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A dearth of Mesozoic-aged, three-dimensional fossils hinders understanding of the origin of the distinctive skull and brain of modern (crown) birds 1 . Here we report Navaornis hestiae gen. et sp. nov., an exquisitely preserved fossil species from the Late Cretaceous of Brazil. The skull of Navaornis is toothless and large-eyed, with a vaulted cranium closely resembling the condition in crown birds; however, phylogenetic analyses recover Navaornis in Enantiornithes, a highly diverse clade of Mesozoic stem birds. Despite an overall geometry quantitatively indistinguishable from crown birds, the skull of Navaornis retains numerous plesiomorphies including a maxilla-dominated rostrum, an akinetic palate, a diapsid temporal configuration, a small cerebellum and a weakly expanded telencephalon. These archaic neurocranial traits are combined with a crown bird-like degree of brain flexion and a bony labyrinth comparable in shape to those of many crown birds but substantially larger. Altogether, the emergent cranial geometry of Navaornis shows an unprecedented degree of similarity between crown birds and enantiornithines, groups last sharing a common ancestor more than 130 million years ago 2 . Navaornis provides long-sought insight into the detailed cranial and endocranial morphology of stem birds phylogenetically crownward of Archaeopteryx, clarifying the pattern and timing by which the distinctive neuroanatomy of living birds was assembled., Competing Interests: Competing interests The authors declare no competing interests., (© 2024. The Author(s).)

Rhynchosauria is a group of extinct, exclusively Triassic, terrestrial, and herbivorous archosauromorphs, characterized by a peculiar maxillary-mandibular apparatus. They reached global distribution during the Carnian, with the Hyperodapedontinae clade. The rhynchosaurian record from South America is included in the Ladinian-?earliest Carnian Dinodontosaurus Assemblage Zone (Pinheiros-Chiniquá Sequence, Brazil) and Tarjadia Assemblage Zone (Chañares Formation, Argentina), and for strictly Carnian Hyperodapedon Assemblage Zone (Lower Candelária Sequence, Brazil) and Ischigualasto Formation (Argentina). Here, we present the first record of Hyperodapedontinae from the Brazilian Early Carnian Santacruzodon Assemblage Zone (Santa Cruz Sequence). The most diagnostic elements belong to a nearly complete left pes, which presents a combination of hyperodapedontine traits. The presence of a second individual of cf. Hyperodapedontinae is based on a fragmentary, isolated left metatarsal IV. An isolated fragment of premaxilla could belong to the second specimen (based on preservation mode) or to a third individual. This new report fills a gap within the South American rhynchosaurian distribution, strengthening biostratigraphic correlation with other regions from Gondwana (i.e., Madagascar), where similar and coeval faunas are known. The inclusion of these specimens in a phylogenetic dataset resulted in low resolution results, due to missing data because postcranial characters for rhynchosaurs are still poorly explored., (© 2024 American Association for Anatomy.)

The acquisition of the load-bearing dentary-squamosal jaw joint was a key step in mammalian evolution 1-5 . Although this innovation has received decades of study, questions remain over when and how frequently a mammalian-like skull-jaw contact evolved, hindered by a paucity of three-dimensional data spanning the non-mammaliaform cynodont-mammaliaform transition. New discoveries of derived non-mammaliaform probainognathian cynodonts from South America have much to offer to this discussion. Here, to address this issue, we used micro-computed-tomography scanning to reconstruct the jaw joint anatomy of three key probainognathian cynodonts: Brasilodon quadrangularis, the sister taxon to Mammaliaformes 6-8 , the tritheledontid-related Riograndia guaibensis 9 and the tritylodontid Oligokyphus major. We find homoplastic evolution in the jaw joint in the approach to mammaliaforms, with ictidosaurs (Riograndia plus tritheledontids) independently evolving a dentary-squamosal contact approximately 17 million years before this character first appears in mammaliaforms of the Late Triassic period 10-12 . Brasilodon, contrary to previous descriptions 6-8 , lacks an incipient dentary condyle and squamosal glenoid and the jaws articulate solely using a plesiomorphic quadrate-articular joint. We postulate that the jaw joint underwent marked evolutionary changes in probainognathian cynodonts. Some probainognathian clades independently acquired 'double' craniomandibular contacts, with mammaliaforms attaining a fully independent dentary-squamosal articulation with a conspicuous dentary condyle and squamosal glenoid in the Late Triassic. The dentary-squamosal contact, which is traditionally considered to be a typical mammalian feature, therefore evolved more than once and is more evolutionary labile than previously considered., (© 2024. The Author(s).)

