Júlio Cesar de Almeida Marsola

‘Thecodont’ refers to teeth implanted in sockets within the jaw, a condition traditionally associated with living mammals and crocodylians, which also coincidentally have teeth attached by ligaments to the socket walls (gomphosis). For over a century, the bony periodontium of many other amniotes has been described as a single tissue, ‘bone of attachment’, causing confusion over dental tissue homology. The conventional definitions of ‘thecodonty’ exclude species with fused teeth (‘ankylothecodonts’), implying a fundamental difference between mammals, crocodylians and most other vertebrates. However, the stereotypically ‘thecodont’ attachment tissues have been discovered in representatives of all major amniote clades, showing that gomphosis and ankylosis likely stem from heterochronic changes in the timing and extent of cementum and alveolar bone mineralization. This challenges (i) previous hypotheses regarding the evolution of the amniote periodontium, (ii) the ‘bone of attachment’ paradigm, and (iii) the significance of ‘thecodonty’. We suggest a new nomenclatural approach that incorporates recent histological and evolutionary research and divides thecodonty into anatomical categories to clarify their origin and evolution. We propose the terms anisothecodont and isothecodont to denote, respectively, asymmetric and symmetric implantation of teeth in their sockets. Regardless of the geometry of the connection, we propose using ankylosis and gomphosis to denote the mode of tooth attachment.

The study of the connection between the teeth and the jaw is important for understanding the palaeobiology of vertebrates, but inconsistent terminology and incomplete sampling have made it difficult to assess the evolutionary significance of some of the related characters. Among archosauromorphs, tooth attachment in dinosaurs and crocodylians is nearly identical to that of mammals in featuring a ligamentous connection (gomphosis), whereas closely related forms appear to have teeth fused to the jaws (ankylosis), as in most other amniotes. Hence, studying tooth attachment of stem‐archosaurs is pivotal to characterize the main shifts in tooth attachment seen in the lineage. Here, we analyze the tooth attachment of rhynchosaurs — a group of quadrupedal herbivorous archosauromorphs that played a key role as primary consumers in many Triassic communities. Their dentition consists of multiple rows of marginal teeth with posterolingual addition of teeth during growth, but their tooth attachment has not been documented in a modern context. Histological data from three rhynchosaur specimens from the Middle Triassic Manda Beds of Tanzania show that, although ankylosed, rhynchosaur teeth are surrounded by an extensive network of Sharpey’s fibers, layers of cementum, and well‐defined zones of alveolar bone. What has been previously described as “spongy bone of attachment” in fact encompasses the same attachment tissues present in mammals, dinosaurs, and crocodylians, albeit completely mineralized in mature teeth. Analysis of different stages of tooth development shows that ankylosis occurs by the growth of alveolar bone towards the cellular cementum, which eventually mineralizes the soft ligament. This suggests that the tissues conflated as “bone of attachment”—alveolar bone, periodontal ligament, and cellular cementum—are homologous across Archosauromorpha. Our data add to a growing body of evidence that heterochronic changes to the timing and extents of mineralization, not convergent evolution to mammal‐like attachment tissues, led to the independent evolution of gomphosis across many amniote lineages, including archosauromorphs.

Most crocodyliforms of the Bauru Group were found in rocks of the Adamantina Formation, whereas the younger Marília Formation is almost devoid of such fossils. Here, we provide a detailed comparative description of MPMA 02–0005/87, a large skull roof found in Marília Formation deposits of the Monte Alto area, assigning it to a new crocodyliform. Despite its fragmentary nature and puzzling suit of characters, the new taxon possesses enough characters to reject its placement within Notosuchia, which is so far the only crocodyliform clade known from the Bauru Group. We tested its phylogenetic position with twodata matrices, both of which recovered the new taxon within Neosuchia and Eusuchia. Although the material does not preserve any eusuchian synapomorphy, a neosuchian affinity is supported by: anterior extension of the meatal chamber covered by the squamosal; constricted flange of the posterior process of the postorbital; poorly developed posterolateral process of squamosal. Finally, the large size estimated for the specimen, ranging from 2.98 to 5.88 metres, coupled with its possible neosuchian affinity, suggests a possible semiaquatic behaviour, an ecology rarely explored by the predominantly terrestrial crocodyliforms of the Bauru Group. http://www.zoobank.org/urn:lsid:zoobank.org:pub:3C7C9499-FFE3-4D5A-A90B-DE2E3572EF16 http://www.zoobank.org/urn:lsid:zoobank.org:pub:98FFC142-674F-464D-BCFD-721C608A2635

