recovery, filling niches left vacant by therapsids and anapsids. However, the nature of lineage and ecological diversification of diapsids is obscured by the limited number of continuous, well-dated stratigraphic sections at the Permian-Triassic boundary and continuing through the first half of the Triassic. The Karoo Basin of South Africa is one such record, and particularly the late Early Triassic (Olenekian) Driefontein locality fills this gap in the diapsid fossil record. We collected a total of 102 teeth of which 81 are identified as diapsids and the remaining 21 as identified as temnospondyls. From the sample, seven distinct tooth morphotypes of diapsids are recognized, six of which are new to the locality. We used a combination of linear measurements, 3D geomorphometrics, and nMDS ordination to compare these morphotypes and made inferences about their possible diets. Although the morphotypes are readily differentiated in nMDS, the overall morphological disparity is low, and we infer five morphotypes are faunivorous with the other two potentially omnivorous or piscivorous based on their morphological similarities with dentitions from extant diapsids, demonstrating an unsampled taxonomic and ecological diversity of diapsids in the Early Triassic based on teeth. Although ecological specialization at Driefontein may be low, it records a diversity of diapsid taxa, specifically of archosauromorph lineages.

The femora of Drepanosauromorpha (Reptilia: Diapsida): Implications for the functional evolution of the thigh of Sauropsida.

The femora of diapsids have undergone morphological changes related to shifts in postural and locomotor modes, such as the transition from plesiomorphic amniote and diapsid taxa to the apomorphic conditions related to a more erect posture within Archosauriformes. One remarkable clade of Triassic diapsids is the chameleon-like Drepanosauromorpha. This group is known from numerous articulated but heavily compressed skeletons that have the potential to further inform early reptile femoral evolution. For the first time, we describe the three-dimensional osteology of the femora of Drepanosauromorpha, based on undistorted fossils from the Upper Triassic Chinle Formation and Dockum Group of North America. We identify apomorphies and a combination of character states that link these femora to those in crushed specimens of drepanosauromorphs and compare our sample with a range of amniote taxa. Several characteristics of drepanosauromorph femora, including a hemispherical proximal articular surface, prominent asymmetry in the proximodistal length of the tibial condyles, and a deep intercondylar sulcus, are plesiomorphies shared with early diapsids. The femora contrast with those of most diapsids in lacking a crest-like, distally tapering internal trochanter. They bear a ventrolaterally positioned tuberosity on the femoral shaft, resembling the fourth trochanter in Archosauriformes. The reduction of an internal trochanter parallels independent reductions in therapsids and archosauriforms. The presence of a ventrolaterally positioned trochanter is also similar to that of chameleonid squamates. Collectively, these features demonstrate a unique femoral morphology for drepanosauromorphs, and suggest an increased capacity for femoral adduction and protraction relative to most other Permo-Triassic diapsids.

Triassic stem caecilian supports dissorophoid origin of living amphibians.

Living amphibians (Lissamphibia) include frogs and salamanders (Batrachia) and the limbless worm-like caecilians (Gymnophiona). The estimated Palaeozoic era gymnophionan-batrachian molecular divergence1 suggests a major gap in the record of crown lissamphibians prior to their earliest fossil occurrences in the Triassic period2-6. Recent studies find a monophyletic Batrachia within dissorophoid temnospondyls7-10, but the absence of pre-Jurassic period caecilian fossils11,12 has made their relationships to batrachians and affinities to Palaeozoic tetrapods controversial1,8,13,14. Here we report the geologically oldest stem caecilian-a crown lissamphibian from the Late Triassic epoch of Arizona, USA-extending the caecilian record by around 35 million years. These fossils illuminate the tempo and mode of early caecilian morphological and functional evolution, demonstrating a delayed acquisition of musculoskeletal features associated with fossoriality in living caecilians, including the dual jaw closure mechanism15,16, reduced orbits17 and the tentacular organ18. The provenance of these fossils suggests a Pangaean equatorial origin for caecilians, implying that living caecilian biogeography reflects conserved aspects of caecilian function and physiology19, in combination with vicariance patterns driven by plate tectonics20. These fossils reveal a combination of features that is unique to caecilians alongside features that are shared with batrachian and dissorophoid temnospondyls, providing new and compelling evidence supporting a single origin of living amphibians within dissorophoid temnospondyls.