Investigating the evolutionary trajectory of synapsid sensory and cephalic systems is pivotal for understanding the emergence and diversification of mammals. Recent studies using CT-scanning to analyze the rostral foramina and maxillary canals morphology in fossilized specimens of probainognathian cynodonts have contributed to clarifying the homology and paleobiological interpretations of these structures. In the present work, μCT-scannings of three specimens of Riograndia guaibensis, an early Norian cynodont from southern Brazil, were analyzed and revealed an incomplete separation between the lacrimal and maxillary canals, with points of contact via non-ossified areas. While the maxillary canal exhibits a consistent morphological pattern with other Prozostrodontia, featuring three main branches along the lateral region of the snout, the rostral alveolar canal in Riograndia displays variability in the number of extra branches terminating in foramina on the lateral surface of the maxilla, showing differences among individuals and within the same skull. Additionally, pneumatization is observed in the anterior region of the skull, resembling similar structures found in reptiles and mammals. Through this pneumatization, certain branches originating from the maxillary canal extend to the canine alveolus. Further investigation is warranted to elucidate the functionality of this structure and its occurrence in other cynodont groups., (© 2024 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The nasal cavity of living mammals is a unique structural complex among tetrapods, acquired along a series of major morphological transformations that occurred mainly during the Mesozoic Era, within the Synapsida clade. Particularly, non-mammaliaform cynodonts document several morphological changes in the skull, during the Triassic Period, that represent the first steps of the mammalian bauplan. We here explore the nasal cavity of five cynodont taxa, namely Thrinaxodon, Chiniquodon, Prozostrodon, Riograndia, and Brasilodon, in order to discuss the main changes within this skull region. We did not identify ossified turbinals in the nasal cavity of these taxa and if present, as non-ossified structures, they would not necessarily be associated with temperature control or the development of endothermy. We do, however, notice a complexification of the cartilage anchoring structures that divide the nasal cavity and separate it from the brain region in these forerunners of mammals., (© 2024. The Author(s).)

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

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., (© 2024 American Association for Anatomy.)

Body size influences most aspects of an animal's biology, consequently, evolutionary diversification is often accompanied by differentiation of body sizes within a lineage. It is accepted that miniaturization, or the evolution of extremely small body sizes, played a key role in the origin and early evolution of different mammalian characters in non-mammaliaform cynodonts. However, while there are multiple studies on the biomechanical, behavioral, and physiological consequences of smaller sizes, few explore the evolutionary processes that lead to them. Here, we use body mass as a universal size measurement in phylogenetic comparative analyses to explore aspects of body size evolution in Cynodontia, focusing on the cynodont-mammal transition, and test the miniaturization hypothesis for the origin of Mammaliaformes. We estimated the body masses of 29 species, ranging from Theriocephalia to Mammaliaformes, providing the largest collection of Triassic cynodont body mass estimates that we know of, and used these estimates in analyses of disparity through time and RR phylo . Unexpectedly, our results did not support the miniaturization hypothesis. Even though cynodont body size disparity fell during the Late Triassic, and remained lower than expected under a purely Brownian motion model of evolution up until the Early Jurassic, we found that rates of body size evolution were significantly lower in prozostrodontians leading to the first Mammaliaformes than in other lineages. Evolution rates were higher in medium and large-sized taxa, indicating that size was changing more rapidly in those lineages and that small sizes were probably a persistent plesiomorphic character-state in Cynodontia., (© 2024 American Association for Anatomy.)

Prozostrodon brasiliensis and Therioherpeton cargnini are non-mammaliaform cynodonts that lived ~233 million years ago (late Carnian, Late Triassic) in western Gondwana. They represent some of the earliest divergent members of the clade Prozostrodontia, which includes "tritheledontids", tritylodontids, "brasilodontids", and mammaliaforms (including Mammalia as crown group). Here, we studied the endocranial anatomy (cranial endocast, nerves, vessels, ducts, ear region, and nasal cavity) of these two species. Our findings suggest that during the Carnian, early prozostrodonts had a brain with well-developed olfactory bulbs, expanded cerebral hemispheres divided by the interhemispheric sulcus, and absence of an unossified zone and pineal body. The morphology of the maxillary canal represents the necessary condition for the presence of facial vibrissae. A slight decrease in encephalization is observed at the origin of the clade Prozostrodontia. This new anatomical information provides evidence for the evolution of endocranial traits of the first prozotrodonts, a Late Triassic lineage that culminated in the origin of mammals., (© 2023 American Association for Anatomy.)

