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Fossils of embryonic and hatchling individuals can provide invaluable insight into the evolution of prenatal morphologies, heterochronies, and allometric trajectories within Archosauria but are exceptionally rare in the Triassic fossil record, obscuring a critical aspect of archosaurian biology during their evolutionary origins. Microvertebrate sampling at a single bonebed in the Upper Triassic Chinle Formation within Petrified Forest National Park has yielded diminutive archosauriform femora (PEFO 45274, PEFO 45199) with estimated and measured femoral lengths of ~31 mm and ~ 37 mm, respectively. These new specimens provide the unique opportunity to assess the preservation, body size, and growth dynamics of skeletally immature archosauriforms in North America and compare the growth dynamics of archosauromorphs within an evolutionary and ontogenetic context. We assign PEFO 45199 and PEFO 45274 to Phytosauria (Archosauriformes) based on their strongly sigmoidal shape in lateral view, the presence of proximal anterolateral and posteromedial tubera, the absence of an anteromedial tuber of the proximal end, a teardrop‐shaped proximal outline, and a fourth trochanter that is not confluent with the proximal head. Osteohistological analyses of PEFO 45274 reveal a cortex comprising low vascularity, parallel‐fibered bone composed of primary osteons that lacks a hatching line and any lines of arrested growth. We interpret PEFO 45274 as a slow‐growing, post‐hatching individual of less than 1 year of age. Surprisingly, osteohistology of some larger phytosaur femora implies faster growth rates in comparison to PEFO 45274 based on the occasional presence of woven bone and overall higher degrees of vascular density, suggesting the ontogenetic shift from rapid‐to‐slow growth rates might not occur simply or uniformly as expected in Phytosauria and that non‐archosaurian archosauriforms may exhibit size‐dependent histological characteristics. This study highlights the importance of including osteohistology from multiple body sizes to investigate non‐archosaurian archosauriform ancestral growth rates given the phylogenetic position of phytosaurs near the divergence of Archosauria. [ABSTRACT FROM AUTHOR]

Spinosaurids are a highly morphologically specialized clade of large tetanuran theropod dinosaurs, characterized by their proportionally long, narrow snouts and conical teeth with fluted surfaces. The unusual morphology of spinosaurids has been usually compared with modern crocodilians, but distinct differences between them have been reported as well. Here, numerous craniodental characters that are shared by spinosaurids and large, robust phytosaurs often termed as "brachyrostral" forms are reviewed. Phytosaurs are a clade of carnivorous archosauriforms that resemble crocodilians in overall morphology, and as such a similar lifestyle has been inferred for them as well. Although based on preliminary observations, the shared craniodental characters of spinosaurids and "brachyrostral" phytosaurs, including those not shared with crocodilians, raise the possibility that at least some aspects of the lifestyle of these extinct archosauriform clades were more similar to each other rather than to that of crocodilians [ABSTRACT FROM AUTHOR]

The Panchet Formation of northeastern India preserves an Induan (earliest Triassic) vertebrate assemblage with only one valid archosauromorph species, the proterosuchid Samsarasuchus pamelae. Two other archosauromorph species have been named for this unit: “Ankistrodon indicus” and “Teratosaurus(?) bengalensis”. “Ankistrodon indicus”, based on a fragment of maxilla with two partial teeth, is indistinguishable from other proterosuchid species and is considered a nomen dubium. “Teratosaurus(?) bengalensis” is represented by an isolated, almost complete tooth that was distinguished from “Ankistrodon indicus” in the presence of mesial denticles. “Teratosaurus(?) bengalensis” is also indistinguishable from other valid proterosuchid species and considered likely a nomen dubium. However, it remains unresolved if “Teratosaurus(?) bengalensis” represents a second Panchet archosauromorph species different from Samsarasuchus pamelae and “Ankistrodon indicus” because the presence of mesial denticles cannot be determined in these species due to their incompleteness. Here we describe two new Panchet Proterosuchidae specimens collected from the same locality as the holotypes of the three above-mentioned species. These new specimens are maxillary fragments and one of them has almost complete teeth with mesial denticles on the apical portion of the crowns, as in “Teratosaurus(?) bengalensis.” The rest of their morphology is congruent with that of Samsarasuchus pamelae and “Ankistrodon indicus.” As a result, the specimens reported here expand our anatomical knowledge of the Panchet archosauriform assemblage and indicate that there is no current evidence for the presence of more than one valid archosauromorph species in the Panchet Formation.

Archosaurs diversified and became dominant during the Mesozoic Era, but their earliest relatives (non-archosaurian archosauromorphs) were already scarcely present in the late Permian. Here we describe a new species of non-archosaurian archosauriform from the upper Permian of Xinjiang, China. Preserved as a partial hindlimb, it possesses a few derived features shared with other archosauriforms, including a much stouter tibia than fibula, a longer metatarsal III than metatarsal IV, and a hooked metatarsal V. Phylogenetic analysis confirmed the new taxon to be a non-archosaurian archosauriform. The morphology of the knee, crus, and pes shows traits that are commonly related with a parasagittal posture, including an entirely proximo-distal articulation of the femur and fibula, the slender and closely spaced tibia and fibula, and a mesaxonic foot with a reduced fifth toe. The new taxon shows that the parasagittal posture evolved before the end-Permian Mass Extinction. [ABSTRACT FROM AUTHOR]

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.