Scleromochlus and the early evolution of Pterosauromorpha.

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.

The dinosaurian femoral head experienced a morphogenetic shift from torsion to growth along the avian stem.

Significant evolutionary shifts in locomotor behaviour often involve comparatively subtle anatomical transitions. For dinosaurian and avian evolution, medial overhang of the proximal femur has been central to discussions. However, there is an apparent conflict with regard to the evolutionary origin of the dinosaurian femoral head, with neontological and palaeontological data suggesting seemingly incongruent hypotheses. To reconcile this, we reconstructed the evolutionary history of morphogenesis of the proximal end of the femur from early archosaurs to crown birds. Embryological comparison of living archosaurs (crocodylians and birds) suggests the acquisition of the greater overhang of the femoral head in dinosaurs results from additional growth of the proximal end in the medial-ward direction. On the other hand, the fossil record suggests that this overhang was acquired by torsion of the proximal end, which projected in a more rostral direction ancestrally. We reconcile this apparent conflict by inferring that the medial overhang of the dinosaur femoral head was initially acquired by torsion, which was then superseded by mediad growth. Details of anatomical shifts in fossil forms support this hypothesis, and their biomechanical implications are congruent with the general consensus regarding broader morpho-functional evolution on the avian stem.

Ontogenetic growth in the crania of Exaeretodon argentinus (Synapsida: Cynodontia) captures a dietary shift.

Background: An ontogenetic niche shift in vertebrates is a common occurrence where ecology shifts with morphological changes throughout growth. How ecology shifts over a vertebrate's lifetime is often reconstructed in extant species-by combining observational and skeletal data from growth series of the same species-because interactions between organisms and their environment can be observed directly. However, reconstructing shifts using extinct vertebrates is difficult and requires well-sampled growth series, specimens with relatively complete preservation, and easily observable skeletal traits associated with ecologies suspected to change throughout growth, such as diet. Methods: To reconstruct ecological changes throughout the growth of a stem-mammal, we describe changes associated with dietary ecology in a growth series of crania of the large-bodied (∼2 m in length) and herbivorous form, Exaeretodon argentinus (Cynodontia: Traversodontidae) from the Late Triassic Ischigualasto Formation, San Juan, Argentina. Nearly all specimens were deformed by taphonomic processes, so we reconstructed allometric slope using a generalized linear mixed effects model with distortion as a random effect. Results: Under a mixed effects model, we find that throughout growth, E. argentinus reduced the relative length of the palate, postcanine series, orbits, and basicranium, and expanded the relative length of the temporal region and the height of the zygomatic arch. The allometric relationship between the zygomatic arch and temporal region with the total length of the skull approximate the rate of growth for feeding musculature. Based on a higher allometric slope, the zygoma height is growing relatively faster than the length of the temporal region. The higher rate of change in the zygoma may suggest that smaller individuals had a crushing-dominated feeding style that transitioned into a chewing-dominated feeding style in larger individuals, suggesting a dietary shift from possible faunivory to a more plant-dominated diet. Dietary differentiation throughout development is further supported by an increase in sutural complexity and a shift in the orientation of microwear anisotropy between small and large individuals of E. argentinus. A developmental transition in the feeding ecology of E. argentinus is reflective of the reconstructed dietary transition across Gomphodontia, wherein the earliest-diverging species are inferred as omnivorous and the well-nested traversodontids are inferred as herbivorous, potentially suggesting that faunivory in immature individuals of the herbivorous Traversodontidae may be plesiomorphic for the clade.