In this contribution we describe a new genus and species of gomphodontosuchine cynodont from the Late Triassic Hyperodapedon Assemblage Zone (AZ) of the Santa Maria Supersequence of southern Brazil, based on material collected 20 years ago. The new taxon, Santagnathus mariensis gen. et sp. nov., is based on numerous cranial and postcranial remains, which altogether provide data on several parts of the skeleton. Santagnathus mariensis is closely related to Siriusgnathus niemeyerorum and Exaeretodon spp. and expands the knowledge about gomphodontosuchine cynodonts. Morphologically, the new species has a skull bauplan particularly similar to S. niemeyerorum and E. riograndensis, differing from them by a unique combination of traits such as three upper incisors, the absence of a descending process in the jugal, a more posterior position of the postorbital bar and a preorbital region larger than the temporal one. The new traversodontid was found associated with the rhynchosaur Hyperodapedon sp., corroborating the assignment of the cynodont fossils to the Hyperodapedon AZ. We also provide comments on the status of the Argentine traversodontid cynodont Proexaeretodon vincei, typically considered a junior synonym of Exaeretodon argentinus, and here accepted as a valid taxon., (© 2023 American Association for Anatomy.)

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 description of new titanosaur specimens unearthed from deposits of the Serra da Galga Formation (Bauru Group, Late Cretaceous) at the BR-262 site, near Peirópolis (Uberaba, Minas Gerais State, Brazil), sheds light on the taxonomy of two taxa previously known from the same area and geological unit: Baurutitan britoi and Trigonosaurus pricei . A comparative revision indicates that T. pricei represents a junior synonym of Ba. britoi , and that the BR-262 specimens belong to that latter species. The information provided by the new specimens also revealed that the paratype of T. pricei (MCT 1719-R), a caudal vertebral series, actually represents a new taxon, named here as Caieiria allocaudata gen. et sp. nov., Competing Interests: The authors declare that they have no competing interests., (© 2022 Silva Junior et al.)

Irajatherium hernandezi is a poorly known non-mammaliaform cynodont from the Late Triassic of southern Brazil. A new specimen of this cynodont was found in recent fieldwork to the type-locality, Sesmaria do Pinhal (Candelária), providing new insights into the anatomy of this mammalian forerunner. This specimen comprises a partial skull preserving the left canine, two left and three right postcanines, and an isolated exoccipital; the left dentary with the canine and postcanines; a fragment of the right dentary; the proximal portion of the left partial humerus; the right scapula; and indeterminate fragments. Based on new material, it is here suggested that I. hernandezi presents: a rostrum broad and short, possibly long as the temporal region; three foramina on the lateral surface of the maxilla, that could correspond to the external openings of the rostral alveolar, infraorbital, and zygomaticofacial canals; a slender zygomatic arch and an absent postorbital bar; a posteriorly wide temporal fossa; a long secondary palate, slightly surpassing the level of the last postcanine tooth; the cerebral hemispheres of the cranial endocast divided by a median sulcus; the scapular blade long and straight, and the postscapular fossa absent in lateral aspect. Finally, I. hernandezi and other tritheledontids were included in a phylogenetic analysis of Eucynodontia. The analysis recovered unresolved relationships for ictidosaurs/tritheledontids, nested within a polytomy with Tritylodontidae and a clade composed by Pseudotherium argentinus, Botucaraitherium belarminoi, Brasilodon quadrangularis, and Mammaliaformes., (© 2021 American Association for Anatomy.)

Among terrestrial vertebrates, only crown birds (Neornithes) rival mammals in terms of relative brain size and behavioural complexity. Relatedly, the anatomy of the avian central nervous system and associated sensory structures, such as the vestibular system of the inner ear, are highly modified with respect to those of other extant reptile lineages. However, a dearth of three-dimensional Mesozoic fossils has limited our knowledge of the origins of the distinctive endocranial structures of crown birds. Traits such as an expanded, flexed brain, a ventral connection between the brain and spinal column, and a modified vestibular system have been regarded as exclusive to Neornithes. Here, we demonstrate all of these 'advanced' traits in an undistorted braincase from an Upper Cretaceous enantiornithine bonebed in southeastern Brazil. Our discovery suggests that these crown bird-like endocranial traits may have originated prior to the split between Enantiornithes and the more crownward portion of avian phylogeny over 140 Ma, while coexisting with a remarkably plesiomorphic cranial base and posterior palate region. Altogether, our results support the interpretation that the distinctive endocranial morphologies of crown birds and their Mesozoic relatives are affected by complex trade-offs between spatial constraints during development.

