Reassessment of the enigmatic "Prestosuchus" loricatus (Archosauria: Pseudosuchia) from the Middle-Late Triassic of southern Brazil.

Our knowledge of the diversity and evolution of South American Triassic pseudosuchians has greatly improved in the past 15 years, due to new discoveries, but also to the revision of several historically important specimens. One of the earliest descriptions of pseudosuchians from the Triassic of Brazil stems from the classic work of Huene from the first half of the 20th century, who described several species, including such influential taxa as Rauisuchus tiradentes or Prestosuchus chiniquensis, which have recently been reviewed. The more poorly known proposed second species of Prestosuchus, P. loricatus, is the focus of the present work. The original material included some elements of the axial skeleton (cervical and caudal vertebrae, ribs, osteoderms) and the hindlimb (ischia, calcaneum, metatarsus), collected from the Dinodontosaurus Assemblage Zone of the Chiniquá area, west of São Pedro do Sul. "Prestosuchus" loricatus shows numerous differences to P. chiniquensis, including the shape of cervical neural spines, presence of epipophyses on the cervical vertebrae, presence of a pit in the iliac articulation of the ischium, lack of longitudinal furrows in the dorsolateral surface of the ischial shafts, the more slender calcaneal tuber and a less pronounced ventral pit in the calcaneum, and is thus referred to a new genus, Schultzsuchus gen. nov. Phylogenetic analysis indicates an early branching position within Poposauroidea for Schultzsuchus, making it the oldest known member of this clade in South America.

A new early-diverging probainognathian cynodont and a revision of the occurrence of cf. Aleodon from the Chañares Formation, northwestern Argentina: New clues on the faunistic composition of the latest Middle-?earliest Late Triassic Tarjadia Assemblage Zone.

The Chañares Formation (Ischigualasto-Villa Unión Basin) is worldwide known by its exquisitely preserved fossil record of latest Middle-to-early Late Triassic tetrapods, including erpetosuchids, "rauisuchians," proterochampsids, gracilisuchids, dinosauromorphs, pterosauromorphs, kannemeyeriiform dicynodonts, and traversodontid, chiniquodontid and probainognathid cynodonts, coming from the Tarjadia (bottom) and Massetognathus-Chanaresuchus (top) Assemblage Zones of its lower member. Regarding cynodonts, most of its profuse knowledge comes from the traditional layers discovered by Alfred Romer and his team in the 1960s that are now enclosed in the Massetognathus-Chanaresuchus Assemblage Zone (AZ). In this contribution we focus our study on the probainognathian cynodonts discovered in levels of the Tarjadia Assemblage Zone. We describe a new chiniquodontid cynodont with transversely broad postcanine teeth (Riojanodon nenoi gen. et sp. nov.) which is related to the genus Aleodon. In addition, the specimen CRILAR-Pv 567 previously referred to cf. Aleodon is here described, compared, and included in a phylogenetic analysis. It is considered as an indeterminate Aleodontinae nov., a clade here proposed to included chiniquodontids with transversely broad upper and lower postcanines, by having a cuspidated sectorial labial margin and a lingual platform that is twice broader than a lingual cingulum. Cromptodon mamiferoides, from the Cerro de Las Cabras Formation (Cuyo Basin), was also included in the phylogenetic analysis and recovered as an Aleodontinae. The new cynodont and the record of Aleodontinae indet. reinforce the faunal differentiation between the Tarjadia and Massetognathus-Chanaresuchus Assemblage Zones, in the lower member of the Chañares Formation, and inform on the diverse chiniquodontid clade with both sectorial and transversely broad postcanine teeth.

Paleoneurology and neuroanatomical notes on the South American "rauisuchian" Saurosuchus galilei from the Ischigualasto formation, San Juan, Argentina.