Heteropelta boboi is a new archosauriform reptile from the upper Anisian of northeastern Italy represented by a fragment of dorsal armor with a row of neural arches of the dorsal vertebrae. The dorsal armor of the new taxon is composed of two columns of paramedian osteoderms and at least six columns of lateral osteoderms. Unlike other armored archosaurs, the osteoderms are imbricated with the posterior osteoderm overlapping the anterior one. The low neural arches bear small neural spines and long postzygapophyses. The osteoderms of the lateral columns increase in size and change in shape from the most medial to the most lateral columns. Among the Archosauriformes, only the non-archosaur proterochampsians Vancleavea campi, Litorosuchus somnii, and the doswelliids have dorsal armor comprised of more than two columns of osteoderms per side, but the morphology and arrangement of their osteoderms is unlike those of the new Italian taxon. A cladistic analysis of Archosauromorpha positions Heteropelta boboi as either a basal phytosaur or a basal suchian. However, a second cladistic analysis focused on armored archosaurs alternatively positions the new taxon as a basal archosauriform, basal suchian, basal loricatan or crocodylomorph. Better resolution of the phylogenetic relationships of Heteropelta boboi will likely be obtained only with the discovery of cranial and postcranial remains associated with its diagnostic armor elements. [ABSTRACT FROM AUTHOR]

Heteropelta boboi is a new archosauriform reptile from the upper Anisian of northeastern Italy represented by a fragment of dorsal armor with a row of neural arches of the dorsal vertebrae. The dorsal armor of the new taxon is composed of two columns of paramedian osteoderms and at least six columns of lateral osteoderms. Unlike other armored archosaurs, the osteoderms are imbricated with the posterior osteoderm overlapping the anterior one. The low neural arches bear small neural spines and long postzygapophyses. The osteoderms of the lateral columns increase in size and change in shape from the most medial to the most lateral columns. Among the Archosauriformes, only the non-archosaur proterochampsians Vancleavea campi, Litorosuchus somnii, and the doswelliids have dorsal armor comprised of more than two columns of osteoderms per side, but the morphology and arrangement of their osteoderms is unlike those of the new Italian taxon. A cladistic analysis of Archosauromorpha positions Heteropelta boboi as either a basal phytosaur or a basal suchian. However, a second cladistic analysis focused on armored archosaurs alternatively positions the new taxon as a basal archosauriform, basal suchian, basal loricatan or crocodylomorph. Better resolution of the phylogenetic relationships of Heteropelta boboi will likely be obtained only with the discovery of cranial and postcranial remains associated with its diagnostic armor elements.

The Placerias/Downs' Quarry complex in eastern Arizona, USA, is the most diverse Upper Triassic vertebrate locality known. We report a new short-faced archosauriform, Syntomiprosopus sucherorum gen. et sp. nov., represented by four incomplete mandibles, that expands that diversity with a morphology unique among Late Triassic archosauriforms. The most distinctive feature of Syntomiprosopus gen. nov. is its anteroposteriorly short, robust mandible with 3–4 anterior, a larger caniniform, and 1–3 "postcanine" alveoli. The size and shape of the alveoli and the preserved tips of replacement teeth preclude assignment to any taxon known only from teeth. Additional autapomorphies of S. sucherorum gen. et sp. nov. include a large fossa associated with the mandibular fenestra, an interdigitating suture of the surangular with the dentary, fine texture ornamenting the medial surface of the splenial, and a surangular ridge that completes a 90° arc. The external surfaces of the mandibles bear shallow, densely packed, irregular, fine pits and narrow, arcuate grooves. This combination of character states allows an archosauriform assignment; however, an associated and similarly sized braincase indicates that Syntomiprosopus n. gen. may represent previously unsampled disparity in early-diverging crocodylomorphs. The Placerias Quarry is Adamanian (Norian, maximum depositional age ~219 Ma), and this specimen appears to be an early example of shortening of the skull, which occurs later in diverse archosaur lineages, including the Late Cretaceous crocodyliform Simosuchus. This is another case where Triassic archosauriforms occupied morphospace converged upon by other archosaurs later in the Mesozoic and further demonstrates that even well-sampled localities can yield new taxa. [ABSTRACT FROM AUTHOR]

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]

Erythrosuchidae is a clade of early archosauriform reptiles that were large-bodied, hypercarnivorous, possibly apex pred-ators in late Early and Middle Triassic ecosystems following the Permo-Triassic mass extinction. Chalishevia cothurnatafrom the late Middle Triassic (Ladinian) of Russia, is the stratigraphically youngest known erythrosuchid species, but the holotype and referred material of this taxon has received little study. Here, we provide the first detailed anatomical description of C. cothurnata, including comparisons to other erythrosuchids. Although known from relatively fragmen-tary material, the anatomy of C. cothurnata is distinctive, including an autapomorphic strongly slanted ventral border of the antorbital fossa. The presence of a large accessory opening (the “accessory antorbital fenestra”) in the skull between the premaxilla, nasal and maxilla, together with the inferred presence of a narrow postnarial process of the premaxilla that articulated with a slot on the nasal, provides strong evidence for a sister taxon relationship between C. cothurnata and the erythrosuchid Shansisuchus shansisuchus from the early Middle Triassic (Anisian) of China. The inferred basal skull length of C. cothurnata was approximately 80 cm, making it one of the largest erythrosuchids known.