Endothermy underpins the ecological dominance of mammals and birds in diverse environmental settings 1,2 . However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most of the fossil evidence is ambiguous 3-17 . Here we show that this key evolutionary transition can be investigated using the morphology of the endolymph-filled semicircular ducts of the inner ear, which monitor head rotations and are essential for motor coordination, navigation and spatial awareness 18-22 . Increased body temperatures during the ectotherm-endotherm transition of mammal ancestors would decrease endolymph viscosity, negatively affecting semicircular duct biomechanics 23,24 , while simultaneously increasing behavioural activity 25,26 probably required improved performance 27 . Morphological changes to the membranous ducts and enclosing bony canals would have been necessary to maintain optimal functionality during this transition. To track these morphofunctional changes in 56 extinct synapsid species, we developed the thermo-motility index, a proxy based on bony canal morphology. The results suggest that endothermy evolved abruptly during the Late Triassic period in Mammaliamorpha, correlated with a sharp increase in body temperature (5-9 °C) and an expansion of aerobic and anaerobic capacities. Contrary to previous suggestions 3-14 , all stem mammaliamorphs were most probably ectotherms. Endothermy, as a crucial physiological characteristic, joins other distinctive mammalian features that arose during this period of climatic instability 28 ., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Introduction: The worldwide spread of SARS-COV2 had led to a delay in treatment of numerous urological pathologies, even in emergency conditions. We therefore sought to determine whether the timing of diagnosis and treatment and the postoperative outcome of patients with testicular torsion had been changed during the COVID pandemic., Materials and Methods: We considered all patients evaluated in the emergency department (ED) for testicular torsion from February 2018 to August 2019 (pre-COVID period) and from February 2020 to August 2021 (during COVID pandemic). All patients underwent clinical and ultrasound evaluation and subsequently scrotal exploration. Primary outcomes were the time differences from pain onset to ED presentation and from ED presentation to surgical treatment. We also investigated whether the number or orchiectomies required changed during the pandemic., Results: A total of 54 patients were divided in two groups: 40 patients in pre-COVID-19 group and 14 in the COVID-19 cohort. Mean time from symptoms onset to ED access was longer during the pandemic (4.2 ± 5.7 versus 39.6 ± 37.3 hours, p = 0.009). Mean time from ED access to surgery was similar (2.9 ± 1.1 versus 4.2 ± 2.3, p = 0.355). In addition, the number of orchiectomies was higher in COVID-19 group (2.5% versus 28.6%, p < 0.01), compared to a lower number of detorsions (97.5% versus 71.4%, p < 0.01). Elapsed time from pain onset to surgery was directly correlated with the increased white blood cell (WBC) count after surgery (r = 0.399, p = 0.002)., Discussion and Conclusions: The current study identifies a significant delay in presentation of testicular torsion which resulted in a significant increase in orchiectomies with the expected decreased in detorsion/orchiopexy. In addition, there was an increase in the WBC at presentation associated with delayed presentation.

Titanosaurs were successful herbivorous dinosaurs widely distributed in all continents during the Cretaceous, with the major diversity in South America. The success of titanosaurs was probably due to several physiological and ecological factors, in addition to a series of morphological traits they achieved during their evolutionary history. However, the generalist nesting behaviour using different palaeoenvironments and strategies was key to accomplish that success. Titanosaur nesting sites have been found extensively around the world, with notable records in Spain, France, Romania, India, and, especially, Argentina. Here, we describe the first titanosaur nesting site from the Late Cretaceous of Brazil that represents the most boreal nesting site for South America. Several egg-clutches, partially preserved, isolated eggs and many eggshell fragments were discovered in an Inceptisol palaeosol profile of the mining Lafarge Quarry, at the Ponte Alta District (Uberaba Municipality, Minas Gerais State), corresponding to the Serra da Galga Formation (Bauru Group, Bauru Basin). Although classical mechanical preparation and CT scans have not revealed embryonic remains in ovo, the eggs and eggshell features match those eggs containing titanosaurian embryos found worldwide. The morphology of the egg-clutches and observations of the sedimentary characteristics bolster the hypothesis that these sauropods were burrow-nester dinosaurs, as was already suggested for the group based on other nesting sites. The egg-clutches distributed in two levels along the Lafarge outcrops, together with the geopalaeontological data collected, provide clear evidence for the first colonial nesting and breeding area of titanosaur dinosaurs in Brazil., (© 2022. The Author(s).)

Polyphyodonty-multiple tooth generations-in Mesozoic birds has been confirmed since the nineteenth century. Their dental cycle had been assessed through sparse data from tooth roots revealed through broken jawbones and disattached teeth. However, detailed descriptions of their tooth cycling are lacking, and the specifics of their replacement patterns remain largely unknown. Here we present unprecedented µCT data from three enantiornithine specimens from the Upper Cretaceous of southeastern Brazil. The high resolution µCT data show an alternating dental replacement pattern in the premaxillae, consistent with the widespread pattern amongst extinct and extant reptiles. The dentary also reveals dental replacement at different stages. These results strongly suggest that an alternating pattern was typical of enantiornithine birds. µCT data show that new teeth start lingually within the alveoli, resorb roots of functional teeth and migrate labially into their pulp cavities at an early stage, similar to modern crocodilians. Our results imply that the control mechanism for tooth cycling is conserved during the transition between non-avian reptiles and birds. These first 3D reconstructions of enantiornithine dental replacement demonstrate that 3D data are essential to understand the evolution and deep homology of archosaurian tooth cycling., (© 2021. The Author(s).)

