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The stem lineage of Archosauria is populated by a diverse fossil record that remains notably understudied relative to the crown clade. Prominent among these specimens is a beautifully preserved skull of the early mid-Triassic rhynchosaur Mesosuchus browni [Iziko South African Museum (SAM) 6536], whose phylogenetic position has considerable influence on patterns of pan-archosaurian cranial evolution. We used high-resolution, micro-computed tomography to re-examine the anatomy of this specimen, building on previous studies that were either limited to external observations or restricted to the braincase. A digital segmentation of the cranial elements and primary neurovascular canals of SAM-PK-6536 allows for expanded character scoring and constitutes a foundation for future comparative insights. Our data support the phylogenetically oldest instance of a pneumatized maxilla in a pan-archosaur, bringing the record of antorbital pneumatization into closer alignment with that of the neurocranium. The nasal cavity and primary palate of Mesosuchus includes a complex septomaxilla, a novel element anterior to the vomer, and is likely to have supported a well-developed vomeronasal system. The evolution of this system is discussed in terms of both phylogenetic pattern and how the skeletal architecture of Mesosuchus and other fossils could inform the signalling dynamics that pattern the vomeronasal system during development. [ABSTRACT FROM AUTHOR]

Some of the earliest members of the archosaur-lineage (i.e., non-archosauriform archosauromorphs) are characterised by an extremely elongated neck. Recent fossil discoveries from the Guanling Formation (Middle Triassic) of southern China have revealed a dramatic increase in the known ecomorphological diversity of these extremely long-necked archosauromorphs, including the fully marine and viviparous Dinocephalosaurus orientalis. These recent discoveries merit a reinvestigation of enigmatic Triassic diapsid fossils from contemporaneous European deposits housed in historical collections. Here, we provide a redescription of Trachelosaurus fischeri, represented by a single, disarticulated specimen first described in 1918. Due to its unique morphology, which includes short, bifurcating cervical ribs, and a high presacral vertebral count, this taxon has been referred to either as a "protorosaurian" archosauromorph or a sauropterygian. Our revision clearly shows that Trachelosaurus represents the first unambiguous Dinocephalosaurus-like archosauromorph known from outside the Guanling Formation. Our finding has important systematic implications. Trachelosauridae Abel, 1919 represents the senior synonym for the recently identified Dinocephalosauridae Spiekman, Fraser and Scheyer, 2021. Based on our phylogenetic analyses, which employ two extensive datasets, we also corroborate previous findings that tanystropheids and trachelosaurids represent two families within a larger monophyletic group among non-crocopodan archosauromorphs, which is here named Tanysauria (clade nov.). Trachelosauridae is minimally composed of Trachelosaurus fischeri, Dinocephalosaurus orientalis, Pectodens zhenyuensis, and Austronaga minuta, but one of our analyses also found a probably taxonomically broader clade that may also include Gracilicollum latens and Fuyuansaurus acutirostris. Trachelosaurus fischeri considerably expands the known spatial and temporal range of Trachelosauridae to the earliest Anisian and the Central European Basin. Our findings add to the growing evidence for the presence of a diverse group of fully marine reptiles during the Middle Triassic among Tanysauria. These trachelosaurids possess flipper-like limbs, high vertebral counts, and elongate necks, thus superficially resembling long-necked Jurassic and Cretaceous plesiosaurs in some regards. [ABSTRACT FROM AUTHOR]

Although the brain fills nearly the entire cranial cavity in birds, it can occupy a small portion of it in crocodilians. The lack of data regarding the volumetric correspondence between the brain and the cranial cavity hampers thorough assessments of the degree of encephalization in non‐neornithean dinosaurs and other extinct archosaurs and, consequently, informed inferences regarding their cognitive capacities. Existing data suggest that, across extant archosaurs, the degree of endocranial doming and the volume of intracranial nonneural components are inversely related. We build upon this information to develop an equation relating these two anatomical features in non‐neornithean dinosaurs and other extinct archosaurs. We rely on measurements of the endocast doming and brain‐to‐endocranial cavity (BEC) index in extant relatives of non‐neornithean dinosaurs, namely, the crurotarsans Caiman crocodilus, Crocodylus niloticus, and Crocodylus porosus; the paleognaths Struthio camelus and Apteryx mantelli; and the fowl Macrocephalon maleo, Gallus gallus, Meleagris gallopavo, Phasianus colchicus, and Anas platyrhynchos. Applying the equation to representative endocasts from major clades of dinosaurs, we found that BEC varies from about 0.6 in ceratopsians and thyreophorans to around 0.7 in ornithopods, pachycephalosaurians, sauropods, and theropods. We, therefore, warn against the use of a catch‐all value, like 0.5, and instead encourage refinement in the adoption of BEC across archosaurs.Illustration of the endocranial doming, which can provide a proxy for the brain‐to‐endocranial cavity index, using the endocast of an adult of the iguanodont dinosaur Proa valdearinnoensis as an example. [ABSTRACT FROM AUTHOR]

During the Late Triassic and Early Jurassic, the area around Bristol and South Wales was an archipelago of islands occupied by diverse small-sized tetrapods. The largest of these palaeo-islands was Mendip Island, now forming the Mendip Hills, and the location of some famous fossiliferous sites. These sites have not been described in detail before, and we present new data on three of them. Highcroft has yielded only sparse remains of rhynchocephalians, and Batscombe famously the gliding reptile Kuehneosuchus latissimus. Emborough yielded the richest fauna of the three, abundant pseudosuchians including crocodylomorphs as well as the gliding reptile Kuehneosaurus latus , rare trilophosaurs, a probable thalattosaur, rhynchocephalians, and the mammal Kuehneotherium. These include some of the last known taxa of clades that died out in the end-Triassic mass extinction. We report a new taxon of sphenosuchid crocodylomorph similar to Saltoposuchus and a find of Pachystropheus , an aquatic reptile shared with Holwell and the bedded Rhaetian at Blue Anchor Point, Aust and Westbury Garden Cliff. The discovery of a fish vertebra strengthens the model of Emborough fissure filling in a marginal marine location. The Emborough fauna differs from coeval assemblages from Cromhall, Tytherington and Ruthin in the scarcity of sphenodontians and the absence or great rarity of procolophonids as well as the abundance of kuehneosaurids and crocodylomorphs. [ABSTRACT FROM AUTHOR]

The lack of any pterosaur living descendants creates gaps in the knowledge of the biology of this group, including its cervical biomechanics, which makes it difficult to understand their posture and life habits. To mitigate part of this issue, we reconstructed the cervical osteology and arthrology of three pterosaurs, allowing us to make inferences about the position of the neck of these animals at rest. We used scans of threedimensionally preserved cervical series of Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri for the reconstructions, thus representing different lineages. For the recognition of ligaments, joint cartilages, and levels of overlapping of the zygapophyses, we applied the Extant Phylogenetic Bracket method, based on various extant birds and on Caiman latirostris. We inferred that pterosaur intervertebral joints were probably covered by a thin layer of synovial cartilage whose thickness varied along the neck, being thicker in the posterior region. Ignoring this cartilage can affect reconstructions. According to the vertebral angulation, their neck was slightly sinuous when in rest position. Our analyses also indicate that pterosaurs had segmented and supra-segmented articular cervical ligaments, which could confer stabilization, execute passive forces on the neck and store elastic energy. [ABSTRACT FROM AUTHOR]

Early archosaurs included a range of ecological types—from predators to herbivores and from terrestrial to semi-aquatic forms. Members of the family Ornithosuchidae, traditionally considered active predators, occupy a special position. Their characteristic jaw apparatus structure and the morphology of their postcranial skeleton suggest the development of a unique ecological type among archosaurs, namely, a medium-sized hyperanisodont macrophagous predator. At the same time, some analogs can be noted between ornithosuchids, saber-toothed therapsids, and mammals. [ABSTRACT FROM AUTHOR]