Non-crocodylomorph loricatans, traditionally known as "rauisuchians," are considered as the top predators of the Triassic continental faunas that reigned before the emergence of the well-known theropod dinosaurs. In particular, Saurosuchus galilei is a large quadrupedal prestosuchid loricatan found in the Ischigualasto Formation from northwestern Argentina. Here, we reevaluated the braincase of S. galilei and present the first paleoneurological study based on the partial natural casts of the holotype and the digital cranial endocast of the referred specimen PVSJ 32. The braincase of S. galilei was here reinterpreted, identifying the sutures of the supraoccipital, otoccipital, basioccipital, parabasisphenoid, prootic, and laterosphenoid. A unique feature identified in the braincase of S. galilei is the presence of deep paracondylar recesses associated with pharyngotympanic pneumaticity, which has not been identified in any other "rauisuchians" so far. Most of the structures of the encephalon were recognized in the cranial endocast and natural casts including cranial nerves V-XII, olfactory bulbs, main blood vessels and sinuses, and inner ear. These structures allowed us to quantify some of its sensorial capacities and recognize that S. galilei had an enhanced olfactory acuity, with coefficients higher than those expected for its body size, a condition previously observed in living crocodilians and tyrannosaurid dinosaurs. An improved sense of smell might have allowed Saurosuchus to track its prey from long distances and compensate for the poorer development of its other senses like vision and balance.

Palaeobiological inferences of "rauisuchians" Fasolasuchus tenax (Los Colorados Fm., Argentina) and Prestosuchus chiniquensis (Santa Maria Super sequence, Brazil) from the Middle-Upper Triassic of South America based on microstructural analyses.

"Rauisuchia" is a non-monophyletic group of quadrupedal and carnivorous pseudosuchians that inhabited the entire world during the Middle-Upper Triassic period (Anisian/Ladinian-Rhaetian). In South America, "rauisuchians" reached the largest sizes among continental carnivores. Despite their important ecological role, some aspects of their palaeobiology have been poorly examined. Here, we study appendicular bones, dorsal ribs and osteoderms of two genera, the Argentinean Fasolasuchus tenax (PVL 3850, holotype) and the Brazilian Prestosuchus chiniquensis (SNSB-BSPG AS XXV) respectively. The femur of F. tenax is formed by laminar fibrolamellar bone, which is composed of non-fully monorefringent woven-fibred matrix and primary osteons; the dorsal rib has a Haversian bone composition with an external fundamental system recorded and the osteoderm is formed by well-organised parallel-fibred bone. The femur, humerus and fibula of P. chiniquensis are mostly composed of strongly arranged parallel-fibred bone and a laminar vascularisation. The minimal ages obtained correspond to 9 years for F. tenax (based on the maximum number of growth marks in the osteoderm) and 4 years for P. chiniquensis (obtained from the highest count of growth marks in the femur and in the humerus). F. tenax attained somatic and skeletal maturity, while P. chiniquensis was near to reaching skeletal and sexual maturity, but it was somatically immature. The overall rapid growth rate and the high and uniform vascularisation seems to imply that these features are common in most of "rauisuchians", except in P. chiniquensis.

New digital braincase endocasts of two species of Desmatosuchus and neurocranial diversity within Aetosauria (Archosauria: Pseudosuchia).

In the present contribution we revise, figure, and redescribe several isolated braincases of the iconic aetosaur Desmatosuchus from the Placerias Quarry locality, Chinle Formation, Arizona, United States. The detailed study of the isolated braincases from the UCMP collection allowed us to assign them at the species-level and recognize two species of Desmatosuchus for the Placerias Quarry: D. spurensis and D. smalli. The former can be distinguished from the latter by the presence of a transverse sulcus on the parietals, deep median pharyngeal recess on the basisphenoid, almost no gap between the basal tubera and the basipterygoid processes, and the exoccipitals meeting at the midline. The presence of D. smalli at the Placerias Quarry has not been previously reported. Based on the braincases UCMP 27408, 27410, 27407, three new brain endocasts were developed through CT scan images, reconstructing the most complete endocranial casts known for an aetosaur, including the encephalon, cranial nerves, inner ear, and endocranial vasculature. The cranial endocasts also exhibited some differences between both species of Desmatosuchus, with D. spurensis having a distinguishable dural expansion and markedly asymmetric anterior and posterior semicircular canals of the labyrinth. Additionally, the combination of osteological features and the endocranial casts allowed us to identify and discuss the presence of an ossified orbitosphenoid on the anteriormost region of the braincase among aetosaurs. Furthermore, we were able to reinterpret some of the observations made by previous authors on the endocast of the holotype of Desmatosuchus spurensis (UMMP VP 7476) and provide some insight into their neurosensory capabilities.