In the last decades, several discoveries have uncovered the complexity of mammalian evolution during the Mesozoic Era, including important Gondwanan lineages: the australosphenidans, gondwanatherians, and meridiolestidans (Dryolestoidea). Most often, their presence and diversity is documented by isolated teeth and jaws. Here, we describe a new meridiolestidan mammal, Orretherium tzen gen. et sp. nov., from the Late Cretaceous of southern Chile, based on a partial jaw with five cheek teeth in locis and an isolated upper premolar. Phylogenetic analysis places Orretherium as the earliest divergence within Mesungulatidae, before other forms such as the Late Cretaceous Mesungulatum and Coloniatherium, and the early Paleocene Peligrotherium. The in loco tooth sequence (last two premolars and three molars) is the first recovered for a Cretaceous taxon in this family and suggests that reconstructed tooth sequences for other Mesozoic mesungulatids may include more than one species. Tooth eruption and replacement show that molar eruption in mesungulatids is heterochronically delayed with regard to basal dryolestoids, with therian-like simultaneous eruption of the last premolar and last molar. Meridiolestidans seem endemic to Patagonia, but given their diversity and abundance, and the similarity of vertebrate faunas in other regions of Gondwana, they may yet be discovered in other continents.

Recently, the morphology and encephalization of the brain endocast of the Triassic non-mammaliaform probainognathian cynodont Riograndia guaibensis were studied. Here, we analyzed the brain endocast of an additional specimen of this species. The new endocast shows well-defined olfactory bulbs and a median sulcus dividing the hemispheres, traits that were not clearly observed in the first studied specimen. Encephalization quotients were also calculated, revealing similar values to other non-mammaliaform cynodonts and lower than those of the first analyzed specimen. The analyzed cranium is slightly larger than the first studied one and may represent an advanced ontogenetic stage. Hence, these differences may be related to the intraspecific variation of this cynodont or alternatively, to the preservation of each specimen.

Pterosaurs were the first vertebrates to evolve powered flight 1 and comprised one of the main evolutionary radiations in terrestrial ecosystems of the Mesozoic era (approximately 252-66 million years ago), but their origin has remained an unresolved enigma in palaeontology since the nineteenth century 2-4 . These flying reptiles have been hypothesized to be the close relatives of a wide variety of reptilian clades, including dinosaur relatives 2-8 , and there is still a major morphological gap between those forms and the oldest, unambiguous pterosaurs from the Upper Triassic series. Here, using recent discoveries of well-preserved cranial remains, microcomputed tomography scans of fragile skull bones (jaws, skull roofs and braincases) and reliably associated postcrania, we demonstrate that lagerpetids-a group of cursorial, non-volant dinosaur precursors-are the sister group of pterosaurs, sharing numerous synapomorphies across the entire skeleton. This finding substantially shortens the temporal and morphological gap between the oldest pterosaurs and their closest relatives and simultaneously strengthens the evidence that pterosaurs belong to the avian line of archosaurs. Neuroanatomical features related to the enhanced sensory abilities of pterosaurs 9 are already present in lagerpetids, which indicates that these features evolved before flight. Our evidence illuminates the first steps of the assembly of the pterosaur body plan, whose conquest of aerial space represents a remarkable morphofunctional innovation in vertebrate evolution.

South American titanosaurians have been central to the study of the evolution of Cretaceous sauropod dinosaurs. Despite their remarkable diversity, the fragmentary condition of several taxa and the scarcity of records outside Patagonia and southwestern Brazil have hindered the study of continental-scale paleobiogeographic relationships. We describe two new Late Cretaceous titanosaurians from Quebrada de Santo Domingo (La Rioja, Argentina), which help to fill a gap between these main areas of the continent. Our phylogenetic analysis recovers both new species, and several Brazilian taxa, within Rinconsauria. The data suggest that, towards the end of the Cretaceous, this clade spread throughout southern South America. At the same locality, we discovered numerous accumulations of titanosaurian eggs, likely related to the new taxa. With eggs distributed in three levels along three kilometres, the new site is one of the largest ever found and provides further evidence of nesting site philopatry among Titanosauria.