Using Macro- and Microscale Preservation in Vertebrate Fossils as Predictors for Molecular Preservation in Fluvial Environments.

Exceptionally preserved fossils retain soft tissues and often the biomolecules that were present in an animal during its life. The majority of terrestrial vertebrate fossils are not traditionally considered exceptionally preserved, with fossils falling on a spectrum ranging from very well-preserved to poorly preserved when considering completeness, morphology and the presence of microstructures. Within this variability of anatomical preservation, high-quality macro-scale preservation (e.g., articulated skeletons) may not be reflected in molecular-scale preservation (i.e., biomolecules). Excavation of the Hayden Quarry (HQ; Chinle Formation, Ghost Ranch, NM, USA) has resulted in the recovery of thousands of fossilized vertebrate specimens. This has contributed greatly to our knowledge of early dinosaur evolution and paleoenvironmental conditions during the Late Triassic Period (~212 Ma). The number of specimens, completeness of skeletons and fidelity of osteohistological microstructures preserved in the bone all demonstrate the remarkable quality of the fossils preserved at this locality. Because the Hayden Quarry is an excellent example of good preservation in a fluvial environment, we have tested different fossil types (i.e., bone, tooth, coprolite) to examine the molecular preservation and overall taphonomy of the HQ to determine how different scales of preservation vary within a single locality. We used multiple high-resolution mass spectrometry techniques (TOF-SIMS, GC-MS, FT-ICR MS) to compare the fossils to unaltered bone from extant vertebrates, experimentally matured bone, and younger dinosaurian skeletal material from other fluvial environments. FT-ICR MS provides detailed molecular information about complex mixtures, and TOF-SIMS has high elemental spatial sensitivity. Using these techniques, we did not find convincing evidence of a molecular signal that can be confidently interpreted as endogenous, indicating that very good macro- and microscale preservation are not necessarily good predictors of molecular preservation.

Africa's oldest dinosaurs reveal early suppression of dinosaur distribution.

The vertebrate lineages that would shape Mesozoic and Cenozoic terrestrial ecosystems originated across Triassic Pangaea1-11. By the Late Triassic (Carnian stage, ~235 million years ago), cosmopolitan 'disaster faunas' (refs. 12-14) had given way to highly endemic assemblages12,13 on the supercontinent. Testing the tempo and mode of the establishment of this endemism is challenging-there were few geographic barriers to dispersal across Pangaea during the Late Triassic. Instead, palaeolatitudinal climate belts, and not continental boundaries, are proposed to have controlled distribution15-18. During this time of high endemism, dinosaurs began to disperse and thus offer an opportunity to test the timing and drivers of this biogeographic pattern. Increased sampling can test this prediction: if dinosaurs initially dispersed under palaeolatitudinal-driven endemism, then an assemblage similar to those of South America4,19-21 and India19,22-including the earliest dinosaurs-should be present in Carnian deposits in south-central Africa. Here we report a new Carnian assemblage from Zimbabwe that includes Africa's oldest definitive dinosaurs, including a nearly complete skeleton of the sauropodomorph Mbiresaurus raathi gen. et sp. nov. This assemblage resembles other dinosaur-bearing Carnian assemblages, suggesting that a similar vertebrate fauna ranged high-latitude austral Pangaea. The distribution of the first dinosaurs is correlated with palaeolatitude-linked climatic barriers, and dinosaurian dispersal to the rest of the supercontinent was delayed until these barriers relaxed, suggesting that climatic controls influenced the initial composition of the terrestrial faunas that persist to this day.

The developing bird pelvis passes through ancestral dinosaurian conditions.