Thalattosuchia (Early Jurassic to Early Cretaceous) and Dyrosauridea (Late Cretaceous to Early Eocene) are crocodylomorph archosaurs which diversified in fluvial and marine environments and endured extinction events (i.e. Jurassic–Cretaceous boundary for Thalattosuchia; Cretaceous–Palaeogene for Dyrosauridea). Their postcrania remain globally undervalued in anatomical descriptions and diagnoses, shrouding the locomotive adaptations that possibly underpinned their radiations and longevity. We thoroughly surveyed the postcranial morphology of Dyrosauridea and Thalattosuchia, recreated their girdles in three‐dimensions using tens of high‐precisions 3D scans, and analysed their shape using geometric morphometrics. Dyrosauridea and Thalattosuchia have clearly distinct postcrania, even when found within similar environments, suggesting the existence of clade‐specific features limiting the strength of evolutionary convergence. Moreover, the range of postcranial morphologies evolved by dyrosaurids and thalattosuchians is large compared to extant crocodylians, making the latter unsatisfactory functional analogues for every group of extinct crocodylomorphs. Our work reveals the previously unsuspected potential of postcranial anatomy as an abundant source of phylogenetic and taxonomic characters to assess the relationships within Crocodylomorpha. Incorporation of postcranial anatomy therefore appears crucial to fully assess the ecology, disparity, and relationships of crocodylomorphs. [ABSTRACT FROM AUTHOR]

Understanding the evolution of the earliest avemetatarsalian (bird-line) archosaurs and inferring the morphology of the last common ancestor of Archosauria are hampered by a poor fossil record in critical temporal intervals. Here we describe an early-diverging avemetatarsalian from the ?Earliest Late Triassic (~235 Ma) 'basal Isalo II'/Makay Formation of Madagascar, which helps bridge these gaps. This taxon, Mambachiton fiandohana gen. et sp. nov. , is represented by well-preserved postcranial material and possibly a postfrontal bone. Features of the neck region include anteroposteriorly elongated vertebrae with laterally expanded dorsal ends of the neural spines with three pairs of osteoderms per cervical vertebra, lying dorsal to those vertebrae. Inclusion of Mambachiton in a phylogenetic analysis of archosauromorphs recovers it at the base of Avemetatarsalia, outside of the aphanosaur + ornithodiran clade. This new specimen indicates that osteoderms were present in the earliest avemetatarsalians, but were lost in more crownward lineages. The plesiomorphic morphology of the taxon also underscores the difficulty of identifying early avemetatarsalians from incomplete skeletons. This early-diverging avemetatarsalian occurring together with a lagerpetid and silesaurid in the 'basal Isalo II'/Makay Formation of Madagascar documents the co-occurrence of multiple non-dinosaurian avemetatarsalian clades in Gondwana near the Middle–Late Triassic transition. Translated abstract (Malagasy and French) is provided in the Supplementary information. [ABSTRACT FROM AUTHOR]

Crocodylomorpha is the stem-lineage of modern crocodylians and the only pseudosuchian (i.e. crocodylian-line archosaurs) clade that survived the Triassic–Jurassic mass extinction event. Its earliest members, the non-crocodyliform crocodylomorphs, also known as 'sphenosuchians', were terrestrial and mostly small-bodied (<2 m long), although some large-bodied forms are known. Saltoposuchus connectens is one of the first described crocodylomorph species but it remains poorly studied, in part due to its contentious taxonomy. Here, all referred Saltoposuchus specimens are described in detail for the first time and its taxonomy is revised, with additional taxonomic implications for the British crocodylomorph Terrestrisuchus gracilis and the coelophysoid theropod Procompsognathus triassicus. Saltoposuchus connectens is clearly distinguished from Terrestrisuchus gracilis based on both cranial and postcranial features. The phylogenetic analysis finds that Saltoposuchus connectens , Terrestrisuchus gracilis , and Litargosuchus leptorhynchus form a clade of gracile, long-legged crocodylomorphs, identified as Saltoposuchidae Crush 1984. A histological section of a femur reveals highly vascularized fibrolamellar tissue in the second-largest specimen of Saltoposuchus connectens (SMNS 12596), indicating sustained high growth rates. A similar pattern was previously observed in Terrestrisuchus sp. contrasting with slower growth rates in the crocodylomorph Hesperosuchus agilis. These findings suggest that saltoposuchids had a high resting metabolic rate and active lifestyle. [ABSTRACT FROM AUTHOR]

The lack of any pterosaur living descendants creates gaps in the knowledge of the biology of this group, including its cervical biomechanics, which makes it difficult to understand their posture and life habits. To mitigate part of this issue, we reconstructed the cervical osteology and arthrology of three pterosaurs, allowing us to make inferences about the position of the neck of these animals at rest. We used scans of three-dimensionally preserved cervical series of Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri for the reconstructions, thus representing different lineages. For the recognition of ligaments, joint cartilages, and levels of overlapping of the zygapophyses, we applied the Extant Phylogenetic Bracket method, based on various extant birds and on Caiman latirostris. We inferred that pterosaur intervertebral joints were probably covered by a thin layer of synovial cartilage whose thickness varied along the neck, being thicker in the posterior region. Ignoring this cartilage can affect reconstructions. According to the vertebral angulation, their neck was slightly sinuous when in rest position. Our analyses also indicate that pterosaurs had segmented and supra-segmented articular cervical ligaments, which could confer stabilization, execute passive forces on the neck and store elastic energy.

Ornithosuchids are a Late Triassic pseudosuchian archosaur group, consisting of four species (three from South America, and one from Scotland). All of them have triangular skulls with a protruding premaxilla, large nostrils, an extensive diastema in their narrow snout, a short jaw that does not reach the anterior end of the skull, and serrated posteriorly curved teeth. For this clade, carnivorous and scavenger habits have been previously proposed. Within the Ornithosuchidae, Riojasuchus tenuisceps (from Argentina) has the most morphologically extreme characteristics. Based on CT scans of the preserved skulls we generated a 3D model, and over this, we estimated the volumes of the adductors and abductor muscles and the force exerted by each. From these data we built the finite element model and measured the bite force (1.8–2.3 kN). Lateral, tractive, and torsional forces were applied to the end of the snout to evaluate the structural response of the skull during feeding. The results show that R. tenuisceps could resist tractive and torsional stresses better than lateral stress. Additionally, we analysed the peculiar morphological characteristics of the skull and their functional implications. We observed that the upper and lower dental rows were laterally separated from each other, preventing the generation of a cutting line during occlusion, and therefore, R. tenuisceps would have fed on small-sized prey that it could swallow whole. The curved premaxilla and the short mandible would not allow it to bite with the tip of the snout (ruling out the scavenging hypothesis), but were instead more adequate to capturing prey suspended in a fluid. This set of results allows us to propose that R. tenuisceps could have had a zoophagous diet and a wading habit, being able to feed on fish, amphibians, or any small animals that they could catch from the shoreline.