The Late Triassic Ischigualasto Formation at Cerro Las Lajas (La Rioja, Argentina): fossil tetrapods, high-resolution chronostratigraphy, and faunal correlations.

Present knowledge of Late Triassic tetrapod evolution, including the rise of dinosaurs, relies heavily on the fossil-rich continental deposits of South America, their precise depositional histories and correlations. We report on an extended succession of the Ischigualasto Formation exposed in the Hoyada del Cerro Las Lajas (La Rioja, Argentina), where more than 100 tetrapod fossils were newly collected, augmented by historical finds such as the ornithosuchid Venaticosuchus rusconii and the putative ornithischian Pisanosaurus mertii. Detailed lithostratigraphy combined with high-precision U-Pb geochronology from three intercalated tuffs are used to construct a robust Bayesian age model for the formation, constraining its deposition between 230.2 ± 1.9 Ma and 221.4 ± 1.2 Ma, and its fossil-bearing interval to 229.20 + 0.11/- 0.15-226.85 + 1.45/- 2.01 Ma. The latter is divided into a lower Hyperodapedon and an upper Teyumbaita biozones, based on the ranges of the eponymous rhynchosaurs, allowing biostratigraphic correlations to elsewhere in the Ischigualasto-Villa Unión Basin, as well as to the Paraná Basin in Brazil. The temporally calibrated Ischigualasto biostratigraphy suggests the persistence of rhynchosaur-dominated faunas into the earliest Norian. Our ca. 229 Ma age assignment to Pi. mertii partially fills the ghost lineage between younger ornithischian records and the oldest known saurischians at ca. 233 Ma.

The evolution of dermal shield vascularization in Testudinata and Pseudosuchia: phylogenetic constraints versus ecophysiological adaptations.

Studies on living turtles have demonstrated that shells are involved in the resistance to hypoxia during apnea via bone acidosis buffering; a process which is complemented with cutaneous respiration, transpharyngeal and cloacal gas exchanges in the soft-shell turtles. Bone acidosis buffering during apnea has also been identified in crocodylian osteoderms, which are also known to employ heat transfer when basking. Although diverse, many of these functions rely on one common trait: the vascularization of the dermal shield. Here, we test whether the above ecophysiological functions played an adaptive role in the evolutionary transitions between land and aquatic environments in both Pseudosuchia and Testudinata. To do so, we measured the bone porosity as a proxy for vascular density in a set of dermal plates before performing phylogenetic comparative analyses. For both lineages, the dermal plate porosity obviously varies depending on the animal lifestyle, but these variations prove to be highly driven by phylogenetic relationships. We argue that the complexity of multi-functional roles of the post-cranial dermal skeleton in both Pseudosuchia and Testudinata probably is the reason for a lack of obvious physiological signal, and we discuss the role of the dermal shield vascularization in the evolution of these groups. This article is part of the theme issue 'Vertebrate palaeophysiology'.

New Specimen Sheds Light on the Anatomy and Taxonomy of the Early Late Triassic Dinosauriforms from the Chañares Formation, NW Argentina.