Proterochampsians are a South American endemic group of non-archosaurian archosauriforms with morphological characteristics recollecting Recent crocodilians, and therefore have been proposed as aquatic species. However, this has not been based on careful examination of anatomical and histological features. We provide a review of the morphological and histological evidence present in the skeleton of proterochampsids and discuss its implications for inferring the lifestyles of these organisms. Anatomical features such as a secondary palate, marginal dentition, palatine teeth, morphology of the tail, limb modification, and dermal armor are reviewed, and details of histological structures are described based on bone thin sections. Histological examination reveals a predominance of fibrolamellar bone tissue, suggesting rapid periosteal osteogenesis and therefore overall fast bone growth. The existence of discontinuities (LAGs) demonstrates that these animals responded to changes in their environment. Ecomorphological features do not provide definitive evidence for the lifestyles of proterochampsids, but allow us to propose a terrestrial/amphibious condition. The same is true of the histological features, particularly compactness of the bone.

Erythrosuchid archosauriforms are quadrupedal carnivorous reptiles with a proportionally huge skull. They represent one of the first evolutionary radiations of medium to large predatory diapsids after the Permo–Triassic mass extinction. Erythrosuchids are known from Lower– Middle Triassic rocks of South Africa, Russia, and China, and there have been preliminary reports from the Middle Triassic Yerrapalli Formation of south-central India. Here we describe, compare and figure for the first time these Indian erythrosuchid remains. We erect the new genus and species Bharitalasuchus tapani based on a holotype and paratype that preserve tooth-bearing cranial fragments, at least 17 presacral vertebrae, some ribs and probable intercentra, and partial shoulder and pelvic girdles and hindlimb and allow recognizing a series of autapomorphies and unique combination of character states among erythrosuchids. Our phylogenetic analysis recovered Bharitalasuchus tapani most closely related to Shansisuchus shansisuchus and Chalishevia cothurnata from the late Anisian of China and Ladinian of Russia, respectively. The phylogenetic affinities of this new taxon and a revision of the tetrapod assemblage of the Yerrapalli Formation shed light on the age of this unit. The presence of the Wadiasaurus-Rechnisaurus-Bharitalasuchus association in the Yerrapalli Formation closely resembles the Sinokannemeyeria- Shansisuchus dicynodont-erythrosuchid association of late Anisian to early Ladinian Chinese units. This evidence supports a post-early–middle Anisian age, even possibly early Ladinian, for the Yerrapalli Formation. The presence of possibly one of the last erythrosuchids in India would indicate that the clade still retained both a northern and southern Pangean distribution before its extinction. [ABSTRACT FROM AUTHOR]

Proterochampsidae is a clade of non-archosaurian archosauriforms restricted to the Middle to the Late Triassic of the Ischigualasto-Villa Unión Basin of Argentina and the Santa Maria Supersequence of Brazil. A reappraisal of proterochampsid specimens from the Brazilian Dinodontosaurus Assemblage Zone (AZ) of the Pinheiros-Chiniquá Sequence (late Ladinian-early Carnian) is presented here. One of the specimens was preliminary assigned to Chanaresuchus sp., whose type species comes from the Massetognathus-Chanaresuchus AZ of the Chañares Formation of Argentina. However, our revision indicates that it differs from Chanaresuchus, being more closely related to the middle-late Carnian Rhadinosuchus gracilis. We therefore propose the new taxon, Pinheirochampsa rodriguesi, to reallocate this specimen. Additionally, we present a revision of other putative Chanaresuchus occurrences in Brazil, including the only known specimen described for the Santacruzodon AZ (Santa Cruz do Sul Sequence; early Carnian), also proposing it as a new taxon: Kuruxuchampsa dornellesi. Both new species are characterized, among other features, by transverse expansion of the anterior end of the rostrum, similar to the condition present in Rhadinosuchus, but absent in Chanaresuchus, Gualosuchus, Pseudochampsa, and non-rhadinosuchine proterochampsids. These two new species expand the growing knowledge of the non-archosaurian archosauriform diversity during the Middle-Late Triassic in South America and enhance faunal and chronological comparisons between approximately coeval geological units between Argentina and Brazil., (© 2023 American Association for Anatomy.)