Ornithosuchids are a Late Triassic pseudosuchian archosaur group, consisting of four species (three from South America, and one from Scotland). All of them have triangular skulls with a protruding premaxilla, large nostrils, an extensive diastema in their narrow snout, a short jaw that does not reach the anterior end of the skull, and serrated posteriorly curved teeth. For this clade, carnivorous and scavenger habits have been previously proposed. Within the Ornithosuchidae, Riojasuchus tenuisceps (from Argentina) has the most morphologically extreme characteristics. Based on CT scans of the preserved skulls we generated a 3D model, and over this, we estimated the volumes of the adductors and abductor muscles and the force exerted by each. From these data we built the finite element model and measured the bite force (1.8–2.3 kN). Lateral, tractive, and torsional forces were applied to the end of the snout to evaluate the structural response of the skull during feeding. The results show that R. tenuisceps could resist tractive and torsional stresses better than lateral stress. Additionally, we analysed the peculiar morphological characteristics of the skull and their functional implications. We observed that the upper and lower dental rows were laterally separated from each other, preventing the generation of a cutting line during occlusion, and therefore, R. tenuisceps would have fed on small-sized prey that it could swallow whole. The curved premaxilla and the short mandible would not allow it to bite with the tip of the snout (ruling out the scavenging hypothesis), but were instead more adequate to capturing prey suspended in a fluid. This set of results allows us to propose that R. tenuisceps could have had a zoophagous diet and a wading habit, being able to feed on fish, amphibians, or any small animals that they could catch from the shoreline. [ABSTRACT FROM AUTHOR]

The independent origins of middle ears in testudinates, lepidosaurs, and archosaurs in the Triassic led to lineage-specific developments in their auditory epithelia. In comparison to the inferred ancestral state, little changed in testudinates, but archosaurs and most lepidosaurs evolved longer, differentiated auditory papillae. In archosaurs, sensory hair cells specialized across and along the papillae, resulting in mainly sensory tall hair cells and mainly mechanically active short hair cells. Crocodilians have 5 mm-long papillae, but relatively low upper frequency limits at around 4 kHz. Avian papillae are mostly 3 to 5 mm in length, being shorter in small species; in owls they exceptionally reach almost 12 mm and have an upper limit of above 10 kHz. Lepidosaurs retained the ancestral papilla as a low-frequency responsive area with one type of hair cell, but most added newly evolved areas (of max. 2 mm length) consisting of oppositely oriented hair cells responding to frequencies above 1 kHz. Initially, these areas flanked both ends of the ancestral area and were redundant in their responses to sound. The evolution of specific configurations in most families eliminated this redundancy, but the paths taken resulted in diverse anatomies that show high degrees of family specificity. Despite these differences, the high-frequency hearing limit of lepidosaurs rarely exceeds 5 kHz. [ABSTRACT FROM AUTHOR]

Crocodilians (alligators, crocodiles, and gharials) are the closet living relatives to birds and, as such, represent a key clade to understand the evolution of the avian brain. However, many aspects of crocodilian neurobiology remain unknown. In this paper, we address an important knowledge gap as there are no published studies of cerebellar connections in any crocodilian species. We used injections of retrograde tracers into the cerebellum of the American alligator (Alligator mississippiensis) to describe for the first time the origin of climbing and mossy fiber inputs. We found that inputs to the cerebellum in the American alligator are similar to those of other nonavian reptiles and birds. Retrograde labeled cells were found in the spinal cord, inferior olive, reticular formation, vestibular and cerebellar nuclei, as well as in nucleus ruber and surrounding tegmentum. Additionally, we found no retrogradely labeled cells in the anterior rhombencephalon which suggest that, like other nonavian reptiles, crocodilians may lack pontine nuclei. Similar to birds and other nonavian reptiles, we found inputs to the cerebellum from the pretectal nucleus lentiformis mesencephali. Additionally, we found retrogradely labeled neurons in two nuclei in the pretectum: the nucleus circularis and the interstitial nucleus of the posterior commissure. These pretectal projections have not been described in any other nonavian reptile to date, but they do resemble projections from the nucleus spiriformis medialis of birds. Our results show that many inputs to the cerebellum are highly conserved among sauropsids and that extensive pretectal inputs to the cerebellum are not exclusive to the avian brain. Finally, we suggest that the pontine nuclei of birds are an evolutionary novelty that may have evolved after the last common ancestor between birds and crocodilians, and may represent an intriguing case of convergent evolution with mammals. [ABSTRACT FROM AUTHOR]

Birds and crocodylians are the only remaining members of Archosauria (ruling reptiles) and they exhibit major differences in posture and gait, which are polar opposites in terms of locomotor strategies. Their broader lineages (Avemetatarsalia and Pseudosuchia) evolved a multitude of locomotor modes in the Triassic and Jurassic periods, including several occurrences of bipedalism. The exact timings and frequencies of bipedal origins within archosaurs, and thus their ancestral capabilities, are contentious. It is often suggested that archosaurs ancestrally exhibited some form of bipedalism. Euparkeria capensis is a central taxon for the investigation of locomotion in archosaurs due to its phylogenetic position and intermediate skeletal morphology, and is argued to be representative of facultative bipedalism in this group. However, no studies to date have biomechanically tested if bipedality was feasible in Eupakeria. Here, we use musculoskeletal models and static simulations in its hindlimb to test the influences of body posture and muscle parameter estimation methods on locomotor potential. Our analyses show that the resulting negative pitching moments around the centre of mass were prohibitive to sustainable bipedality. We conclude that it is unlikely that Euparkeria was facultatively bipedal, and was probably quadrupedal, rendering the inference of ancestral bipedal abilities in Archosauria unlikely.

Characterized by an elongated snout, proterochampsids are carnivorous non-archosaur archosauriforms. The clade is endemic to South America and its fossil record extends from the early Carnian to the late Carnian/early Norian. Nesting close to Archosauria, it is a key clade for understanding the origin and evolution of archosaurian traits. Unfortunately, hind limb elements are usually poorly preserved for the group. Therefore, the hind limb anatomy of proterochampsids still lacks detailed descriptions. In the present study, we partially fill this gap with the description of a new proterochampsid represented by a complete and well-preserved hind limb. Stenoscelida aurantiacus gen. et sp. nov. was excavated from the late Carnian/early Norian (Late Triassic) beds of southern Brazil. A phylogenetic investigation recovers the new taxon as a non-rhadinosuchine proterochampsid. The species bears an unusual set of traits for the group, which provides clues on the evolutionary origins of some muscle attachment structures. For instance, the femur of Stenoscelida aurantiacus gen. et sp. nov. possesses an anterior trochanter and an anterolateral scar. So far, these features have not been reported in other non-archosaurian archosauriforms. Therefore, the new specimen indicates that some typical archosaurian features evolved earlier than previously thought. The taxon also carries additional uncommon features for proterochampsids, such as an iliofibularis tubercle on the anterior margin of the fibula and a vestigial phalanx in digit V. In sum, Stenoscelida aurantiacus has one of the best-preserved hind limbs within Proterochampsidae and sheds light on the polarization of important traits regarding the evolutionary context of Archosauria. [ABSTRACT FROM AUTHOR]

Understanding the evolution of the tetrapod brain is essential to trace the history of ecomorphological diversification of modern clades. While previous studies focused on the morphological transformation of the nervous system along the dinosaur–bird transition, little is known about the brain anatomy of archosauriformes and early archosaurs. Here, we describe the endocast of Euparkeria capensis, a small bodied, terrestrial archosauriform closely related to Archosauria, with the goal of resolving the current uncertainties surrounding the ancestral condition of the archosaurian nervous system. The endocast of Euparkeria is sigmoidal, with large olfactory bulbs, an expanded cerebral hemisphere and an elongated flocculus. We suggest that this pivotal taxon was an active predator with a remarkable olfactory acuity. Overall, the endocast of Euparkeria resembles the ones observed in phytosaurs, crocodilians and early dinosaurs, implying that modern crocodilians retain an archosaurian plesiomorphic brain morphology. [ABSTRACT FROM AUTHOR]

We present the pelvic and hindlimb musculature of the abelisaurid Skorpiovenator bustingorryi, constituting the most comprehensive muscle reconstruction to date in ceratosaur theropods. Using extant phylogenetic bracket method, we reconstructed 39 muscles that can commonly found in extant archosaurs. Through the identification of bone correlates, we recognized thigh and hindlimb muscles including knee extensors, m. iliofibularis, m. flexor tibialis externus, mm. caudofemorales, mm. puboischiofemorales, and crus muscles important in foot extension and flexion (e.g., m. tibialis anterior, mm. gastrocnemii). Also, autopodial intrinsic muscles were reconstructed whose function involve extension (m. extensor digiti 2-4), flexion (mm. flexor digitorum brevis superficialis), interdigital adduction (m. interosseus dorsalis) and abduction (m. interosseous plantaris, m. abductor 4). Abelisaurids like Skorpiovenator show a deep pre- and postacetabular blade of the ilia and enlarged cnemial crests, which would have helped increasing the moment arm of muscles related to hip flexion and hindlimb extension. Also, pedal muscles related to pronation were probably present but reduced (e.g., m. pronator profundus). Despite some gross differences in the autopodial morphology in extant outgroups (e.g., crocodilian metatarsus and avian tarsometatarsus), the present study allows us to hypothesize several pedal muscles in Skorpiovenator. These muscles would not be arranged in tendinous bundles as in Neornithes, but rather the condition would be similar to that of crocodilians with several layers formed by fleshy bellies on the plantar and dorsal aspects of the metatarsus. The musculature of Skorpiovenator is key for future studies concerning abelisaurid biomechanics, including the integration of functional morphology and ichnological data., (© 2024 American Association for Anatomy.)