The lower Carnian levels of the Chañares Formation (Ischigualasto-Villa Unión Basin, La Rioja Province) have yielded some of the most informative dinosaur precursor species known so far. However, these species are based on partial skeletons that in several cases hamper the comparison between them because of the absence of overlapping bones. This has generated a contentious debate during the last 20 years about the synonymy between two of these species, Lewisuchus admixtus and Pseudolagosuchus major. Here, we describe a new dinosauriform partial skeleton (CRILAR-Pv 552) recently collected in the Chañares Formation that preserves previously unknown anatomical regions for the dinosaur precursors of this unit (e.g., premaxilla, inner ear, anterior zeugopodium) and allows comparisons with other dinosauriform specimens. CRILAR-Pv 552 is referred to Lewisuchus admixtus because it possesses a proportionally large skull, a laterally projected, shelf-like ridge on the jugal, and recurved, finely serrated middle-posterior maxillary and dentary teeth ankylosed to the bone, and the absence of a coracoid foramen. The new specimen preserves a dorsally bowed dentary with a lateroventral shelf that is identical to a dentary associated with the holotype of Lewisuchus admixtus. Additionally, the morphology of the new specimen is completely congruent with that of specimens of Pseudolagosuchus major, bolstering the hypothesis that the latter species is a subjective junior synonym of Lewisuchus admixtus. A preliminary phylogenetic analysis with updated scorings for Lewisuchus admixtus found this species at the base of Silesauridae. Anat Rec, 303:1393-1438, 2020. © 2019 American Association for Anatomy.

Deep faunistic turnovers preceded the rise of dinosaurs in southwestern Pangaea.

The Triassic period documents the origin and diversification of modern amniote lineages and the Late Triassic fossil record of South America has been crucial to shed light on these early evolutionary histories. However, the faunistic changes that led to the establishment of Late Triassic ecosystems are largely ignored because of the global scarcity of fossils from assemblages a few million years older. Here we contribute to fill this gap with the description of a new tetrapod assemblage from the lowermost levels of the Chañares Formation (uppermost Middle-lower Late Triassic epochs) of Argentina, which is older than the other vertebrate assemblages of the same basin. The new assemblage is composed of therapsids, rhynchosaurids and archosaurs, and clearly differs from that of the immediately overlying and well-known historical Chañares vertebrate assemblage. The new tetrapod association is part of a phase of relatively rapidly changing vertebrate assemblage compositions, in a time span shorter than 6 million years, before the diversification of dinosaurs and other common Late Triassic tetrapods in southwestern Pangaea.

Osteoderm histology of Proterochampsia and Doswelliidae (Reptilia: Archosauriformes) and their evolutionary and paleobiological implications.

Postcranial osteoderms are commonly developed in the major lineages of Archosauriformes, including forms such as proterochampsids and doswelliids. Here, we survey the histology of osteoderms of the doswelliids Archeopelta arborensis and Tarjadia ruthae, and the proterochampsids Chanaresuchus bonapartei and Pseudochampsa ischigualastensis to understand better the morphogenesis of these skeletal elements. Whereas, the Doswelliid osteoderms possess a trilaminar organization, in which two cortices (external and basal) can be differentiated from an internal core of cancellous bone, these elements are compact structures in proterochampsids. The osteoderms of P. ischigualastensis are avascular and they consist entirely of parallel-fibered bone. Conversely, the osteoderms of C. bonapartei are well vascularized structures composed of zones of woven-fibered bone and annuli of parallel-fibered bone. The rather simple microstructure observed in P. ischigualastensis osteoderms suggests that these elements grew at a constant, low rate. Compared with proterochampsids, doswelliid osteoderms possess a more complex histology, which appears to be linked to variations in the growth rate during the osteoderm formation and also to the development of the external ornamentation. A comparison of our findings with the results of earlier studies on other archosauriforms (phytosaurs and pseudosuchians) reveals that the general osteoderm histology of doswelliids bears a closer resemblance to that of phytosaurs and pseudosuchians than the proterochampsid osteoderm microstructure. If all archosauriform osteoderms are homologous structures, the closer resemblance of doswellid osteoderm microstructures to that of phytosaurs and pseudosuchians is in agreement with the hypothesis that doswellids are more closely related to archosaurs than proterochampsids. J. Morphol. 276:385-402, 2015. © 2015 Wiley Periodicals, Inc.

A new aetosaur from the Upper Triassic of the Santa Maria Formation, southern Brazil.