The six known specimens of Scleromochlus taylori and casts made from their negative impressions were examined to reassess the osteological evidence that has been used to interpret Scleromochlus’s locomotion and phylogenetic relationships. It was found that the trunk was dorsoventrally compressed. The upper temporal fenestra was on the lateral surface of skull and two-thirds the size of the lower, the jaw joint posteriorly placed with short retroarticular process, and teeth short and subconical, but no evidence of external nares or antorbital fossae was found. The posterior trunk was covered with ~20 rows of closely spaced transversely elongate dorsal osteoderms. The coracoid was robust and elongate. The acetabulum was imperforate and the femoral head hemispherical and only weakly inturned such that the hip joint was unsuited to swinging in a parasagittal plane. The presence of four distal tarsals is confirmed. The marked disparity of tibial and fibular shaft diameters and of proximal tarsal dimensions indicates that the larger proximal tarsal is the astragalus and the significantly smaller tarsal is the calcaneum. The astragalus and calcaneum bear little resemblance to those of Lagosuchus, and the prominent calcaneal tuber confirms that the ankle was crurotarsal. There is no evidence that preserved body and limb postures are unnatural, and most specimens are preserved in what is interpreted as a typical sprawling resting pose. A principal component analysis of skeletal measurements of Scleromochlus and other vertebrates of known locomotor type found Scleromochlus to plot with frogs, and that finding combined with skeletal morphology suggests Scleromochlus was a sprawling quadrupedal hopper. Phylogenetic analyses found that Scleromochlus was not an ornithodiran, but was either within the Doswelliidae or outside the clade consisting of the most recent common ancestor of the Erythrosuchidae and Archosauria and all its descendants.

Erythrosuchidae is a clade of early archosauriform reptiles that were large-bodied, hypercarnivorous, possibly apex predators in late Early and Middle Triassic ecosystems following the Permo-Triassic mass extinction. Chalishevia cothurnata from the late Middle Triassic (Ladinian) of Russia, is the stratigraphically youngest known erythrosuchid species, but the holotype and referred material of this taxon has received little study. Here, we provide the first detailed anatomical description of C. cothurnata, including comparisons to other erythrosuchids. Although known from relatively fragmentary material, the anatomy of C. cothurnata is distinctive, including an autapomorphic strongly slanted ventral border of the antorbital fossa. The presence of a large accessory opening (the "accessory antorbital fenestra") in the skull between the premaxilla, nasal and maxilla, together with the inferred presence of a narrow postnarial process of the premaxilla that articulated with a slot on the nasal, provides strong evidence for a sister taxon relationship between C. cothurnata and the erythrosuchid Shansisuchus shansisuchus from the early Middle Triassic (Anisian) of China. The inferred basal skull length of C. cothurnata was approximately 80 cm, making it one of the largest erythrosuchids known. [ABSTRACT FROM AUTHOR]

Proterosuchids are an important group of carnivorous basal archosauriforms characterized by a bizarre and enigmatic downturned premaxilla that overhangs the lower jaw. They are particularly significant because they radiated in the immediate aftermath of the Permian–Triassic mass extinction and represent one of the best known 'disaster taxa' following that event. While traditionally considered to be semi‐aquatic, recent histological studies and geological data have suggested that it is more likely that they inhabited terrestrial environments. By using computed tomographic (CT) data, we virtually reconstruct the brain endocast and endosseous labyrinths of two adult specimens of Proterosuchus fergusi from the earliest Triassic of South Africa, in an attempt to understand its life habits within the context of basal archosauriform evolution. Endocasts reveal that the brain cavity is tubular in shape and the endosseous labyrinths are highly pyramidal. The angle of the lateral semicircular canal suggests that P. fergusi naturally held its head upwards ~17°, while the length of the cochlear duct suggests its auditory abilities were specialized towards low‐frequency sounds. Furthermore, beam theory analysis suggests that the rostrum of P. fergusi is highly resistant to both bending and torsion when compared to modern crocodilians, although this resistance is neither enhanced nor reduced by the overhanging premaxilla. Comparative anatomical analyses suggest P. fergusi was probably a semi‐aquatic, generalist apex predator capable of surviving the harsh environmental perturbations of the Early Triassic. [ABSTRACT FROM AUTHOR]

The six known specimens of Scleromochlus taylori and casts made from their negative impressions were examined to reassess the osteological evidence that has been used to interpret Scleromochlus's locomotion and phylogenetic relationships. It was found that the trunk was dorsoventrally compressed. The upper temporal fenestra was on the lateral surface of skull and two-thirds the size of the lower, the jaw joint posteriorly placed with short retroarticular process, and teeth short and subconical, but no evidence of external nares or antorbital fossae was found. The posterior trunk was covered with ~20 rows of closely spaced transversely elongate dorsal osteoderms. The coracoid was robust and elongate. The acetabulum was imperforate and the femoral head hemispherical and only weakly inturned such that the hip joint was unsuited to swinging in a parasagittal plane. The presence of four distal tarsals is confirmed. The marked disparity of tibial and fibular shaft diameters and of proximal tarsal dimensions indicates that the larger proximal tarsal is the astragalus and the significantly smaller tarsal is the calcaneum. The astragalus and calcaneum bear little resemblance to those of Lagosuchus, and the prominent calcaneal tuber confirms that the ankle was crurotarsal. There is no evidence that preserved body and limb postures are unnatural, and most specimens are preserved in what is interpreted as a typical sprawling resting pose. A principal component analysis of skeletal measurements of Scleromochlus and other vertebrates of known locomotor type found Scleromochlus to plot with frogs, and that finding combined with skeletal morphology suggests Scleromochlus was a sprawling quadrupedal hopper. Phylogenetic analyses found that Scleromochlus was not an ornithodiran, but was either within the Doswelliidae or outside the clade consisting of the most recent common ancestor of the Erythrosuchidae and Archosauria and all its descendants. [ABSTRACT FROM AUTHOR]