Recent studies suggest that both stem- and crown-group Archosauria encompassed high ecological diversity during their initial Triassic radiation. We describe a new pseudosuchian archosaur, Benggwigwishingasuchus eremicarminis gen. et sp. nov., from the Anisian (Middle Triassic) Fossil Hill Member of the Favret Formation (Nevada, USA), a pelagic setting in the eastern Panthalassan Ocean characterized by the presence of abundant ammonoids and large-bodied ichthyosaurs. Coupled with archosauriforms from the eastern and western Tethys Ocean, Benggwigwishingasuchus reveals that pseudosuchians were also components of Panthalassan ocean coastal settings, establishing that the group occupied these habitats globally during the Middle Triassic. However, Benggwigwishingasuchus , Qianosuchus , and Ticinosuchus (two other pseudosuchians known from marine sediments) are not recovered in a monophyletic group, demonstrating that a nearshore marine lifestyle occurred widely across Archosauriformes during this time. Benggwigwishingasuchus is recovered as part of an expanded Poposauroidea, including several taxa (e.g. Mandasuchus , Mambawakalae ) from the Middle Triassic Manda Beds of Tanzania among its basally branching members. This implies a greater undiscovered diversity of poposauroids during the Early Triassic, and supports that the group, and pseudosuchians more broadly, diversified rapidly following the End-Permian mass extinction.

In this contribution we introduce a new Late Triassic archosaur, Incertovenator longicollum gen. et sp. nov., with an unusual combination of character states that are present in certain early avemetatarsalian and pseudosuchian archosaur clades. The holotype consists of a partial postcranial skeleton, preserving most of the axial skeleton and displaying a marked anteroposterior elongation in the cervical vertebrae. We include I. longicollum gen. et sp. nov. into one of the most comprehensive early archosaur phylogenetic data sets available, and recover it as either an early diverging avemetatarsalian, closely associated with the clade Aphanosauria and Ornithodira, or as an early diverging loricatan closely related to Mandasuchus tanyauchen in the most parsimonious trees. We further evaluate which alternative phylogenetic positions can I. longicollum gen. et sp. nov. take in the suboptimal trees, and determined which character states support those alternative positions in comparison with those of the unconstrained analysis. The analyses recover the new taxon in three main general phylogenetic placements within Archosauria, as well as one position outside this clade, highlighting widespread morphological evolutionary convergence towards neck elongation in several clades of Triassic archosauriforms.

The independent origins of middle ears in testudinates, lepidosaurs, and archosaurs in the Triassic led to lineage-specific developments in their auditory epithelia. In comparison to the inferred ancestral state, little changed in testudinates, but archosaurs and most lepidosaurs evolved longer, differentiated auditory papillae. In archosaurs, sensory hair cells specialized across and along the papillae, resulting in mainly sensory tall hair cells and mainly mechanically active short hair cells. Crocodilians have 5 mm-long papillae, but relatively low upper frequency limits at around 4 kHz. Avian papillae are mostly 3 to 5 mm in length, being shorter in small species; in owls they exceptionally reach almost 12 mm and have an upper limit of above 10 kHz. Lepidosaurs retained the ancestral papilla as a low-frequency responsive area with one type of hair cell, but most added newly evolved areas (of max. 2 mm length) consisting of oppositely oriented hair cells responding to frequencies above 1 kHz. Initially, these areas flanked both ends of the ancestral area and were redundant in their responses to sound. The evolution of specific configurations in most families eliminated this redundancy, but the paths taken resulted in diverse anatomies that show high degrees of family specificity. Despite these differences, the high-frequency hearing limit of lepidosaurs rarely exceeds 5 kHz.

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. [ABSTRACT FROM AUTHOR]

Faxinalipterus minimus was originally described as a purported pterosaur from the Late Triassic (early Norian) Caturrita Formation of southern Brazil. Its holotype comprises fragmentary postcranial elements, whereas a partial maxilla was referred to the species. The assignment of Faxinalipterus minimus to Pterosauria has been questioned by some studies, but the specimen has never been accessed in detail after its original description. Here we provide a reassessment of Faxinalipterus minimus after additional mechanical preparation of the holotype. Our interpretations on the identity of several bones differ from those of the original description, and we found no support favoring pterosaur affinities for the taxon. The maxilla previously referred to Faxinalipterus minimus is disassociated from this taxon and referred to a new putative pterosauromorph described here from a partial skull and fragmentary postcranial elements. Maehary bonapartei gen. et sp. nov. comes from the same fossiliferous site that yielded Faxinalipterus minimus, but the lack of overlapping bones hampers comparisons between the two taxa. Our phylogenetic analysis places Faxinalipterus minimus within Lagerpetidae and Maehary bonapartei gen. et sp. nov. as the earliest-diverging member of Pterosauromorpha. Furthermore, the peculiar morphology of the new taxon reveals a new dental morphotype for archosaurs, characterized by conical, unserrated crowns, with a pair of apicobasally oriented grooves. These two enigmatic archosaurs expand our knowledge on the Caturrita Formation fauna and reinforce the importance of its beds on the understanding of Late Triassic ecosystems. [ABSTRACT FROM AUTHOR]

Faxinalipterus minimus was originally described as a purported pterosaur from the Late Triassic (early Norian) Caturrita Formation of southern Brazil. Its holotype comprises fragmentary postcranial elements, whereas a partial maxilla was referred to the species. The assignment of Faxinalipterus minimus to Pterosauria has been questioned by some studies, but the specimen has never been accessed in detail after its original description. Here we provide a reassessment of Faxinalipterus minimus after additional mechanical preparation of the holotype. Our interpretations on the identity of several bones differ from those of the original description, and we found no support favoring pterosaur affinities for the taxon. The maxilla previously referred to Faxinalipterus minimus is disassociated from this taxon and referred to a new putative pterosauromorph described here from a partial skull and fragmentary postcranial elements. Maehary bonapartei gen. et sp. nov. comes from the same fossiliferous site that yielded Faxinalipterus minimus, but the lack of overlapping bones hampers comparisons between the two taxa. Our phylogenetic analysis places Faxinalipterus minimus within Lagerpetidae and Maehary bonapartei gen. et sp. nov. as the earliest-diverging member of Pterosauromorpha. Furthermore, the peculiar morphology of the new taxon reveals a new dental morphotype for archosaurs, characterized by conical, unserrated crowns, with a pair of apicobasally oriented grooves. These two enigmatic archosaurs expand our knowledge on the Caturrita Formation fauna and reinforce the importance of its beds on the understanding of Late Triassic ecosystems.

Based on the restudying of all materials on early archosaurs from the Early Triassic Donskaya Luka locality, the genus and species Scythosuchus basileus Sennikov, 1999 is considered as a junior synonym of Tsylmosuchus donensis Sennikov, 1990. A revised diagnosis of the genus Tsylmosuchus, which is provisionally attributed to the family Ctenosauriscidae, is given. Tsylmosuchus was lightly built and capable of fast locomotion. The position of its limbs was probably already partially parasagittal. [ABSTRACT FROM AUTHOR]

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.