Aetosaurs are armored pseudosuchian archosaurs widespread in Upper Triassic units. In South America, four taxa were previously recorded: Aetosauroides scagliai, Neoaetosauroides engaeus, Aetobarbakinoides brasiliensis, and Chilenosuchus forttae. Herein we describe a new Late Triassic juvenile aetosaur from the Santa Maria Formation of southern Brazil, Polesinesuchus aurelioi gen. et sp. nov., increasing the paleobiodiversity of this interesting group to five taxa in Western Gondwana. The holotype is composed of cranial (parietal and braincase) and postcranial elements (cervical, dorsal, sacral, caudal vertebrae, both scapulae, a humerus, ilium, pubis, ischium, tibia, a partial right pes, and anterior and mid-dorsal paramedian osteoderms). It belongs to a juvenile individual, as its neurocentral sutures are open in all vertebrae, and also due to its small size. However, future paleohistological investigation is necessary to fully corroborate this assumption. This new taxon is distinguished from all other aetosaurs by the presence of an unique combination of character states (not controlled by ontogeny) such as: cervical vertebrae with prezygapophyses widely extending laterally through most of the anterior edge of the diapophyses; absence of hyposphene articulations in both cervical and mid-dorsal vertebrae; presence of a ventral keel in cervical vertebrae; anterior and mid-dorsal vertebrae without a lateral fossa in their centra; expanded proximal end of scapula; anteroposteriorly expanded medial portion of scapular blade; a short humerus with a robust shaft; and a dorsoventral and very low iliac blade with a long anterior process which slightly exceeds the pubic peduncle. Regarding its phylogenetic relationships, the present analysis placed Polesinesuchus as the sister taxon of Aetobarbakinoides and both as sister taxa of the unnamed monophyletic clade Desmatosuchinae plus Typothoracisinae. 

Bone histology of phytosaur, aetosaur, and other archosauriform osteoderms (Eureptilia, Archosauromorpha).

As in other archosauriforms, phytosaurs and aetosaurs are characterized by the presence of well-developed osteoderms. Here we provide a comparative study on the microstructure of phytosaur (five taxa) and aetosaur (thirteen taxa) osteoderms. For outgroup comparison, we sampled osteoderms of the sister taxon to Aetosauria, Revueltosaurus callenderi, and the doswelliid Jaxtasuchus salomoni. Phytosaur, aetosaur, and Jaxtasuchus osteoderms are composed of a diploe structure, whereas the Revueltosaurus osteoderm microanatomy is more compact. The external cortex of phytosaurs, Revueltosaurus and Jaxtasuchus osteoderms is mainly composed of parallel-fibered bone. In aetosaurs, the external cortex mainly consists of lamellar bone, with lines of resorption within the primary bone indicating successive cycles of bone erosion and deposition. The basal cortex in all the specimens is composed of parallel-fibered bone, with the cancellous internal core being more strongly developed in aetosaurs than in phytosaurs. Woven or fibro-lamellar bone was recorded in both phytosaurian and aetosaurian taxa, as well as in Jaxtasuchus. Structural fibers, which at least partly suggest metaplastic origin, were only recorded in the internal core of two phytosaurs and in the basal cortex of one aetosaur. Osteoderm thickness and cancellous to compact bone ratios appear to be subject to ontogenetic change. Minimum growth mark counts in osteoderms sampled indicate that some aetosaurs and phytosaurs lived for at least two decades. Bone microstructures are more uniform in phytosaur osteoderms and show a higher level of disparity among aetosaur osteoderms, and at least in the latter, histological features are potentially apomorphic for species/genus level.

The oldest known communal latrines provide evidence of gregarism in Triassic megaherbivores.

Defecation in communal latrines is a common behaviour of extant mammals widely distributed among megaherbivores. This behaviour has key social functions with important biological and ecological implications. Herbivore communal latrines are only documented among mammals and their fossil record is exceptionally restricted to the late Cenozoic. Here we report the discovery of several massive coprolite associations in the Middle-Late Triassic of the Chañares Formation, Argentina, which represent fossil communal latrines based on a high areal density, small areal extension and taphonomic attributes. Several lines of evidence (size, morphology, abundance and coprofabrics) and their association with kannemeyeriiform dicynodonts indicate that these large synapsids produced the communal latrines and had a gregarious behaviour comparable to that of extant megaherbivores. This is the first evidence of megaherbivore communal latrines in non-mammal vertebrates, indicating that this mammal-type behaviour was present in distant relatives of mammals, and predates its previous oldest record by 220 Mya.