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 early evolution of lepidosaurs is marked by an extremely scarce fossil record during the Triassic. Importantly, most Triassic lepidosaur specimens are represented by disarticulated individuals from high energy accretion deposits in Laurasia, thus greatly hampering our understanding of the initial stages of lepidosaur evolution. Here, we describe the fragmentary remains of an associated skull and mandible of Clevosaurus hadroprodon sp. nov., a new taxon of sphenodontian lepidosaur from the Late Triassic (Carnian; 237-228 Mya) of Brazil. Referral to Sphenodontia is supported by the combined presence of a marginal dentition ankylosed to the apex of the dentary, maxilla, and premaxilla; the presence of 'secondary bone' at the bases of the marginal dentition; and a ventrally directed mental process at the symphysis of the dentary. Our phylogenetic analyses recover Clevosaurus hadroprodon as a clevosaurid, either in a polytomy with the Late Triassic to Early Jurassic Clevosaurus and Brachyrhinodon (under Bayesian inference), or nested among different species of Clevosaurus (under maximum parsimony). Clevosaurus hadroprodon represents the oldest known sphenodontian from Gondwana, and its clevosaurid relationships indicates that these sphenodontians achieved a widespread biogeographic distribution much earlier than previously thought.

Hypsodonty, the occurrence of high-crowned teeth, is widespread among mammals with diets rich in abrasive material, such as plants or soil, because it increases the durability of dentitions against wear. Hypsodont postcanine teeth evolved independently in multiple mammalian lineages and in the closely related mammaliaforms since the Jurassic period. Here, we report the oldest record, to our knowledge, of hypsodont postcanines in the non-mammaliaform stem-mammal, Menadon besairiei, from the early Late Triassic. The postcanines are long and columnar, with open roots. They were not replaced in older individuals and remained functional after the total wear of the crown enamel. Dental histology suggests that, convergently to hypsodont mammals, wear was compensated by the prolonged growth of each postcanine, resulting in dentine hypsodont teeth most similar to extant xenarthran mammals. These findings highlight the constraints imposed by limited tooth replacement and tooth wear in the evolutionary trajectories of herbivorous mammals and stem-mammals.

Brasilodon quadrangularis (Cynodontia, Probainognathia) is an iconic non-mammaliaform cynodont from the Late Triassic of Brazil (Riograndia Assemblage Zone, Candelária Sequence), being considered as the sister taxon of Mammaliaformes. Although its phylogenetic position is very important, several aspects of its postcranial anatomy remain unclear or unstudied. Here, we present a detailed description of the postcranial elements referred to Brasilodon, including previously mentioned specimens and new ones, which add relevant information about its postcranial morphology and provide a new insight into the anatomical transition between advanced non-mammaliaform cynodonts and early mammaliaforms. Functional and ecological implications are also investigated, based on the postcranial morphology and muscular reconstructions. The postcranium of Brasilodon differs from most non-mammaliaform cynodonts and presents similarities with tritylodontids, early mammaliaforms and extant therians, such as a ventrally oriented scapular glenoid facet, a distinct and ossified greater humeral tubercle, lack of ectepicondylar foramen, olecranon process, hemispherical humeral and femoral heads and a prominent intertrochanteric crest. The humeral torsion, the length of the deltopectoral crest, the large bicipital groove and the well-developed lesser tubercle, indicate that the forelimb of Brasilodon was hold in a semi-sprawling position, with well-developed adductor muscles to maintain the body off the ground. The short femoral neck and the strong medial projection of the femoral head indicate the femur was held in a more erect posture than in basal non-mammaliaform cynodonts. The anterodorsally projected iliac blade with reduced postacetabular process, reduction of the anterior part of the pubis, medially located lesser trochanter indicate a basically mammalian pattern of pelvic musculature, able to swing the femur in a nearly parasagittal plane., Competing Interests: The authors have declared that no competing interests exist.

Uberabatitan ribeiroi is a Late Cretaceous titanosaur (Dinosauria, Sauropoda) from southeastern Brazil. Here we provide a detailed revision of all its available specimens, including new elements from the type-locality. One new autopomorphy is added to diagnosis of the taxon: astragalus with a well-developed anteroposterior crest that mediodistally delimits the tibial articulation. Linear regressions were conducted in an attempt to circumscribe specimens within the type-series, revealing that it is composed of several individuals, with inferred total body lengths varying from 7 to 26 meters. Phylogenetic analyses including U. ribeiroi show that the Brazilian taxon corresponds to a non-saltasaurid lithostrotian titanosaur.