Proa valdearinnoensis is a relatively large‐headed and stocky iguanodontian dinosaur from the latest Early Cretaceous of Spain. Its braincase is known from three specimens. Similar to that of other dinosaurs, it shows a mosaic ossification pattern in which most of the bones seem to have fused together indistinguishably while a few (frontoparietal, basioccipital) might have remained loosely attached. The endocasts of the three specimens are described based on CT data and digital reconstructions. They show unmistakable morphological similarities with the endocast of closely related taxa, such as Sirindhorna khoratensis (which is close in age but from Thailand). This supports a high conservatism of the endocranial cavity. The issue of volumetric correspondence between endocranial cavity and brain in dinosaurs is analyzed. Although a brain‐to‐endocranial cavity (BEC) index of 0.50 has been traditionally used, we employ instead 0.73. This is indeed the mid‐value between the situation in adults of Alligator mississippiensis and Gallus gallus, which are members of the extant bracketing taxa of dinosaurs (Crocodilia and Aves). We thence gauge the level of encephalization of P. valdearinnoensis through the calculation of the encephalization quotient (EQ), which remains valuable as a metric for assessing the degree of cognitive function in extinct taxa, especially those with fully ossified braincases like dinosaurs and other archosaurs. The EQ obtained for P. valdearinnoensis (3.611) suggests that this species was significantly more encephalized than most if not all extant nonavian, nonmammalian amniotes. Our work adds to the growing body of data concerning theoretical cognitive capabilities in dinosaurs and supports the idea that an increasing encephalization was fostered not only in theropods but also in parallel in the shorter‐lived lineage of ornithopods. P. valdearinnoensis was ill‐equipped to respond to theropod dinosaurs and possibly lived in groups as a strategy to mitigate the risk of being predated upon. We hypothesize that group‐living and protracted caring of juveniles in this and possibly many other iguanodontian ornithopods favored a degree of encephalization that was outstanding by reptile standards. [ABSTRACT FROM AUTHOR]

Birds and mammals are key elements of modern ecosystems, and many biologists explain their great success by their endothermy, or warm-bloodedness. New palaeontological discoveries point to the origins of endothermy in the Triassic, and that birds (archosaurs) and mammals (synapsids) likely acquired endothermy in parallel. Here, a further case is made, that the emergence of endothermy in a stepwise manner began in the Late Permian but accelerated in the Early Triassic. The trigger was the profound destruction wrought by the Permian-Triassic mass extinction (PTME). In the oceans, this was the beginning of the Mesozoic Marine Revolution (MMR), and a similar revolution occurred on land, termed here the Triassic Terrestrial Revolution (TTR). Among tetrapods, both synapsids and archosaurs survived into the Triassic, but numbers were heavily depleted. However, the survivors were marked by the acquisition of endothermy, as shown by bone histology, isotopic analyses, and the acquisition of insulating pelage. Both groups before the PTME had been sprawlers; after the event they adopted parasagittal (erect) gait. The new posture and the new physiology enabled both groups to compete in their ecosystems at a faster rate than before the PTME. The new world of the Triassic was characterised by a fast-paced arms race between synapsids and archosauromorphs in which the latter, as both dinosaurs and pterosaurs, initially prevailed. [Display omitted] • Birds and mammals likely acquired endothermy (warm-bloodedness) at the same time • This time was the Early Triassic, as suggested by phylogenetic macroevolutionary studies of both archosaurs and synapsids • Life remodelled itself most significantly in the aftermath of the devastating end-Permian mass extinction • The switchover happened in parallel with a shift in posture from sprawling to parasagittal in both major lineages • Cynodonts and avemetatarsalian archosaurs engaged in arms races through the Triassic, as their metabolic rates speeded up [ABSTRACT FROM AUTHOR]

The reconstruction of jaw muscles is critical in establishing potential cranial functions; however, myological studies of extinct groups that have no descendants are difficult to perform and test. This is particularly true for pterosaurs, a group of extinct flying reptiles that present a plethora of cranial morphologies, suggesting different functions and feeding habits. Here we present a first attempt to reconstruct the adductor musculature of the pterodactyloid skull in detail, using osteological correlates and the extant phylogenetic bracketing method. Using these reconstructions, we estimate bite force for nine selected species and investigate implications for potential dietary habits. [ABSTRACT FROM AUTHOR]

rnithosuchidae is one of the most enigmatic clades of Triassic pseudosuchians. The group is composed by three carnivorous species that were excavated from Upper Triassic beds of Scotland and Argentina. We describe the first ornithosuchid from the Upper Triassic sediments of Brazil and explore its phylogenetic affinities and implications for the evolution of the group. Dynamosuchus collisensis gen. et sp. nov. was found as the sister taxon of the Argentinean form Venaticosuchus rusconii. These relationships reject a potential endemic radiation of ornithosuchids from the Ischigualasto-Villa Unión Basin and would better support multiple diversification events. Our findings with ornithosuchids is consistent with the pattern reported for proterochampsid and erpetosuchid archosauriforms from Ischigualasto-Villa Unión and the Paraná basins. In addition, the presence of an ornithosuchid in the Late Triassic of Brazil suggests that ornithosuchids were more widespread than previously thought in the southern hemisphere. The new ornithosuchid further demonstrates a faunistic link between the Argentinean and Brazilian basins during the Carnian. Finally, the discovery of the new species provides the first clue of a putative necrophagous vertebrate from the oldest dinosaur-bearing beds and expands our knowledge regarding the trophic structure of the Late Triassic of Brazil.

Archosauria originated around the Earth's largest biotic crisis that severely affected all ecosystems globally, the Permotriassic Mass extinction event, and comprises two crown-group lineages: the bird-lineage and the crocodylian lineage. The bird lineage includes the iconic pterosaurs, as well as dinosaurs and birds, whereas the crocodylian lineage includes clades such as aetosaurs, poposaurs, "rauisuchians," as well as Crocodylomorpha; the latter being represented today only by less than 30 extant species of Crocodylia. Despite playing important roles during Mesozoic and Cenozoic ecosystems, both on land and in water, Pseudosuchia received far less attention compared to the bird-lineage, which is also reflected in number and scope of histological studies so far. Lately, the field has seen a shift of focus toward pseudosuchians, however, and the symposium on "Paleohistological Inferences of Paleobiological Traits in Pseudosuchia" held during the International Congress of Vertebrate Morphology 2023 in Cairns, Queensland, Australia, is the latest proof of that. To put these novel aspects of paleohistological and paleobiological research into context, an overview of the non-extant pseudosuchian taxa whose postcranial bones were studied so far is provided here (c. 80 species out of a total of more than 700 extinct species described) and recent trends in pseudosuchian osteohistology are highlighted. In addition, histological studies on cranial and dental material and other potential hard tissues, such as eggshells and otoliths, are briefly reviewed as well., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Tarjadia ruthae is a quadrupedal terrestrial pseudosuchian from the Middle-early Upper Triassic of the Chañares Formation, La Rioja Province, Argentina. Originally, this species was identified as an indeterminate archosaur and later as a doswelliid archosauriform based on very fragmentary specimens characterized by the ornamentation of the skull roof and osteoderms. Additional specimens (including skulls and postcrania) recovered in the last decade show that Tarjadia is an erpetosuchid, an enigmatic pseudosuchian group composed of six species registered in Middle-Upper Triassic continental units of Tanzania, Germany, Scotland, North America, Brazil, and Argentina. Tarjadia ruthae from Argentina and Parringtonia gracilis from Tanzania are the best preserved and more abundant species. Although the monophyly of Erpetosuchidae is well supported, alternative high-level positions within Archosauria have been suggested, such as sister taxon to Crocodylomorpha, Aetosauria, or Ornithosuchidae. In order to improve the knowledge about the erpetosuchids, we performed a detailed description and paleoneurological reconstruction of the skull of Tarjadia ruthae, based on two articulated partial skulls (CRILAR-Pv 478 and CRILAR-Pv 495) and other fragmentary specimens. We analyzed the stratigraphic and geographic occurrence of historical and new specimens of Tarjadia and provided a new emended diagnosis (the same for the genus as for the species, due to monotypy) along with a comparative description of the cranial endocast. The skull of Tarjadia is robust, with a thick and strongly ornamented skull roof, triangular in dorsal view, with concave lateral margins at mid-length that form an abrupt widened posterior region. The external nares are the smallest openings of the skull. The antorbital fossa is deeply excavated and has a small heart-shaped fenestra with both lobes pointing anteriorly. The supratemporal fenestrae are as large and rounded as the orbits, and the infratemporal fenestrae are L-shaped with an extensive excavation along the jugal, quadratojugal and quadrate. The hemimandibles are low, slightly concave on the dentigerous region and strongly convex on the posterior region, conferring them a S-shaped profile in dorsal view. The external mandibular fenestra is small and elliptic, being twice longer than high. The maxillary dentition is restricted to the anterior to mid region of the rostrum. Since the braincase of both specimens is partially damaged, the dorsal surface of the brain could not be entirely reconstructed. As a result, the endocast is anteroposteriorly elongated and seemingly flat, and the cephalic flexure seems to be lower than expected for a suchian. The labyrinth is twice wider than high, the semicircular canals are remarkably straight, and the anterior canal is longer than the posterior one., (© 2024 American Association for Anatomy.)

