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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.

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Riojasuchus tenuisceps was a pseudosuchian archosaur from the Late Triassic period in Argentina. Like other ornithosuchids, it had unusual morphology such as a unique "crocodile-reversed" ankle joint, a lesser trochanter as in dinosaurs and a few other archosaurs, robust vertebrae, and somewhat shortened, gracile forelimbs. Such traits have fuelled controversies about its locomotor function-were its limbs erect or "semi-erect"? Was it quadrupedal or bipedal, or a mixture thereof? These controversies seem to persist because analyses have been qualitative (functional morphology) or correlative (morphometrics) rather than explicitly, quantitatively testing mechanistic hypotheses about locomotor function. Here, we develop a 3D whole-body model of R. tenuisceps with the musculoskeletal apparatus of the hindlimbs represented in detail using a new muscle reconstruction. We use this model to quantify the body dimensions and hindlimb muscle leverages of this enigmatic taxon, and to estimate joint ranges of motion and qualitative joint functions. Our model supports prior arguments that R. tenuisceps used an erect posture, parasagittal gait and plantigrade pes. However, some of our inferences illuminate the rather contradictory nature of evidence from the musculoskeletal system of R. tenuisceps-different features support (or are ambiguous regarding) quadrupedalism or bipedalism. Deeper analyses of our biomechanical model could move toward a consensus regarding ornithosuchid locomotion. Answering these questions would not only help understand the palaeobiology and bizarre morphology of this clade, but also more broadly if (or how) locomotor abilities played a role in the survival versus extinction of various archosaur lineages during the end-Triassic mass extinction event.

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The clade Pseudosuchia appeared 250 million years ago. The exclusively semi-aquatic Crocodylia, which includes crocodiles, alligators, caimans, and gharials is the only surviving subgroup. Investigating Crocodylia biology is pivotal for inferring traits of extinct pseudosuchians. Alligator femur length is widely used for modeling pseudosuchian body mass, but the regression is influenced by sex and captivity status, leading to potential accuracy problems. An alternative model results from the correlation between alligator femur volume and body mass, which is unaffected by those covariates. Here, an alligator femur volume-based regression is applied to estimate the masses of non-crocodylian pseudosuchians, encompassing goniopholids, dyrosaurs, notosuchians, and thalattosuchians. For each, femur volume as the predictor yields lower body masses than does femur length. Morphological resemblances to existing crocodylians support the inference that extinct goniopholids and dyrosaurs were semi-aquatic. Therefore, body masses predicted from femur length and volume should be reasonable, although larger body masses obtained from femur length may reflect sensitivity to sex or environmental factors. Fully terrestrial notosuchians had proportionately longer femora for their body sizes compared to semi-aquatic crocodylians, suggesting that the higher body masses predicted from alligator femur length are overestimates. Fully aquatic thalattosuchians, skeletally adapted for buoyancy and with reduced reliance on the femur for locomotion, pose challenges for both femur length and volume-based models. The results of this study advocate for the use of femur volume to predict body mass, particularly for semi-aquatic and terrestrial pseudosuchians, and encourage further exploration of volumetric models as body size predictors for extinct vertebrates.

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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.

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Maximum individual body size in pseudosuchian archosaurs is not well constrained in the fossil record, but it may be influenced by a variety of factors including basal metabolic rate, evolutionary relationships, and environmental conditions. Body size varies among the Aetosauria in which estimated total length ranges between 1 m (e.g., Coahomasuchus kahleorum) and 5 m (e.g., Desmatosuchus spurensis). A new, very large specimen of the aetosaurian Typothorax coccinarum from Petrified Forest National Park in northeastern Arizona is nearly twice the size of all other known specimens of Typothorax and is the largest aetosaur specimen currently known worldwide. The specimen lacks co-ossified neurocentral sutures in the trunk vertebrae which may suggest that the individual had not yet reached skeletal maturity, yet smaller specimens of T. coccinarum exhibit partially or fully co-ossified neurocentral sutures in the same region. If body size correlates with skeletal maturity in aetosaurs, this discrepancy warns that timing of neurocentral suture co-ossification in aetosaurs may not be a reliable indicator of ontogenetic stage. Osteohistological observations of a trunk rib demonstrate that although PEFO 42506 shows a large body size, the specimen did not deposit an external fundamental system despite depositing as many as 19 growth lines, further indicating that it had not yet reached skeletal maturity. Thus, at least within Aetosauria, neurocentral suture co-ossification and skeletal maturity may correlate, whereas body size can be incongruent in comparison. Furthermore, this specimen indicates that non-desmatosuchin aetosaurs could exhibit large body sizes and suggests that some aetosaurs may have experienced indeterminate growth.