Field work conducted by the staff of the Centro de Pesquisas Paleontológicas Llewellyn Ivor Price of the Universidade Federal do Triângulo Mineiro since 2009 at Campina Verde municipality (MG) have resulted in the discovery of a diverse vertebrate fauna from the Adamantina Formation (Bauru Basin). The baurusuchid Campinasuchus dinizi was described in 2011 from Fazenda Três Antas site and after that, preliminary descriptions of a partial crocodyliform egg, abelisaurid teeth, and fish remains have been done. Recently, the fossil sample has been considerably increased including the discovery of several, partially articulated fish remains referred to Lepisosteiformes and an almost complete and articulated skeleton referred to a new species of Caipirasuchus (Notosuchia, Sphagesauridae), which is the main subject of this contribution. At present, this genus was restricted to the Adamantina Formation cropping out in São Paulo state, with the species Caipirasuchus montealtensis , Caipirasuchus paulistanus , and Caipirasuchus stenognathus . The new material represents the holotype of a new species, Caipirasuchus mineirus n. sp., diferenciated from the previously ones due to the following traits: last two maxillary teeth located posterior to anterior edge of infraorbital fenestra, elongated lateroventral maxillo-jugal suture-about ½ the anteroposterior maxillar length-and contact between posterior crest of quadrate and posterior end of squamosal forming an almost 90° flaring roof of the squamosal, among others. C. mineirus was found in the same outcrop than Campinasuchus but stratigraphically the former occurs in the lower portion of the section with no unambiguous data supporting the coexistance of both taxa., Competing Interests: The authors declare that they have no competing interests.

The Prozostrodontia includes a group of Late Triassic-Early Cretaceous eucynodonts plus the clade Mammaliaformes, in which Mammalia is nested. Analysing their growth patterns is thus important for understanding the evolution of mammalian life histories. Obtaining material for osteohistological analysis is difficult due to the rare and delicate nature of most of the prozostrodontian taxa, much of which comprises mostly of crania or sometimes even only teeth. Here we present a rare opportunity to observe the osteohistology of several postcranial elements of the basal prozostrodontid Prozostrodon brasiliensis , the tritheledontid Irajatherium hernandezi , and the brasilodontids Brasilodon quadrangularis and Brasilitherium riograndensis from the Late Triassic of Brazil (Santa Maria Supersequence). Prozostrodon and Irajatherium reveal similar growth patterns of rapid early growth with annual interruptions later in ontogeny. These interruptions are associated with wide zones of slow growing bone tissue. Brasilodon and Brasilitherium exhibit a mixture of woven-fibered bone tissue and slower growing parallel-fibered and lamellar bone. The slower growing bone tissues are present even during early ontogeny. The relatively slower growth in Brasilodon and Brasilitherium may be related to their small body size compared to Prozostrodon and Irajatherium. These brasilodontids also exhibit osteohistological similarities with the Late Triassic/Early Jurassic mammaliaform Morganucodon and the Late Cretaceous multituberculate mammals Kryptobaatar and Nemegtbaatar. This may be due to similar small body sizes, but may also reflect their close phylogenetic affinities as Brasilodon and Brasilitherium are the closest relatives to Mammaliaformes. However, when compared with similar-sized extant placental mammals, they may have grown more slowly to adult size as their osteohistology shows it took more than one year for growth to attenuate. Thus, although they exhibit rapid juvenile growth, the small derived, brasilodontid prozostrodontians still exhibit an extended growth period compared to similar-sized extant mammals., Competing Interests: The authors declare there are no competing interests.

The early evolution of mammals is associated with the linked evolutionary origin of diphyodont tooth replacement, rapid juvenile growth and determinate adult growth. However, specific relationships among these characters during non-mammalian cynodont evolution require further exploration. Here, polarized light microscopy revealed incremental lines, resembling daily laminations of extant mammals, in histological sections of enamel in eight non-mammalian cynodont species. In the more basal non-probainognathian group, enamel extends extremely rapidly from cusp to cervix. By contrast, the enamel of mammaliamorphs is gradually accreted, with slow rates of crown extension, more typical of the majority of non-hypsodont crown mammals. These results are consistent with the reduction in dental replacement rate across the non-mammalian cynodont lineage, with greater rates of crown extension required in most non-probainognathians, and slower crown extension rates permitted in mammaliamorphs, which have reduced patterns of dental replacement in comparison with many non-probainognathians. The evolution of mammal-like growth patterns, with faster juvenile growth and more abruptly terminating adult growth, is linked with this reduction in dental replacement rates and may provide an additional explanation for the observed pattern in enamel growth rates. It is possible that the reduction in enamel extension rates in mammaliamorphs reflects an underlying reduction in skeletal growth rates at the time of postcanine formation, due to a more abruptly terminating pattern of adult growth in these more mammal-like, crownward species., Competing Interests: The authors declare no competing interests.

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.