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., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

In this contribution we introduce a new Late Triassic archosaur, Incertovenator longicollum gen. et sp. nov., with an unusual combination of character states that are present in certain early avemetatarsalian and pseudosuchian archosaur clades. The holotype consists of a partial postcranial skeleton, preserving most of the axial skeleton and displaying a marked anteroposterior elongation in the cervical vertebrae. We include I. longicollum gen. et sp. nov. into one of the most comprehensive early archosaur phylogenetic data sets available, and recover it as either an early diverging avemetatarsalian, closely associated with the clade Aphanosauria and Ornithodira, or as an early diverging loricatan closely related to Mandasuchus tanyauchen in the most parsimonious trees. We further evaluate which alternative phylogenetic positions can I. longicollum gen. et sp. nov. take in the suboptimal trees, and determined which character states support those alternative positions in comparison with those of the unconstrained analysis. The analyses recover the new taxon in three main general phylogenetic placements within Archosauria, as well as one position outside this clade, highlighting widespread morphological evolutionary convergence towards neck elongation in several clades of Triassic archosauriforms. [ABSTRACT FROM AUTHOR]

Adaptive radiations have played a major role in generating modern and deep-time biodiversity. The Triassic radiation of the Archosauromorpha was one of the most spectacular vertebrate radiations, giving rise to many highly ecomorphologically varied lineages—including the dinosaurs, pterosaurs, and stem-crocodylians—that dominated the larger-bodied land fauna for the following 150 Ma, and ultimately gave rise to today’s > 10,000 species of birds and crocodylians. This radiation provides an outstanding testbed for hypotheses relating to adaptive radiations more broadly. Recent studies have started to characterize the tempo and mode of the archosauromorph early adaptive radiation, indicating very high initial rates of evolution, non-competitive niche-filling processes, and previously unrecognized morphological disparity even among non-crown taxa. However, these analyses rested primarily either on discrete characters or on geometric morphometrics of the cranium only, or even failed to fully include phylogenetic information. Here we expand previous 2D geometric morphometric cranial datasets to include new taxa and reconstructions, and create an analogous dataset of the pelvis, thereby allowing comparison of anatomical regions and the transition from “sprawling” to “upright” posture to be examined. We estimated morphological disparity and evolutionary rates through time. All sampled clades showed a delayed disparity peak for sum of variances and average nearest neighbor distances in both the cranium and pelvis, with disparity likely not saturated by the end of the studied time span (Late Jurassic); this contrasts with smaller radiations, but lends weight to similar results for large, ecomorphologically-varied groups. We find lower variations in pelvic than cranial disparity among Triassic-Jurassic archosaurs, which may be related to greater morphofunctional constraints on the pelvis. Contrasting with some previous work, but also confirming some previous findings during adaptive radiations, we find relatively widespread evidence of correlation between sampled diversity and disparity, especially at the largest phylogenetic scales and using average displacement rather than sum of variances as disparity metric; this also demonstrates the importance of comparing disparity metrics, and the importance of phylogenetic scale. Stem and crown archosauromorphs show a morphological diversification of both the cranium and pelvis with higher initial rates (Permian–Middle Triassic and at the base of major clades) followed by lower rates once diversification into niches has occurred (Late Triassic–Jurassic), indicating an “early burst” pattern sensu lato. Our results provide a more detailed and comprehensive picture of the early archosauromorph radiation and have significant bearing on the understanding of deep-time adaptive radiations more broadly, indicating widespread patterns of delayed disparity peaks, initial correlation of diversity and disparity, and evolutionary early bursts.

Dyrosaurids were highly specialized, largely marine, relatives of living crocodylians, and one of the few archosaur lineages to survive the K-Pg extinction. Dyrosaurids lived during the Cretaceous to the Eocene and represent a unique combination of morphology and ecology not seen in living crocodylians. Little is known about their endocranial anatomy, leaving many questions about their neurosensory adaptations unaddressed. Recently, µCT (micro-computed tomography) scans were made of a well-preserved skull of Rhabdognathus, a Paleocene dyrosaurid from Mali. This marks the first time the braincase and neurosensory features of a dyrosaurid have been examined using CT. We focus our attention to three specific internal structures: the cranial endocast; the inner ear; and the paratympanic sinuses. The cranial endocast of Rhabdognathus revealed novel features including a unique conformation of its paratympanic system, a prominent dorsal venous system that communicates with the external skull table, extremely enlarged tympanic vestibules that meet at the midline of the endocranium, a prominent spherical cerebrum, and elongate olfactory tracts accounting for half the total endocast length. The bizarre laterally facing lateral Eustachian foramen of dyrosaurids is now understood to be a complex fossa including both a ventrally directed lateral Eustachian foramen and a laterally directed foramen for the basioccipital diverticulum. A novel median pterygopharyngeal canal was discovered connecting the pharynx to the adductor chamber. These revelations require a reinterpretation of the associated external foramina visible on the posterior of the skull in dyrosaurids and potentially their close relatives the pholidosaurids. The olfactory tract terminates in an enlarged olfactory region possessing complex bony projections—a unique morphology perhaps serving to increase surface area for olfaction. The inner ear of Rhabdognathus exhibits characteristics seen in both Pelagosaurus and Gavialis. The vestibule is spherical, as in Gavialis, but is significantly expanded. The semicircular canals are enlarged but pyramidal in shape as in the thalattosuchian Pelagosaurus. The proportion of the cochlear length to total endosseous labyrinth height is roughly 0.5 in Rhabdognathus implying that the hearing capabilities resemble that of thalattosuchians. A suite of expanded sense organs (e.g., bony olfactory lamina; hypertrophied vestibule of the inner ear), and the clear expansion of the cerebrum to a more symmetrical and spherical shape suggest that dyrosaurids possess neuroanatomical modifications facilitating an agile predatory near-shore ecology.

