Diphyodont tooth replacement of Brasilodon-A Late Triassic eucynodont that challenges the time of origin of mammals.

Two sets of teeth (diphyodonty) characterise extant mammals but not reptiles, as they generate many replacement sets (polyphyodonty). The transition in long-extinct species from many sets to only two has to date only been reported in Jurassic eucynodonts. Specimens of the Late Triassic brasilodontid eucynodont Brasilodon have provided anatomical and histological data from three lower jaws of different growth stages. These reveal ordered and timed replacement of deciduous by adult teeth. Therefore, this diphyodont dentition, as contemporary of the oldest known dinosaurs, shows that Brasilodon falls within a range of wide variations of typically mammalian, diphyodont dental patterns. Importantly, these three lower jaws represent distinct ontogenetic stages that reveal classic features for timed control of replacement, by the generation of only one replacement set of teeth. This data shows that the primary premolars reveal a temporal replacement pattern, importantly from directly below each tooth, by controlled regulation of tooth resorption and regeneration. The complexity of the adult prismatic enamel structure with a conspicuous intra-structural Schmelzmuster array suggests that, as in the case of extant mammals, this extinct species would have probably sustained higher metabolic rates than reptiles. Furthermore, in modern mammals, diphyodonty and prismatic enamel are inextricably linked, anatomically and physiologically, to a set of other traits including placentation, endothermy, fur, lactation and even parental care. Our analysis of the osteodental anatomy of Brasilodon pushes back the origin of diphyodonty and consequently, its related biological traits to the Norian (225.42 ± 0.37 myr), and around 25 myr after the End-Permian mass extinction event.

Cordão Formation: loess deposits in the southern coastal plain of the state of Rio Grande do Sul, Brazil.

Loess consists of silt-dominated sediments that cover ~10% of the Earth's surface. In southern South America it occurs in Argentina, Bolivia, Paraguay and Uruguay, and its presence in southern Brazil was never studied in detail. Here is proposed a new lithostratigraphic unit, Cordão Formation, consisting of loess deposits in the southern Brazilian coastal plain. It consists of fine-very fine silt with subordinate sand and clay, found mostly in lowland areas between Pleistocene coastal barriers. These sediments are pale-colored (10YR hue) and forms ~1,5-2,0 meter-thick stable vertical walls. The clay minerals include illite, smectite, interstratified illite/smectite and kaolinite, the coarser fraction is mostly quartz and plagioclase. Caliche and iron-manganese nodules are also present. The only fossils found so far are rodent teeth and a tooth of a camelid (Hemiauchenia paradoxa). Luminescence ages indicate that this loess was deposited in the latest Pleistocene, between ~30 and 10 kyrs ago, and its upper portion was modified by erosion and accumulation of clay and organic matter in the Holocene. The estimated accumulation rate was ~630 g/m2/year. The probable source of this loess is the Pampean Aeolian System of Argentina and it would have been deposited by the increased aeolian processes of the last glacial.

New rhadinosuchine proterochampsids from the late Middle-early Late Triassic of southern Brazil enhance the diversity of archosauriforms.

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.

New information on Riograndia guaibensis Bonaparte, Ferigolo & Ribeiro, 2001 (Eucynodontia, Tritheledontidae) from the Late Triassic of southern Brazil: anatomical and biostratigraphic implications.

The tritheledontid Riograndia guaibensis was the first cynodont described for the "Caturrita Formation" fauna from the Late Triassic of southern Brazil (Santa Maria 2 Sequence). The type materials did not preserve anatomical information regarding braincase, occiput, basicranium, zygomatic arch, postdentary bones and craniomandibular joint. Here new materials are described and supply the missing information. Riograndia shows a suite of important anatomical features quite derived among the non-mammaliaform eucynodonts, such as the partial closure of the medial orbital wall and braincase, extensive secondary osseous palate, wide primary palate, basicranium with jugular foramen separated from the periphery of fenestra rotunda, narrow zygomatic arch and much reduced postdentary bones. Many of these features constitute synapomorphies shared only with the other members of mammaliamorpha. Thus, the almost complete cranial, mandibular and dental information from the new fossils of Riograndia can bring a significant improve in the understanding of the anatomy and phylogenetic relationships of the tritheledontids and help to elucidate the transformational steps involved in the cynodont-mammal transition. Additionally, Riograndia is a key taxon in refining the "Caturrita Formation" biostratigraphy, enabling the connection of several fossiliferous outcrops that have a rich tetrapod fauna that can be correlated with other Triassic faunas from Gondwana and Laurasia.

A New Clevosaurid from the Triassic (Carnian) of Brazil and the Rise of Sphenodontians in Gondwana.

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.

Spinosaur taxonomy and evolution of craniodental features: Evidence from Brazil.

Fossil sites from Brazil have yielded specimens of spinosaurid theropods, among which the most informative include the cranial remains of Irritator, Angaturama, and Oxalaia. In this work some of their craniodental features are reinterpreted, providing new data for taxonomic and evolutionary issues concerning this particular clade of dinosaurs. The mesial-most tooth of the left maxilla of the holotype of Irritator is regarded as representing the third tooth position, which is also preserved in the holotype of Angaturama. Thus, both specimens cannot belong to the same individual, contrary to previous assumptions, although they could have been the same taxon. In addition, the position of the external nares of Irritator is more comparable to those of Baryonyx and Suchomimus instead of other spinosaurine spinosaurids. In fact, with regards to some craniodental features, Brazilian taxa represent intermediate conditions between Baryonychinae and Spinosaurinae. Such a scenario is corroborated by our cladistic results, which also leave open the possibility of the former subfamily being non-monophyletic. Furthermore, the differences between spinosaurids regarding the position and size of the external nares might be related to distinct feeding habits and degrees of reliance on olfaction. Other issues concerning the evolution and taxonomy of Spinosauridae require descriptions of additional material for their clarification.