In this contribution we report the first occurrence of the enigmatic African probainognathian genus Aleodon in the Middle-early Late Triassic of several localities from the state of Rio Grande do Sul in southern Brazil. Aleodon is unusual among early probainognathians in having transversely-expanded postcanine teeth, similar to those of gomphodont cynognathians. This genus was previously known from the Manda Beds of Tanzania and the upper Omingonde Formation of Namibia. The Brazilian record of this genus is based upon multiple specimens representing different ontogenetic stages, including three that were previously referred to the sectorial-toothed probainognathian Chiniquodon theotonicus. We propose a new species of Aleodon (A. cromptoni sp. nov.) based on the specimens from Brazil. Additionally, we tentatively refer one specimen from the upper Omingonde Formation of Namibia to this new taxon, strengthening biostratigraphic correlations between these strata. Inclusion of A. cromptoni in a phylogenetic analysis of eucynodonts recovers it as the sister-taxon of A. brachyrhamphus within the family Chiniquodontidae. The discovery of numerous specimens of Aleodon among the supposedly monospecific Chiniquodon samples of Brazil raises concerns about chiniquodontid alpha taxonomy, particularly given the extremely broad geographic distribution of Chiniquodon. The discovery of Brazilian Aleodon and new records of the traversodontid Luangwa supports the hypothesis that at least two subzones can be recognized in the Dinodontosaurus Assemblage Zone.

We describe two new cynodonts from the early Late Triassic of southern Brazil. One taxon, Bonacynodon schultzi gen. et sp. nov., comes from the lower Carnian Dinodontosaurus AZ, being correlated with the faunal association at the upper half of the lower member of the Chañares Formation (Ischigualasto-Villa Unión Basin, Argentina). Phylogenetically, Bonacynodon is a closer relative to Probainognathus jenseni than to any other probainognathian, bearing conspicuous canines with a denticulate distal margin. The other new taxon is Santacruzgnathus abdalai gen. et sp. nov. from the Carnian Santacruzodon AZ. Although based exclusively on a partial lower jaw, it represents a probainognathian close to Prozostrodon from the Hyperodapedon AZ and to Brasilodon, Brasilitherium and Botucaraitherium from the Riograndia AZ. The two new cynodonts and the phylogenetic hypothesis presented herein indicate the degree to which our knowledge on probainognathian cynodonts is incomplete and also the relevance of the South American fossil record for understanding their evolutionary significance. The taxonomic diversity and abundance of probainognathians from Brazil and Argentina will form the basis of deep and complex studies to address the evolutionary transformations of cynodonts leading to mammals., Competing Interests: The authors have declared that no competing interests exist.

We report here on a new prozostrodontian cynodont, Botucaraitherium belarminoi gen. et sp. nov., from the Late Triassic Riograndia Assemblage Zone (AZ) of the Candelária Sequence (Santa Maria Supersequence), collected in the Botucaraí Hill Site, Candelária Municipality, state of Rio Grande do Sul, Brazil. The new taxon is based on a single specimen (holotype MMACR-PV-003-T) which includes the left lower jaw, without postdentary bones, bearing the root of the last incisor, canine and four postcanines plus one partial crown inside the dentary, not erupted, and two maxillary fragments, one with a broken canine and another with one postcanine. The features of the lower jaw and lower/upper postcanines resemble those of the prozostrodontians Prozostrodon brasiliensis from the older Hyperodapedon AZ and Brasilodon quadrangularis and Brasilitherium riograndensis from the same Riograndia AZ. The inclusion of Botucaraitherium within a broad phylogenetic analysis, positioned it as a more derived taxon than tritylodontids, being the sister-taxon of Brasilodon, Brasilitherium plus Mammaliaformes. Although the new taxon is based on few cranial elements, it represents a additional faunal component of the Triassic Riograndia AZ of southern Brazil, in which small-sized derived non-mammaliaform cynodonts, closely related to the origin of mammaliaforms, were ecologically well succeed and taxonomically diverse.

The record of non-mosasaur squamates (Reptilia, Squamata) is sparse in the Cretaceus fossil record of Brazil and include six putative reports, three from the Aptian-Albian of the Araripe Basin (Tijubina pontei Bonfim-Júnior and Marques, Olindalacerta brasiliensis Evans and Yabumoto, and a lizard indet.) and three from the Upper Cretaceous of the Bauru Group (Pristiguana brasiliensis Estes and Price, Anilioidae gen. et sp. indet., and Squamata gen. et sp. indet.). In this contribution, a new genus and species of lizard, Brasiliguana prudentis gen. et sp. nov., is described based on an isolated left maxilla with teeth. The material was discovered in an outcrop of the Upper Cretaceous Adamantina Formation (Bauru Group) located in the proximity of Presidente Prudente Municipality, São Paulo State, Brazil. The new taxon is considered a basal non-Priscagamidae+Acrodonta iguanian based on the presence of a weakly inclined anterior margin of the maxillary nasal process and maxillary tooth shape and tooth implantation similar to that of iguanians rather than of other lizard groups (e.g. teiids). This finding significantly increases the squamate lizard diversity of South America, which is still poorly understood and sparsely represented in the fossil record.