Four turtle taxa are previously documented from the Cenomanian Arlington Archosaur Site (AAS) of the Lewisville Formation (Woodbine Group) in Texas. Herein, we describe a new side-necked turtle (Pleurodira), Pleurochayah appalachius gen. et sp. nov., which is a basal member of the Bothremydidae. Pleurochayah appalachius gen. et sp. nov. shares synapomorphic characters with other bothremydids, including shared traits with Kurmademydini and Cearachelyini, but has a unique combination of skull and shell traits. The new taxon is significant because it is the oldest crown pleurodiran turtle from North America and Laurasia, predating bothremynines Algorachelus peregrinus and Paiutemys tibert from Europe and North America respectively. This discovery also documents the oldest evidence of dispersal of crown Pleurodira from Gondwana to Laurasia. Pleurochayah appalachius gen. et sp. nov. is compared to previously described fossil pleurodires, placed in a modified phylogenetic analysis of pelomedusoid turtles, and discussed in the context of pleurodiran distribution in the mid-Cretaceous. Its unique combination of characters demonstrates marine adaptation and dispersal capability among basal bothremydids. [ABSTRACT FROM AUTHOR]

Birds are characterized by evolutionary specializations of both locomotion (e.g., flapping flight) and digestive system (toothless, crop, and gizzard), while the potential selection pressures responsible for these evolutionary specializations remain unclear. Here we used a recently developed molecular phyloecological method to reconstruct the diets of the ancestral archosaur and of the common ancestor of living birds (CALB). Our results suggest a trophic shift from carnivory to herbivory (fruit, seed, and/or nut eater) at the archosaur-to-bird transition. The evolutionary shift of the CALB to herbivory may have essentially made them become a low-level consumer and, consequently, subject to relatively high predation risk from potential predators such as gliding non-avian maniraptorans, from which birds descended. Under the relatively high predation pressure, ancestral birds with gliding capability may have then evolved not only flapping flight as a possible anti-predator strategy against gliding predatory non-avian maniraptorans but also the specialized digestive system as an evolutionary tradeoff of maximizing foraging efficiency and minimizing predation risk. Our results suggest that the powered flight and specialized digestive system of birds may have evolved as a result of their tropic shift-associated predation pressure. Yonghua Wu applies recently developed molecular phyloecological methods to identify a trophic shift from carnivory to herbivory in the earliest birds. Predation pressure from becoming low-level consumers may have stimulated the evolution of powered flight as an anti-predator strategy against gliding predatory non-avian maniraptorans. [ABSTRACT FROM AUTHOR]

Reptiles, including birds, exhibit a range of behaviorally relevant adaptations that are reflected in changes to the structure of the inner ear. These adaptations include the capacity for flight and sensitivity to high-frequency sound. We used three-dimensional morphometric analyses of a large sample of extant and extinct reptiles to investigate inner ear correlates of locomotor ability and hearing acuity. Statistical analyses revealed three vestibular morphotypes, best explained by three locomotor categories—quadrupeds, bipeds and simple fliers (including bipedal nonavialan dinosaurs), and high-maneuverability fliers. Troodontids fall with Archaeopteryx among the extant low-maneuverability fliers. Analyses of cochlear shape revealed a single instance of elongation, on the stem of Archosauria. We suggest that this transformation coincided with the origin of both high-pitched juvenile location, alarm, and hatching-synchronization calls and adult responses to them. [ABSTRACT FROM AUTHOR]

Dyrosaurids were highly specialized, largely marine, relatives of living crocodylians, and one of the few archosaur lineages to survive the K-Pg extinction. Dyrosaurids lived during the Cretaceous to the Eocene and represent a unique combination of morphology and ecology not seen in living crocodylians. Little is known about their endocranial anatomy, leaving many questions about their neurosensory adaptations unaddressed. Recently, µCT (micro-computed tomography) scans were made of a well-preserved skull of Rhabdognathus, a Paleocene dyrosaurid from Mali. This marks the first time the braincase and neurosensory features of a dyrosaurid have been examined using CT. We focus our attention to three specific internal structures: the cranial endocast; the inner ear; and the paratympanic sinuses. The cranial endocast of Rhabdognathus revealed novel features including a unique conformation of its paratympanic system, a prominent dorsal venous system that communicates with the external skull table, extremely enlarged tympanic vestibules that meet at the midline of the endocranium, a prominent spherical cerebrum, and elongate olfactory tracts accounting for half the total endocast length. The bizarre laterally facing lateral Eustachian foramen of dyrosaurids is now understood to be a complex fossa including both a ventrally directed lateral Eustachian foramen and a laterally directed foramen for the basioccipital diverticulum. A novel median pterygopharyngeal canal was discovered connecting the pharynx to the adductor chamber. These revelations require a reinterpretation of the associated external foramina visible on the posterior of the skull in dyrosaurids and potentially their close relatives the pholidosaurids. The olfactory tract terminates in an enlarged olfactory region possessing complex bony projections--a unique morphology perhaps serving to increase surface area for olfaction. The inner ear of Rhabdognathus exhibits characteristics seen in both Pelagosaurus and Gavialis. The vestibule is spherical, as in Gavialis, but is significantly expanded. The semicircular canals are enlarged but pyramidal in shape as in the thalattosuchian Pelagosaurus. The proportion of the cochlear length to total endosseous labyrinth height is roughly 0.5 in Rhabdognathus implying that the hearing capabilities resemble that of thalattosuchians. A suite of expanded sense organs (e.g., bony olfactory lamina; hypertrophied vestibule of the inner ear), and the clear expansion of the cerebrum to a more symmetrical and spherical shape suggest that dyrosaurids possess neuroanatomical modifications facilitating an agile predatory near-shore ecology. [ABSTRACT FROM AUTHOR]

Prestosuchus chiniquensis is the most famous “rauisuchian” described by Friedrich von Huene, eight decades ago, and several specimens have been assigned to this taxon since then. In the present contribution, we provide the first detailed description of a complete and very well preserved skull (including the braincase) assigned to Prestosuchus chiniquensis from the Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence of southern Brazil. The detailed description of the skull of Prestosuchus chiniquensis, besides increasing the knowledge about this taxon, may help elucidate the taxonomic relationships of pseudosuchians even further, since most of the characters used in phylogenetic analyzes are cranial. The presence of the subnarial fenestra, a controvertial extra opening on the skull of “rauisuchians”, is thoroughly discussed considering the evidence provided by this new specimen. We consider that the small slit-opening between the premaxilla and the maxilla in Prestosuchus chiniquensis, can not safely be considered a true fenestra, but indicates more likely the existence of some degree of cranial kinesis between these elements which can result in different relative positions of the bones after definitive burial and fossilization, so that the size and shape of this opening is taphonomically controlled. Complementarily, the digital reconstruction of its cranial endocast was developed both from the observation of the preserved braincase and from CT scan images, which resulted in obtaining the first endocast known for a “rauisuchian”. The endocast of Prestosuchus chiniquensis exhibited some remarkable convergences with that of theropod dinosaurs, which could be a reflection of the similar niches they occupied, since “rauisuchians” were the top predators at the end of the Late Triassic, before the extinction of all non-crocodylomorph pseudosuchians.

Two isolated teeth, a dorsal vertebra, fragments of a humerus and femur, a fragmentary pubic “boot” and part of an ischium shaft, identified here as belonging to a large predatory archosaur were discovered in the Upper Triassic site at Marciszów near Zawiercie (southern Poland). Comparisons of the new fossils from Marciszów with the dorsal vertebrae, pubic “boot”, ischium and femur of the theropod-like Smok wawelski from Lisowice (Silesia) reveal that the two taxa are very similar. Nevertheless, due to the lack of more diagnostic elements (e.g., braincase or cranial elements), we prefer to consider all described specimens from Marciszów as Smok sp. Smok sp. shares a low mound-like, anterior trochanter with trochanteric shelf on the femur, a massive pubic “boot” with a distinct depression (= bevelled area), and a transversely lenticular ischium shaft in cross-section with S. wawelski. Some observed characters of the dorsal vertebra (e.g., lack of some lamina, shape and position of zygapophyses), however, are different from S. wawelski and may also suggest that the new findings represent a second species of the genus in the Upper Triassic of Poland. The discovery of Smok sp. at Marciszów is significant because it is the second example of the co-occurrence of this genus with: (i) bones of a large dicynodont; and (ii) record of gnawed tetrapod bones. The discovery of Smok sp. and the lack of significant morphologic divergence from S. wawelski suggest that this taxon is the only large-bodied predator currently known from the Upper Triassic of Poland. This new evidence expands the record of the genus and contributes, in some measure, to our knowledge of the stratigraphical distribution of large predatory archosaurs from the Polish Upper Triassic bone-bearing levels.