Erythrosuchidae are a globally distributed and important group of apex predators that occupied Early and Middle Triassic terrestrial ecosystems following the Permo-Triassic mass extinction. The stratigraphically oldest known genus of Erythrosuchidae is Garjainia Ochev, 1958, which is known from the late Early Triassic (late Olenekian) of European Russia and South Africa. Two species of Garjainia have been reported from Russia: the type species, Garjainia prima Ochev, 1958, and ‘Vjushkovia triplicostata’ von Huene, 1960, which has been referred to Garjainia as either congeneric (Garjainia triplicostata) or conspecific (G. prima). The holotype of G. prima has received relatively extensive study, but little work has been conducted on type or referred material attributed to ‘V. triplicostata’. However, this material includes well-preserved fossils representing all parts of the skeleton and comprises seven individuals. Here, we provide a comprehensive description and review of the cranial anatomy of material attributed to ‘V. triplicostata’, and draw comparisons with G. prima. We conclude that the two Russian taxa are indeed conspecific, and that minor differences between them result from a combination of preservation or intraspecific variation. Our reassessment therefore provides additional information on the cranial anatomy of G. prima. Moreover, we quantify relative head size in erythrosuchids and other early archosauromorphs in an explicit phylogenetic context for the first time. Our results show that erythrosuchids do indeed appear to have disproportionately large skulls, but that this is also true for other early archosauriforms (i.e. proterosuchids), and may reflect the invasion of hypercarnivorous niches by these groups following the Permo-Triassic extinction.

Erythrosuchidae is a clade of early archosauriform reptiles that were large-bodied, hypercarnivorous, possibly apex predators in late Early and Middle Triassic ecosystems following the Permo-Triassic mass extinction. Chalishevia cothurnata from the late Middle Triassic (Ladinian) of Russia, is the stratigraphically youngest known erythrosuchid species, but the holotype and referred material of this taxon has received little study. Here, we provide the first detailed anatomical description of C. cothurnata, including comparisons to other erythrosuchids. Although known from relatively fragmentary material, the anatomy of C. cothurnata is distinctive, including an autapomorphic strongly slanted ventral border of the antorbital fossa. The presence of a large accessory opening (the "accessory antorbital fenestra") in the skull between the premaxilla, nasal and maxilla, together with the inferred presence of a narrow postnarial process of the premaxilla that articulated with a slot on the nasal, provides strong evidence for a sister taxon relationship between C. cothurnata and the erythrosuchid Shansisuchus shansisuchus from the early Middle Triassic (Anisian) of China. The inferred basal skull length of C. cothurnata was approximately 80 cm, making it one of the largest erythrosuchids known. [ABSTRACT FROM AUTHOR]

Erythrosuchids are Early–Middle Triassic bulky, quadrupedal archosauromorphs with proportionally large heads, and they were among the first archosauromorphs to occupy the role of terrestrial hypercarnivores. Recent work has expanded anatomical and phylogenetic knowledge of erythrosuchids, but a thorough understanding is still lacking, and Erythrosuchus africanus is the only species for which a thorough anatomical description of most of the skeleton has been published. Garjainia prima, from the upper Olenekian of Russia, represents one of the oldest known erythrosuchids and includes the best-preserved skull of an Early Triassic erythrosuchid. Here, we expand understanding of erythrosuchid anatomy by providing the first detailed and comprehensive description of the holotype of Garjainia prima. We consider ' Vjushkovia triplicostata ' to be a junior synonym of Garjainia prima, because of their extremely similar morphology, with the two character states that differ between them being variable intraspecifically in some other diapsids. A new phylogenetic analysis recovered a monophyletic Garjainia as sister to Middle Triassic erythrosuchids and recovered Fugusuchus hejiapanensis as sister to all other erythrosuchids. Branch support for Erythrosuchidae is low, but the Middle Triassic erythrosuchid clade is well supported by characters with low homoplasy, indicating that their morphology was specialized in ways rarely explored by other Triassic archosauromorphs. [ABSTRACT FROM AUTHOR]

Proterochampsians are a South American endemic group of non-archosaurian archosauriforms with morphological characteristics recollecting Recent crocodilians, and therefore have been proposed as aquatic species. However, this has not been based on careful examination of anatomical and histological features. We provide a review of the morphological and histological evidence present in the skeleton of proterochampsids and discuss its implications for inferring the lifestyles of these organisms. Anatomical features such as a secondary palate, marginal dentition, palatine teeth, morphology of the tail, limb modification, and dermal armor are reviewed, and details of histological structures are described based on bone thin sections. Histological examination reveals a predominance of fibrolamellar bone tissue, suggesting rapid periosteal osteogenesis and therefore overall fast bone growth. The existence of discontinuities (LAGs) demonstrates that these animals responded to changes in their environment. Ecomorphological features do not provide definitive evidence for the lifestyles of proterochampsids, but allow us to propose a terrestrial/amphibious condition. The same is true of the histological features, particularly compactness of the bone. [ABSTRACT FROM AUTHOR]