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Osteoderms are mineralized structures embedded in the dermis, known for nonavian archosaurs, squamates, xenarthrans, and amphibians. Herein, we compared the osteoderm histology of Brazilian Notosuchia of Cretaceous age using three neosuchians for comparative purposes. Microanatomical analyses showed that most of them present a diploe structure similar to those of other pseudosuchians, lizards, and turtles. This structure contains two cortices (the external cortex composed of an outer and an inner layers, and the basal cortex) and a core in-between them. Notosuchian osteoderms show high bone compactness (>0.85) with varying degrees of cancellous bone in the core. The neosuchian Guarinisuchus shows the lowest bone compactness with a well-developed cancellous layer. From an ontogenetic perspective, most tissues are formed through periosteal ossification, although the mineralized tissues observed in baurusuchid LPRP/USP 0634 suggest a late metaplastic development. Histology suggests that the ossification center of notosuchian osteoderm is located at the keel. Interestingly, we identified Sharpey's fibers running perpendicularly to the outer layer of the external cortex in Armadillosuchus arrudai, Itasuchus jesuinoi, and Baurusuchidae (LPRP/USP 0642). This feature indicates a tight attachment within the dermis, and it is evidence for the presence of an overlying thick leathery layer of skin over these osteoderms. These data allow a better understanding of the osteohistological structure of crocodylomorph dermal bones, and highlight their structural diversity. We suggest that the vascular canals present in some sampled osteoderms connecting the inner layer of the external cortex and the core with the external surface may increase osteoderm surface and the capacity of heat transfer in terrestrial notosuchians.

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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.

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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.

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Fossil trackways provide a glimpse into the behavior of extinct animals. However, while providing information of the trackmaker size, stride, and even speed, the actual gait of the organism can be ambiguous. This is especially true of quadrupedal animals, where disparate gaits can have similar trackway patterns. Here, predictive simulation using trajectory optimization can help distinguish gaits used by trackmakers. First, we demonstrated that a planar, five-link quadrupedal biomechanical model can generate the qualitative trackway patterns made by domestic dogs, although a systematic error emerges in the track phase (relative distance between ipsilateral pes and manus prints). Next, we used trackway dimensions as inputs to a model of Batrachotomus kupferzellensis, a long-limbed, crocodile-line archosaur (clade Pseudosuchia) from the Middle Triassic of Germany. We found energetically optimal gaits and compared their predicted track phases to those of fossil trackways of Isochirotherium and Brachychirotherium. The optimal results agree with trackways at slow speeds but differ at faster speeds. However, all simulations point to a gait transition around a non-dimensional speed of 0.4 and another at 1.0. The trackways likewise exhibit stark differences in the track phase at these speeds. In all cases, including when simulations are constrained to the fossil track phase, the optimal simulations after the first gait transition do not correspond to a trot, as often used by living crocodiles. Instead, they are a diagonal sequence gait similar to the slow tölt of Icelandic horses. This is the first evidence that extinct pseudosuchians may have exhibited different gaits than their modern relatives and of a gait transition in an extinct pseudosuchian. The results of this analysis highlight areas where the models can be improved to generate more reliable predictions for fossil data while also showcasing how simple models can generate insights about the behavior of extinct animals.

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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).

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Aetosaurs comprise a clade of quadrupedal, armored, omnivores to herbivores that lived across much of the supercontinent of Pangea during the Late Triassic. Their relative abundance in many units, and the rarity of other Triassic herbivores, points to them as key components of Late Triassic ecosystems. Debate persists about whether they were growing more or less slowly when compared to extant crocodylians, and bone histology is sparsely sampled within the group. We undertook a histological examination of Coahomasuchus chathamensis to address its ontogenetic trajectory and characterize its histology. We sampled a paramedian osteoderm from the holotype specimen, as well as five osteoderms (two paramedian, one lateral, and two of uncertain position) and two incomplete limb bones (radius and fibula), from referred specimens discovered at the type locality. Using these we estimated specimen ages with lines of arrested growth (LAGs) to determine that the study individuals reached from 2 to 7 years old. All of the sampled elements contained woven-fibered bone with extensive vasculature within the internal cortex. In some specimens, more poorly vascularized, parallel-fibered bone is evident externally. The holotype of C. chathamensis represents a juvenile individual, and raises the possibility that the holotype of C. kahleorum is a juvenile as well, complicating aetosaur systematics and diversity. When compared to aetosaurs of similar size, it is clear that C. chathamensis was growing comparatively rapidly, about 3 times the rate of similarly sized specimens of Aetosauroides scagliai from Argentina. This discovery reveals the presence of disparate growth strategies within Aetosauria. Anat Rec, 302:1504-1515, 2019. © 2018 American Association for Anatomy.