Archosaurs displayed an evolutionary trend toward increasing bipedalism in their evolutionary history, that is, forelimbs tend to be reduced in contrast to the development of hindlimbs becoming major weight‐bearing and locomotor appendages. The archosaurian locomotion has been extensively discussed based on their limb morphology because the latter reflects their locomotor modes very well. However, despite some attempts of reconstructing the hindlimb musculature in Archosauria, that of the most distal portion, the pes, has often been neglected. In order to rectify this trend, detailed homologies of pedal muscles among sauropsids were established based on dissections and literature reviews of adult conditions. As a result, homologies of some pedal muscles between non‐avian sauropsids and avians were revised, challenging classical hypotheses. The present new hypothesis postulates that the avian m. tibialis cranialis and non‐avian m. extensor digitorum longus, as well as the avian m. extensor digitorum longus and non‐avian m. tibialis anterior, are homologous with each other, respectively. This is more plausible because it requires no drastical change in the attachment sites between the avian and non‐avian homologues unlike the classical hypothesis. Many interosseous muscles in non‐archosaurian sauropsids that have long been regarded as a part of short digital extensors or flexors are also divided into multiple distinct muscles so that they can be homologized with short pedal muscles among all sauropsids. In addition, osteological correlates of attachments are identified for most of the pedal muscles, contributing to future attempts of reconstruction of this muscle system in fossil archosaurs. [ABSTRACT FROM AUTHOR]

Ever since Darwin, biologists have debated the relative roles of external and internal drivers of large-scale evolution. The distributions and ecology of living crocodilians are controlled by environmental factors such as temperature. Crocodilians have a rich history, including amphibious, marine and terrestrial forms spanning the past 247 Myr. It is uncertain whether their evolution has been driven by extrinsic factors, such as climate change and mass extinctions, or intrinsic factors like sexual selection and competition. Using a new phylogeny of crocodilians and their relatives, we model evolutionary rates using phylogenetic comparative methods. We find that body size evolution follows a punctuated, variable rate model of evolution, consistent with environmental drivers of evolution, with periods of stability interrupted by periods of change. Regression analyses show warmer environmental temperatures are associated with high evolutionary rates and large body sizes. We confirm that environmental factors played a significant role in the evolution of crocodiles. Stockdale and Benton use a new phylogeny of living and extinct crocodilians and their closest relatives to model evolutionary rates and identify the relative influences of extrinsic and intrinsic drivers of evolution in this order. Their results show that environmental factors played a significant role in the evolution of crocodiles. [ABSTRACT FROM AUTHOR]

Loricatan pseudosuchians (known as “rauisuchians”) typically consist of poorly understood fragmentary remains known worldwide from the Middle Triassic to the end of the Triassic Period. Renewed interest and the discovery of more complete specimens recently revolutionized our understanding of the relationships of archosaurs, the origin of Crocodylomorpha, and the paleobiology of these animals. However, there are still few loricatans known from the Middle to early portion of the Late Triassic and the forms that occur during this time are largely known from southern Pangea or Europe. Heptasuchus clarki was the first formally recognized North American “rauisuchian” and was collected from a poorly sampled and disparately fossiliferous sequence of Triassic strata in North America. Exposed along the trend of the Casper Arch flanking the southeastern Big Horn Mountains, the type locality of Heptasuchus clarki occurs within a sequence of red beds above the Alcova Limestone and Crow Mountain formations within the Chugwater Group. The age of the type locality is poorly constrained to the Middle—early Late Triassic and is likely similar to or just older than that of the Popo Agie Formation assemblage from the western portion of Wyoming. The holotype consists of associated cranial elements found in situ, and the referred specimens consist of crania and postcrania. Thus, about 30% of the osteology of the taxon is preserved. All of the pseudosuchian elements collected at the locality appear to belong to Heptasuchus clarki and the taxon is not a chimera as previously hypothesized. Heptasuchus clarki is distinct from all other archosaurs by the presence of large, posteriorly directed flanges on the parabasisphenoid and a distinct, orbit-overhanging postfrontal. Our phylogenetic hypothesis posits a sister-taxon relationship between Heptasuchus clarki and the Ladinian-aged Batrachotomus kupferzellensis from current-day Germany within Loricata. These two taxa share a number of apomorphies from across the skull and their phylogenetic position further supports ‘rauisuchian’ paraphyly. A minimum of three individuals of Heptasuchus are present at the type locality suggesting that a group of individuals died together, similar to other aggregations of loricatans (e.g., Heptasuchus, Batrachotomus, Decuriasuchus, Postosuchus).

Dietary habits in extinct species cannot be directly observed; thus, in the absence of extraordinary evidence, they must be reconstructed with a combination of morphological proxies. Such proxies often include information on dental organization and function such as tooth formation time and tooth replacement rate. In extinct organisms, tooth formation times and tooth replacement rate are calculated, in part via extrapolation of the space between incremental lines in dental tissues representing daily growth (von Ebner Line Increment Width; VEIW). However, to date, little work has been conducted testing assumptions about the primary data underpinning these calculations, specifically, the potential impact of differential sampling and data extrapolation protocols. To address this, we tested a variety of intradental, intramandibular, and ontogentic sampling effects on calculations of mean VEIW, tooth formation times, and replacement rates using histological sections and CT reconstructions of a growth series of three specimens of the extant archosaurian Alligator mississippiensis. We find transect position within the tooth and transect orientation with respect to von Ebner lines to have the greatest impact on calculations of mean VEIW—a maximum number of VEIW measurements should be made as near to the central axis (CA) as possible. Measuring in regions away from the central axis can reduce mean VEIW by up to 36%, causing inflated calculations of tooth formation time. We find little demonstrable impact to calculations of mean VEIW from the practice of subsampling along a transect, or from using mean VEIW derived from one portion of the dentition to extrapolate for other regions of the dentition. Subsampling along transects contributes only minor variations in mean VEIW (

Archosauria (birds, crocodilians and their extinct relatives) form a major part of terrestrial ecosystems today, with over 10 000 living species, and came to dominate the land for most of the Mesozoic (over 150 Myr) after radiating following the Permian–Triassic extinction. The archosaur skull has been essential to this diversification, itself diversified into myriad forms. The archosauriform Euparkeria capensis from the Middle Triassic (Anisian) of South Africa has been of great interest since its initial description in 1913, because its anatomy shed light on the origins and early evolution of crown Archosauria and potentially approached that of the archosaur common ancestor. Euparkeria has been widely used as an outgroup in phylogenetic analyses and when investigating patterns of trait evolution among archosaurs. Although described monographically in 1965, subsequent years have seen great advances in the understanding of early archosaurs and in imaging techniques. Here, the cranium and mandible of Euparkeria are fully redescribed and documented using all fossil material and computed tomographic data. Details previously unclear are fully described, including vomerine dentition, the epiptergoid, number of premaxillary teeth and palatal arrangement. A new diagnosis and cranial and braincase reconstruction is provided, and an anatomical network analysis is performed on the skull of Euparkeria and compared with other amniotes. The modular composition of the cranium suggests a flexible skull well adapted to feeding on agile food, but with a clear tendency towards more carnivorous behaviour, placing the taxon at the interface between ancestral diapsid and crown archosaur ecomorphology, corresponding to increases in brain size, visual sensitivity, upright locomotion and metabolism around this point in archosauriform evolution. The skull of Euparkeria epitomizes a major evolutionary transition, and places crown archosaur morphology in an evolutionary context.