For the first time, a phylogenetic analysis including all putative euparkeriid taxa is conducted, using a large data matrix analysed with maximum parsimony and Bayesian analysis. Using parsimony, the putative euparkeriid Dorosuchus neoetus from Russia is the sister taxon to Archosauria + Phytosauria. Euparkeria capensis is placed one node further from the crown, and forms a euparkeriid clade with the Chinese taxa Halazhaisuchus qiaoensis and ‘Turfanosuchus shageduensis’ and the Polish taxon Osmolskina czatkowicensis. Using Bayesian methods, Osmolskina and Halazhaisuchus are sister taxa within Euparkeriidae, in turn sister to ‘Turfanosuchus shageduensis’ and then Euparkeria capensis. Dorosuchus is placed in a polytomy with Euparkeriidae and Archosauria + Phytosauria. Although conclusions remain tentative owing to low node support and incompleteness, a broad phylogenetic position close to the base of Archosauria is confirmed for all putative euparkeriids, and the ancestor of Archosauria +Phytosauria is optimized as similar to euparkeriids in its morphology. Ecomorphological characters and traits are optimized onto the maximum parsimony strict consensus phylogeny presented using squared change parsimony. This optimization indicates that the ancestral archosaur was probably similar in many respects to euparkeriids, being relatively small, terrestrial, carnivorous and showing relatively cursorial limb morphology; this Bauplan may have underlain the exceptional radiaton and success of crown Archosauria.

European Russia has yielded several fragmentary but potentially important archosauriform specimens from the Middle Triassic, but these have been only briefly described in the literature. One of these puzzling taxa isDongusuchus efremoviSennikov, 1988, described from the Donguz Svita. We present a redescription ofDongusuchus efremovi, which includes the first photographic atlas and thorough anatomical description of the holotype and referred specimens. This taxon is shown to be a gracile, probably fast-running species with elongate and slender limbs. A phylogenetic analysis recoversDongusuchus efremovias an early-diverging, non-archosaurian archosauriform. Previous work had suggested that this taxon was a ‘rauisuchid’. The gracile proportions of the femur and somewhat wedge-shaped head, however, are unusual for basal archosauriforms and are similar to the plesiomorphic state in crocodile and avian-line crown archosaurs. Several Early-Middle Triassic basal archosauriforms and early members of the crocodile and avian lineages were gracile with elongate, slender limbs. This suggests that the limb morphology ofDongusuchus efremovimay be plesiomorphic for Archosauria and proximal clades. [ABSTRACT FROM AUTHOR]

Here we present archosauriform remains from ‘Abra de los Colorados’, a fossiliferous locality at Sierra de Guayaguas, NW San Luis Province. Two fossiliferous levels were identified in outcrops of the Quebrada del Barro Formation (Norian), which represent the southernmost outcrops of the Marayes–El Carrizal Basin. These levels are composed by massive muddy lithofacies, interpreted as floodplain deposits. The specimens consist of one incomplete maxilla (MIC-V718), one caudal vertebra (MIC-V719), one metatarsal (MIC-V720) and one indeterminate appendicular bone (MIC-V721). The materials can be assigned to Archosauriformes but the fragmentary nature and lack of unambiguous synapomorphies preclude a more precise taxomic assignment. The maxilla is remarkably large and robust and represents the posterior process. It preserved one partially erupted tooth with ziphodont morphology. This bone shows some anatomical traits and size match with ‘rauisuchians’ and theropods. MIC-V719 corresponds to a proximal caudal vertebra. It has a high centrum, a ventral longitudinal furrow, expanded articular processes for the chevrons, a posteriorly displaced diapophysis located below the level of the prezygapophyses, and short prezygapophyses. This vertebra would be from an indeterminate archosauriform. MIC-V720 presents a cylindrical diaphysis, with a well-developed distal trochlea, which present resemblances with metatarsals of theropods, pseudosuchians, and silesaurids, although the size matches better with theropods. MIC-V721 has a slender diaphysis and a convex triangular articular surface, and corresponds to an indeterminate archosauriform. Despite being fragmentary, these materials indicate the presence of a diverse archosauriforms association from Late Triassic beds of San Luis. Thus, they add to the faunal assemblage recently reported from this basin at San Juan Province, which is much rich and diverse than the coeval paleofauna well known from Los Colorados Formation in the Ischigualasto–Villa Union Basin. [ABSTRACT FROM AUTHOR]