The African cynodont Aleodon (Cynodontia, Probainognathia) in the Triassic of southern Brazil and its biostratigraphic significance.

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.

An exceptional fossil skull from South America and the origins of the archosauriform radiation.

Birds, dinosaurs, crocodilians, pterosaurs and their close relatives form the highly diverse clade Archosauriformes. Archosauriforms have a deep evolutionary history, originating in the late Permian, prior to the end-Permian mass extinction, and radiating in the Triassic to dominate Mesozoic ecosystems. However, the origins of this clade and its extraordinarily successful body plan remain obscure. Here, we describe an exceptionally preserved fossil skull from the Lower Triassic of Brazil, representing a new species, Teyujagua paradoxa, transitional in morphology between archosauriforms and more primitive reptiles. This skull reveals for the first time the mosaic assembly of key features of the archosauriform skull, including the antorbital and mandibular fenestrae, serrated teeth, and closed lower temporal bar. Phylogenetic analysis recovers Teyujagua as the sister taxon to Archosauriformes, and is congruent with a two-phase model of early archosauriform evolution, in response to two mass extinctions occurring at the end of the Guadalupian and the Permian.

An unusual pterosaur specimen (Pterodactyloidea, ?azhdarchoidea) from the early cretaceous Romualdo formation of Brazil, and the evolution of the pterodactyloid palate.

A new and unusual specimen of a probable azhdarchoid pterosaur is described for the Early Cretaceous (Albian) Romualdo Formation of Brazil. The specimen consists of a palate that, although fragmentary, has a unique morphology differing from all other known pterosaurs with preservation of palatal elements. The new specimen probably indicates the presence of a yet undescribed pterodactyloid taxon for Romualdo Formation and brings new information on pterosaur diversity of this sedimentary unity. Mainly due to the rarity of pterodactyloid specimens with palate preservation, this structure has been overlooked in this clade. Here, we reassess the palatal anatomy of Pterodactyloidea, revealing an intriguing variety of morphotypes and evolutionary trends, some of them described here for the first time. The morphological disparity displayed by different pterodactyloid taxa may be further evidence of the presence of diverse feeding strategies within the clade.

New information on Riograndia guaibensis Bonaparte, Ferigolo & Ribeiro, 2001 (Eucynodontia, Tritheledontidae) from the Late Triassic of southern Brazil: anatomical and biostratigraphic implications

The tritheledontid Riograndia guaibensis was the first cynodont described for the "Caturrita Formation" fauna from the Late Triassic of southern Brazil (Santa Maria 2 Sequence). The type materials did not preserve anatomical information regarding braincase, occiput, basicranium, zygomatic arch, postdentary bones and craniomandibular joint. Here new materials are described and supply the missing information. Riograndia shows a suite of important anatomical features quite derived among the non-mammaliaform eucynodonts, such as the partial closure of the medial orbital wall and braincase, extensive secondary osseous palate, wide primary palate, basicranium with jugular foramen separated from the periphery of fenestra rotunda, narrow zygomatic arch and much reduced postdentary bones. Many of these features constitute synapomorphies shared only with the other members of mammaliamorpha. Thus, the almost complete cranial, mandibular and dental information from the new fossils of Riograndia can bring a significant improve in the understanding of the anatomy and phylogenetic relationships of the tritheledontids and help to elucidate the transformational steps involved in the cynodont-mammal transition. Additionally, Riograndia is a key taxon in refining the "Caturrita Formation" biostratigraphy, enabling the connection of several fossiliferous outcrops that have a rich tetrapod fauna that can be correlated with other Triassic faunas from Gondwana and Laurasia.

O triteledontídeo Riograndia guiabensis foi o primeiro cinodonte descrito para a fauna da "Formação Caturrita" do Triássico Superior do sul do Brasil (Sequência Santa Maria 2). Os materiais da série-tipo não forneceram informações anatômicas relativas à caixa craniana, ociput, basicrânio, arco zigomático, ossos pós-dentários e articulação crânio-mandibular. Neste artigo são descritos novos materiais que suprem as informações anatômicas não contempladas anteriormente. Riograndia apresenta um conjunto de importantes aspectos anatômicos derivados dentre os eucinodontes não-mammaliaformes, tais como fechamento parcial da parede orbital medial e da caixa craniana, extenso palato ósseo secundário, palato primário alargado, basicrânio com separação do forame jugular e da fenestra rotunda, arco zigomático estreito e ossos pós-dentários extremamente reduzidos. Muitas destas feições constituem sinapomorfias compartilhadas somente com os demais membros do clado Mammaliamorpha. Assim, as praticamente completas informações crânio-mandibulares e dentárias de Riograndia trazem uma significante contribuição ao conhecimento da anatomia dos triteledontídeos e suas relações filogenéticas e auxiliam na elucidação sobre os passos transformacionais envolvidos na transição cinodonte-mamífero. Adicionalmente, Riograndia figura como um táxon-chave no refinamento bioestratigráfico da "Formação Caturrita", através da integração de vários afloramentos fossilíferos os quais apresentam uma rica fauna de tetrápodes que pode ser correlacionada com outras faunas triássicas do Gondwana e da Laurásia.