Living birds (Aves) have bodies substantially modified from the ancestral reptilian condition. The avian pelvis in particular experienced major changes during the transition from early archosaurs to living birds1,2. This stepwise transformation is well documented by an excellent fossil record2-4; however, the ontogenetic alterations that underly it are less well understood. We used embryological imaging techniques to examine the morphogenesis of avian pelvic tissues in three dimensions, allowing direct comparison with the fossil record. Many ancestral dinosaurian features2 (for example, a forward-facing pubis, short ilium and pubic 'boot') are transiently present in the early morphogenesis of birds and arrive at their typical 'avian' form after transitioning through a prenatal developmental sequence that mirrors the phylogenetic sequence of character acquisition. We demonstrate quantitatively that avian pelvic ontogeny parallels the non-avian dinosaur-to-bird transition and provide evidence for phenotypic covariance within the pelvis that is conserved across Archosauria. The presence of ancestral states in avian embryos may stem from this conserved covariant relationship. In sum, our data provide evidence that the avian pelvis, whose early development has been little studied5-7, evolved through terminal addition-a mechanism8-10 whereby new apomorphic states are added to the end of a developmental sequence, resulting in expression8,11 of ancestral character states earlier in that sequence. The phenotypic integration we detected suggests a previously unrecognized mechanism for terminal addition and hints that retention of ancestral states in development is common during evolutionary transitions.

A new pseudosuchian archosaur, Mambawakale ruhuhu gen. et sp. nov., from the Middle Triassic Manda Beds of Tanzania.

The Manda Beds of southwest Tanzania have yielded key insights into the early evolutionary radiation of archosaurian reptiles. Many key archosaur specimens were collected from the Manda Beds in the 1930s and 1960s, but until recently, few of these had been formally published. Here, we describe an archosaur specimen collected in 1963 which has previously been referred to informally as Pallisteria angustimentum. We recognize this specimen as the type of a new taxon, Mambawakale ruhuhu gen. et sp. nov. The holotype and only known specimen of M. ruhuhu comprises a partial skull of large size (greater than 75 cm inferred length), lower jaws and fragments of the postcranium, including three anterior cervical vertebrae and a nearly complete left manus. Mambawakale ruhuhu is characterized by several cranial autapomorphies that allow it to be distinguished with confidence from all other Manda Beds archosaurs, with the possible exception of Stagonosuchus nyassicus for which comparisons are highly constrained due to very limited overlapping material. Phylogenetic analysis suggests that M. ruhuhu is an early diverging pseudosuchian, but more precise resolution is hampered by missing data. Mambawakale ruhuhu is one of the largest known pseudosuchians recovered to date from the Middle Triassic.

Histological analysis of ankylothecodonty in Silesauridae (Archosauria: Dinosauriformes) and its implications for the evolution of dinosaur tooth attachment.

Dinosaurs possess a form of tooth attachment wherein an unmineralized periodontal ligament suspends each tooth within a socket, similar to the condition in mammals and crocodylians. However, little information is known about tooth attachment and implantation in their close relatives, the silesaurids. We conducted a histological survey of several silesaurid taxa to determine the nature of tooth attachment in this phylogenetically and paleoecologically important group of archosaurs. Our histological data demonstrate that these early dinosauriforms do not exhibit the crocodilian/dinosaur condition of a permanent gomphosis, nor the rapid ankylosis that is plesiomorphic for amniotes. Instead, all sampled silesaurids exhibit delayed ankylosis, a condition in which teeth pass through a prolonged stage where the teeth are suspended in sockets by a periodontal ligament, followed by eventual mineralization and fusion of the tooth to the jaws. This suggests that tooth attachment in crocodylians and dinosaurs represent the further retention of an early ontogenetic stage compared to silesaurids, a paedomorphic trend that is mirrored in the evolution of synapsid tooth attachment. It also suggests that the dinosaur and crocodylian gomphosis was convergently acquired via heterochrony or, less likely, that the silesaurid condition represents a reversal to a plesiomorphic state. Moreover, if Silesauridae is nested within Ornithischia, a permanent gomphosis could be convergent between the two main dinosaur lineages, Ornithischia and Saurischia. These results demonstrate that dental characters in early archosaur phylogenies must be chosen and defined carefully, taking into account the relative duration of the different phases of dental ontogeny.