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Stagonolepis robertsoni, from the Late Triassic of Scotland, was the first named aetosaurian. Known mostly from a series of natural molds from two localities, the osteology of this taxon has been difficult to interpret. Detailed work on this material in the late 1950s resulted in a monograph that set the standard for the understanding of aetosaurians, making Stagonolepis robertsoni the best known aetosaurian; however, little has been done with this material since. Reanalysis of this material shows that despite its limitations the early 1960s reconstruction work depicts the preserved characteristics faithfully, especially in the skull. The first cervical rib is extremely anteroposteriorly elongate as in Alligator, a character not previously recognized in aetosaurians. Diapophyseal and zygapophyseal vertebral laminae are present in the cervical and trunk vertebrae. The ilium is autapomorphic with distinct pre- and post-processes of the iliac blade. The osteoderms differ from North and South American material that has been ascribed to the genus. Those assignments are based on plesiomorphies within Aetosauria, such as a radial ornamentation and a posteriorly located and medially offset dorsal eminence. Biostratigraphic correlations using taxonomic conclusions based on plesiomorphic characters should not be used. The holotype specimen of S. robertsoni is currently diagnostic, in part because ventral osteoderms are not known for many aetosaurian taxa and the surface ornamentation of randomly distributed, closely packed oblong pits found in S. robertsoni is unique within Aetosauria.

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Phytosaurs are a clade of large, carnivorous pseudosuchian archosaurs from the Late Triassic with a near cosmopolitan distribution. Their superficial resemblance to longirostrine (long-snouted) crocodylians, such as gharials, has often been used in the past to infer ecological and behavioural convergence between the two groups. Although more than thirty species of phytosaur are currently recognised, little is known about the endocranial anatomy of this clade. Here, we describe the endocranial anatomy (including the brain, inner ear, neurovascular structures and sinus systems) of the two non-mystriosuchine phytosaurs Parasuchus angustifrons (="Paleorhinus angustifrons") and Ebrachosuchus neukami from the Late Triassic of Germany based on digital reconstructions. Results show that the endocasts of both taxa are very similar to each other in their rostrocaudally elongate morphology, with long olfactory tracts, weakly demarcated cerebral regions and dorsoventrally short endosseous labyrinths. In addition, several sinuses, including large antorbital sinuses and prominent dural venous sinuses, were reconstructed. Comparisons with the endocranial anatomy of derived phytosaurs indicate that Phytosauria is united by the presence of elongate olfactory tracts and longitudinally arranged brain architecture-characters which are also shared with Crocodyliformes. However, a substantial morphological variability is observed in the cephalic and pontine flexure and the presence of a pineal organ across the different phytosaur species. These results suggest that the endocranial anatomy in Phytosauria generally follows a plesiomorphic pattern, with moderate variation within the clade likely resulting from divergent sensory and behavioural adaptations.

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The 'rauisuchians' are a group of Triassic pseudosuchian archosaurs that displayed a near global distribution. Their problematic taxonomic resolution comes from the fact that most taxa are represented only by a few and/or mostly incomplete specimens. In the last few decades, renewed interest in early archosaur evolution has helped to clarify some of these problems, but further studies on the taxonomic and paleobiological aspects are still needed. In the present work, we describe new material attributed to the 'rauisuchian' taxon Prestosuchus chiniquensis, of the Dinodontosaurus Assemblage Zone, Middle Triassic (Ladinian) of the Santa Maria Supersequence of southern Brazil, based on a comparative osteologic analysis. Additionally, we present well supported evidence that these represent juvenile forms, due to differences in osteological features (i.e., a subnarial fenestra) that when compared to previously described specimens can be attributed to ontogeny and indicate variation within a single taxon of a problematic but important osteological structure in the study of 'rauisuchians.'

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Aetosauria is an early-diverging clade of pseudosuchians (crocodile-line archosaurs) that had a global distribution and high species diversity as a key component of various Late Triassic terrestrial faunas. It is one of only two Late Triassic clades of large herbivorous archosaurs, and thus served a critical ecological role. Nonetheless, aetosaur phylogenetic relationships are still poorly understood, owing to an overreliance on osteoderm characters, which are often poorly constructed and suspected to be highly homoplastic. A new phylogenetic analysis of the Aetosauria, comprising 27 taxa and 83 characters, includes more than 40 new characters that focus on better sampling the cranial and endoskeletal regions, and represents the most comprenhensive phylogeny of the clade to date. Parsimony analysis recovered three most parsimonious trees; the strict consensus of these trees finds an Aetosauria that is divided into two main clades: Desmatosuchia, which includes the Desmatosuchinae and the Stagonolepidinae, and Aetosaurinae, which includes the Typothoracinae. As defined Desmatosuchinae now contains Neoaetosauroides engaeus and several taxa that were previously referred to the genus Stagonolepis, and a new clade, Desmatosuchini, is erected for taxa more closely related to Desmatosuchus. Overall support for some clades is still weak, and Partitioned Bremer Support (PBS) is applied for the first time to a strictly morphological dataset demonstrating that this weak support is in part because of conflict in the phylogenetic signals of cranial versus postcranial characters. PBS helps identify homoplasy among characters from various body regions, presumably the result of convergent evolution within discrete anatomical modules. It is likely that at least some of this character conflict results from different body regions evolving at different rates, which may have been under different selective pressures.