The cranial osteology and feeding ecology of the metriorhynchid crocodylomorph genera Dakosaurus and Plesiosuchus from the late Jurassic of Europe.

BACKGROUND: Dakosaurus and Plesiosuchus are characteristic genera of aquatic, large-bodied, macrophagous metriorhynchid crocodylomorphs. Recent studies show that these genera were apex predators in marine ecosystems during the latter part of the Late Jurassic, with robust skulls and strong bite forces optimized for feeding on large prey. METHODOLOGY/PRINCIPAL FINDINGS: Here we present comprehensive osteological descriptions and systematic revisions of the type species of both genera, and in doing so we resurrect the genus Plesiosuchus for the species Dakosaurus manselii. Both species are diagnosed with numerous autapomorphies. Dakosaurus maximus has premaxillary 'lateral plates'; strongly ornamented maxillae; macroziphodont dentition; tightly fitting tooth-to-tooth occlusion; and extensive macrowear on the mesial and distal margins. Plesiosuchus manselii is distinct in having: non-amblygnathous rostrum; long mandibular symphysis; microziphodont teeth; tooth-crown apices that lack spalled surfaces or breaks; and no evidence for occlusal wear facets. Our phylogenetic analysis finds Dakosaurus maximus to be the sister taxon of the South American Dakosaurus andiniensis, and Plesiosuchus manselii in a polytomy at the base of Geosaurini (the subclade of macrophagous metriorhynchids that includes Dakosaurus, Geosaurus and Torvoneustes). CONCLUSIONS/SIGNIFICANCE: The sympatry of Dakosaurus and Plesiosuchus is curiously similar to North Atlantic killer whales, which have one larger 'type' that lacks tooth-crown breakage being sympatric with a smaller 'type' that has extensive crown breakage. Assuming this morphofunctional complex is indicative of diet, then Plesiosuchus would be a specialist feeding on other marine reptiles while Dakosaurus would be a generalist and possible suction-feeder. This hypothesis is supported by Plesiosuchus manselii having a very large optimum gape (gape at which multiple teeth come into contact with a prey-item), while Dakosaurus maximus possesses craniomandibular characteristics observed in extant suction-feeding odontocetes: shortened tooth-row, amblygnathous rostrum and a very short mandibular symphysis. We hypothesise that trophic specialisation enabled these two large-bodied species to coexist in the same ecosystem.

Tooth-on-tooth interlocking occlusion suggests macrophagy in the mesozoic marine crocodylomorph dakosaurus.

Metriorhynchidae was a peculiar but long-lived group of marine Mesozoic crocodylomorphs adapted to a pelagic lifestyle. Recent discoveries show that metriorhynchids evolved a wide range of craniodental morphotypes and inferred feeding strategies. One genus, Dakosaurus, is arguably the most aberrant marine crocodylomorph due to its large, robust, ziphodont teeth; very low tooth count; and brevirostrine/oreinirostral snout. We here report an additional unusual feature of Dakosaurus that is unique among marine crocodylomorphs: tightly fitting tooth-to-tooth occlusion, whose inference is supported by reception pits along the upper and lower tooth rows, indicative of vertically orientated crowns that were in close contact during occlusion, and three distinct types of dental wear. These include irregular spalled surfaces near the apex (probably caused by tooth-food contact), semi-circular wear near the base, and elongate surfaces extending along the mesial and distal margins of the teeth, obliterating the carinae (including the denticles). Scanning electron micrographs show that these latter surfaces are marked by parallel apicobasal striations, which in extant mammals reflect tooth-tooth contact. As such, we interpret the carinal wear facets in Dakosaurus as being formed by repeated tooth-tooth contact between the mesial and distal margins of the teeth of the upper and lower jaw. We posit that this increased the available shearing surface on their high crowns. Together, these wear patterns suggest that occlusion in Dakosaurus was specialized for cutting large and abrasive prey items into portions small enough to swallow, making it a prime example of an aquatic reptile with macrophagous feeding habits.