The South African species Euparkeria capensis is of great importance for understanding the early radiation of archosauromorphs (including archosaurs) following the Permo–Triassic mass extinction, as most phylogenetic analyses place it as the sister taxon to crown group Archosauria within the clade Archosauriformes. Although a number of species from Lower–Middle Triassic deposits worldwide have been referred to the putative clade Euparkeriidae, the monophyly of Euparkeriidae is controversial and has yet to be demonstrated by quantitative phylogenetic analysis. Three Chinese taxa have been recently suggested to be euparkeriids: Halazhaisuchus qiaoensis, ‘Turfanosuchus shageduensis’, and Wangisuchus tzeyii, all three of which were collected from the Middle Triassic Ermaying Formation of northern China. Here, we reassess the taxonomy and systematics of these taxa. We regard Wangisuchus tzeyii as a nomen dubium, because the holotype is undiagnostic and there is no convincing evidence that the previously referred additional specimens represent the same taxon as the holotype. We also regard ‘Turfanosuchus shageduensis’ as a nomen dubium as we are unable to identify any diagnostic features. We refer the holotype to Archosauriformes, and more tentatively to Euparkeriidae. Halazhaisuchus qiaoensis and the holotype of ‘Turfanosuchus shageduensis’ are resolved as sister taxa in a phylogenetic analysis, and are in turn the sister taxon to Euparkeria capensis, forming a monophyletic Euparkeriidae that is the sister to Archosauria+Phytosauria. This is the first quantitative phylogenetic analysis to recover a non-monospecific, monophyletic Euparkeriidae, but euparkeriid monophyly is only weakly supported and will require additional examination. Given their similar sizes, stratigraphic positions and phylogenetic placement, the holotype of ‘Turfanosuchus shageduensis’ may represent a second individual of Halazhaisuchus qiaoensis, but no apomorphies or unique character combination can be identified to unambiguously unite the two. Our results have important implications for understanding the species richness and palaeobiogeographical distribution of early archosauriforms.

Proterosuchidae is one of the first clades of Archosauriformes (archosaurs and closely related species) to appear in the fossil record, with the richest sample of the group coming from the Lystrosaurus Assemblage Zone (earliest Triassic) of South Africa. Four nominal proterosuchid species were described from South Africa during the twentieth century ( Proterosuchus fergusi, Chasmatosaurus vanhoepeni, Chasmatosaurus alexanderi and Elaphrosuchus rubidgei), but interpretations of their taxonomy have been widely disparate. The most recent taxonomic revision concluded that P. fergusi is the only valid species and that the other nominal species are junior subjective synonyms of this taxon. This proposal was based on the interpretation that anatomical differences between the nominal species could be explained as a result of ontogenetic changes and/or post-mortem deformation. The recent discoveries of multiple new South African proterosuchid specimens provide an impetus to revisit their taxonomy. Based upon a comprehensive re-examination of all known specimens, as well as examination of other proterosuchid taxa in collections worldwide, we conclude that the holotype of Proterosuchus fergusi is undiagnostic. As a result, we propose a neotype ( RC 846) for the species. ' Chasmatosaurus vanhoepeni' and ' Elaphrosuchus rubidgei' are considered subjective junior synonyms of P. fergusi. ' Chasmatosaurus' alexanderi is considered a valid species, for which we propose the new combination P. alexanderi comb. nov. A third species, P. goweri sp. nov., is erected on the basis of a single specimen ( NMQR 880). All three species recognized here are taxonomically distinct from a previously described archosauriform maxilla from the lower Lystrosaurus AZ. As a result, we recognize a minimum of four archosauriform species following the Permo- Triassic mass extinction in South Africa. Our results suggest a greater species richness of earliest Triassic archosauriforms than previously appreciated, but that archosauriform morphological disparity remained low and did not expand until the late Early Triassic - early Mid- Triassic. [ABSTRACT FROM AUTHOR]

Numerous well-preserved skeletal remains of a distinctive armoured archosauriform reptile from the Lower Keuper (Erfurt Formation; Middle Triassic: Ladinian: Longobardian) of Baden-Württemberg (Germany) represent a new taxon,Jaxtasuchus salomonigen. et sp. nov. The dermal armour ofJaxtasuchuscomprises transverse rows of four rectangular osteoderms each in the cervical, dorsal and caudal regions, with the individual plates closely resembling those ofDoswellia kaltenbachifrom Carnian strata in North America. The long and low maxilla ofJaxtasuchusheld at least 15 teeth. The labial and lingual surfaces of the tall, only slightly recurved crowns of the maxillary teeth bear distinct vertical ridges and smooth mesial and distal carinae. The cervical region of the vertebral column is long. Phylogenetic analysis placesJaxtasuchusas the sister taxon toDoswelliain DoswelliidaesensuDesojoet al. (2011). Doswelliidae is diagnosed by the coarsely reticulate, incised ornamentation of osteoderms composed of central regular pits of subequal size and contour, and a mostly smooth anterior articular lamina on each osteoderm. The discovery ofJaxtasuchusconfirms that Doswelliidae had a wide palaeogeographical distribution during the latter half of the Triassic. http://www.zoobank.org/urn:lsid:zoobank.org:pub:4A400628-07F3-4D1B-9CA8-336C379721FA [ABSTRACT FROM PUBLISHER]

Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ' Paleorhinus biochron'. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein ( Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy-based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic. © 2013 The Linnean Society of London [ABSTRACT FROM AUTHOR]