Osteology and relationships of Revueltosaurus callenderi (Archosauria: Suchia) from the Upper Triassic (Norian) Chinle Formation of Petrified Forest National Park, Arizona, United States.

Once known solely from dental material and thought to represent an early ornithischian dinosaur, the early-diverging pseudosuchian Revueltosaurus callenderi is described from a minimum of 12 skeletons from a monodominant bonebed in the upper part of the Chinle Formation of Arizona. This material includes nearly the entire skeleton and possesses a combination of plesiomorphic and derived character states that help clarify ingroup relationships within Pseudosuchia. A phylogenetic analysis recovers R. callenderi in a clade with Aetosauria and Acaenasuchus geoffreyi that is named Aetosauriformes. Key autapomorphies of R. callenderi include a skull that is longer than the femur, a complete carapace of dermal armor including paramedian and lateral rows, as well as ventral osteoderms, and a tail end sheathed in bone. Histology of the femur and associated osteoderms demonstrate that R. callenderi was slow growing and that the individuals from the bonebed were not young juveniles but had not ceased growing. A review of other material assigned to Revueltosaurus concludes that the genus cannot be adequately diagnosed based on the type materials of the three assigned species and that only R. callenderi can be confidently referred to Revueltosaurus.

Femoral specializations to locomotor habits in early archosauriforms.

The evolutionary history of archosaurs and their closest relatives is characterized by a wide diversity of locomotor modes, which has even been suggested as a pivotal aspect underlying the evolutionary success of dinosaurs vs. pseudosuchians across the Triassic-Jurassic transition. This locomotor diversity (e.g., more sprawling/erect; crouched/upright; quadrupedal/bipedal) led to several morphofunctional specializations of archosauriform limb bones that have been studied qualitatively as well as quantitatively through various linear morphometric studies. However, differences in locomotor habits have never been studied across the Triassic-Jurassic transition using 3D geometric morphometrics, which can relate how morphological features vary according to biological factors such as locomotor habit and body mass. Herein, we investigate morphological variation across a dataset of 72 femora from 36 different species of archosauriforms. First, we identify femoral head rotation, distal slope of the fourth trochanter, femoral curvature, and the angle between the lateral condyle and crista tibiofibularis as the main features varying between bipedal and quadrupedal taxa, all of these traits having a stronger locomotor signal than the lesser trochanter's proximal extent. We show a significant association between locomotor mode and phylogeny, but with the locomotor signal being stronger than the phylogenetic signal. This enables us to predict locomotor modes of some of the more ambiguous early archosauriforms without relying on the relationships between hindlimb and forelimb linear bone dimensions as in prior studies. Second, we highlight that the most important morphological variation is linked to the increase of body size, which impacts the width of the epiphyses and the roundness and proximodistal position of the fourth trochanter. Furthermore, we show that bipedal and quadrupedal archosauriforms have different allometric trajectories along the morphological variation in relation to body size. Finally, we demonstrate a covariation between locomotor mode and body size, with variations in femoral bowing (anteroposterior curvature) being more distinct among robust femora than gracile ones. We also identify a decoupling in fourth trochanter variation between locomotor mode (symmetrical to semi-pendant) and body size (sharp to rounded). Our results indicate a similar level of morphological disparity linked to a clear convergence in femoral robusticity between the two clades of archosauriforms (Pseudosuchia and Avemetatarsalia), emphasizing the importance of accounting for body size when studying their evolutionary history, as well as when studying the functional morphology of appendicular features. Determining how early archosauriform skeletal features were impacted by locomotor habits and body size also enables us to discuss the potential homoplasy of some phylogenetic characters used previously in cladistic analyses as well as when bipedalism evolved in the avemetatarsalian lineage. This study illuminates how the evolution of femoral morphology in early archosauriforms was functionally constrained by locomotor habit and body size, which should aid ongoing discussions about the early evolution of dinosaurs and the nature of their evolutionary "success" over pseudosuchians.