Archosaurs diversified and became dominant during the Mesozoic Era, but their earliest relatives (non-archosaurian archosauromorphs) were already scarcely present in the late Permian. Here we describe a new species of non-archosaurian archosauriform from the upper Permian of Xinjiang, China. Preserved as a partial hindlimb, it possesses a few derived features shared with other archosauriforms, including a much stouter tibia than fibula, a longer metatarsal III than metatarsal IV, and a hooked metatarsal V. Phylogenetic analysis confirmed the new taxon to be a non-archosaurian archosauriform. The morphology of the knee, crus, and pes shows traits that are commonly related with a parasagittal posture, including an entirely proximo-distal articulation of the femur and fibula, the slender and closely spaced tibia and fibula, and a mesaxonic foot with a reduced fifth toe. The new taxon shows that the parasagittal posture evolved before the end-Permian Mass Extinction., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Shansisuchus shansisuchus Young, 1964 was restudied on the basis of a new specimen. Some anatomical features that were either briefly or not described at all in the original study were detailed. The new specimen not only provides further information on the skull anatomy and the vertebral column but also expands the range of the geographical distribution of the taxon. With new information, the diagnosis of S. shansisuchus was emended and its phylogenetic relationships were further analyzed. S. shansisuchus differs from other archosauriforms primarily in the presence of a large subnarial fenestra anterior to the antorbital fenestra, tongue-in-groove articulations between the ascending process of the premaxilla and nasal and between the premaxilla and maxilla, a tall and posterodorsally directed ascending process of the maxilla, a knee-shaped process of the postorbital projecting into the orbit, a broad descending process of squamosal distally well forked and a large, deeply bow-shaped intercentrum tightly anchoring/capping the sharp ventral edges of two neighboring centra together in cervical and at least first eight dorsal vertebrae. With additional information derived from the new specimen, the phylogenetic relationships of S. shansisuchus were reanalyzed; it is closely related to Erythrosuchus-Vjushkovia clade. [ABSTRACT FROM AUTHOR]

The paraphyletic genus ‘Paleorhinus’ is understood currently as a cosmopolitan phytosaur taxon from the Late Triassic. There is no consensus regarding the number of species of ‘Paleorhinus,’ with multiple species and genera synonymised into a single genus or even a single species at various points in its published history. The taxonomy is confounded by historical descriptions without the benefit of comparisons to more recently collected specimens, emphasis on plesiomorphic cranial morphology as diagnostic features of the genus, and lack of cladistic analyses. When included in a recent explicitly cladistic phylogenetic analysis, the holotype of ‘Paleorhinus’ scurriensis (TTU P-00539) was found to be the earliest-branching phytosaur with respect to other North American specimens previously referred to ‘Paleorhinus,’ and is generically distinct from Paleorhinus. ‘Paleorhinus’ scurriensis differs from all known phytosaurs in five unambiguous characters: basitubera widely separated mediolaterally; ridge present on lateral surface of jugal; thickened shelf present along posteroventral edge of expanded pterygoid-quadrate wing; ‘septomaxillae’ separated and excluded from internarial septum; and nasal swelling present posterior to posterior borders of nares. This detailed morphological description of an early-branching phytosaur taxon is a first step towards resolving long-standing issues surrounding specific anatomical features and relationships among early members of the clade. [ABSTRACT FROM PUBLISHER]

Restudy of skulls and available postcrania of the proterochampsian archosauriform Proterochampsa barrionuevoi from the Ischigualasto Formation (Upper Triassic, Carnian) in the San Juan Province, Argentina, confirms that the genus is diagnosed by autapomorphies that include dermal sculpturing consisting of prominent ridges and nodular protuberances, a large hook-like lateral projection on the quadratojugal, an antorbital fossa restricted to a depression along the maxilla, lateral expansion of the premaxilla anterior to the premaxilla-maxilla contact, absence of a supratemporal fossa, exclusion of jugal from suborbital fenestra, basal tubera of parabasisphenoid facing ventrally and reaching laterally beyond the basipterygoid process, and a ventral lamina on the angular. Proterochampsa nodosa is a valid species distinguished from P. barrionuevoi by fewer cranial ridges with larger protuberances, relatively smaller supratemporal fenestrae and width of frontals between orbits less than that of the nasals. A phylogenetic analysis supports the monophyly of Proterochampsia consisting of Proterochampsa, Chanaresuchus bonapartei, Gualosuchus reigi, Tropidosuchus romeri and Cerritosaurus binsfeldi. A temporal separation between the two basal proterochampsians with earliest records in the Late Triassic ( Proterochampsa and Cerritosaurus) and Chanaresuchus, Gualosuchus and Tropidosuchus in the Middle Triassic indicates hidden proterochampsian diversity in the Middle Triassic. [ABSTRACT FROM AUTHOR]

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The two major lineages of extant archosaurs, crocodylians and avians, diverged in the Triassic, but the details and timing of this event are incompletely understood. Fragmentary and phylogenetically uninformative specimens, in addition to poor temporal constraints on rock units from the Early and Middle Triassic, typify obstacles in identifying early archosaurs. This paper re-describes the partial skeleton of the only known specimen of Xilousuchus sapingensis Wu, 1981 from the Early Triassic Heshanggou Formation in north-central China. Originally assigned to the non-archosaurian archosauriform clade Proterosuchidae, an extensive phylogenetic analysis posits X. sapingensis as a crown-group archosaur within Suchia, thus making this taxon the unequivocally oldest known member of Archosauria. The age and phylogenetic position of X. sapingensis indicate that many archosaurs, including all major clades of non-archosaurian archosauriforms, the avianline, ornithosuchids, aetosaurs and paracrocodylomorph lineages, must have diverged by the end of the Early Triassic. X. sapingensis is part of a possible clade of sail-backed poposauroids that were common components of archosaur assemblages during the Early to Middle Triassic. [ABSTRACT FROM PUBLISHER]