Dinosaurs possess a form of tooth attachment wherein an unmineralized periodontal ligament suspends each tooth within a socket, similar to the condition in mammals and crocodylians. However, little information is known about tooth attachment and implantation in their close relatives, the silesaurids. We conducted a histological survey of several silesaurid taxa to determine the nature of tooth attachment in this phylogenetically and paleoecologically important group of archosaurs. Our histological data demonstrate that these early dinosauriforms do not exhibit the crocodilian/dinosaur condition of a permanent gomphosis, nor the rapid ankylosis that is plesiomorphic for amniotes. Instead, all sampled silesaurids exhibit delayed ankylosis, a condition in which teeth pass through a prolonged stage where the teeth are suspended in sockets by a periodontal ligament, followed by eventual mineralization and fusion of the tooth to the jaws. This suggests that tooth attachment in crocodylians and dinosaurs represent the further retention of an early ontogenetic stage compared to silesaurids, a paedomorphic trend that is mirrored in the evolution of synapsid tooth attachment. It also suggests that the dinosaur and crocodylian gomphosis was convergently acquired via heterochrony or, less likely, that the silesaurid condition represents a reversal to a plesiomorphic state. Moreover, if Silesauridae is nested within Ornithischia, a permanent gomphosis could be convergent between the two main dinosaur lineages, Ornithischia and Saurischia. These results demonstrate that dental characters in early archosaur phylogenies must be chosen and defined carefully, taking into account the relative duration of the different phases of dental ontogeny.

Noasaurines form an enigmatic group of small-bodied predatory theropod dinosaurs known from the Late Cretaceous of Gondwana. They are relatively rare, with notable records in Argentina and Madagascar, and possible remains reported for Brazil, India, and continental Africa. In south-central Brazil, the deposits of the Bauru Basin have yielded a rich tetrapod fauna, which is concentrated in the Bauru Group. The mainly aeolian deposits of the Caiuá Group, on the contrary, bear a scarce fossil record composed only of lizards, turtles, and pterosaurs. Here, we describe the first dinosaur of the Caiuá Group, which also represents the best-preserved theropod of the entire Bauru Basin known to date. The recovered skeletal parts (vertebrae, girdles, limbs, and scarce cranial elements) show that the new taxon was just over 1 m long, with a unique anatomy among theropods. The shafts of its metatarsals II and IV are very lateromedially compressed, as are the blade-like ungual phalanges of the respective digits. This implies that the new taxon could have been functionally monodactyl, with a main central weight-bearing digit, flanked by neighbouring elements positioned very close to digit III or even held free of the ground. Such anatomical adaptation is formerly unrecorded among archosaurs, but has been previously inferred from footprints of the same stratigraphic unit that yielded the new dinosaur. A phylogenetic analysis nests the new taxon within the Noasaurinae clade, which is unresolved because of the multiple alternative positions that Noasaurus leali can acquire in the optimal trees. The exclusion of the latter form results in positioning the new dinosaur as the sister-taxon of the Argentinean Velocisaurus unicus.

Sauropodomorphs are the most abundant and diverse clade of Triassic dinosaurs, but the taxonomy of their earliest (Carnian) representatives is still poorly understood. One such taxon is Pampadromaeus barberenai, represented by a nearly complete disarticulated skeleton recovered from the upper part of the Santa Maria Formation of Rio Grande do Sul, Brazil. Here, the osteology of Pam. barberenai is fully described for the first time. Detailed comparisons with other Carnian sauropodomorphs reveal a unique anatomy, corroborating its status as a valid species. Potential autapomorphies of Pam. barberenai can be seen in the articulation of the sacral zygapophyses, the length of the pectoral epipodium, the shape of the distal articulation of the femur and the proximal articulation of metatarsal 1. A novel phylogenetic study shows that relationships among the Carnian sauropodomorphs are poorly constrained, possibly because they belong to a “zone of variability”, where homoplasy abounds. Yet, there is some evidence that Pam. barberenai may nest within Saturnaliidae, along with Saturnalia tupiniquim and Chromogisaurus novasi, which represents the sister group to the larger sauropodomorphs, i.e. Bagualosauria.

Dinosaurs were ubiquitous in terrestrial ecosystems through most of the Mesozoic and are still diversely represented in the modern fauna in the form of birds. Recent efforts to better understand the origins of the group have resulted in the discovery of many new species of early dinosaur and their closest relatives (dinosauromorphs). In addition, recent re‐examinations of early dinosaur phylogeny have highlighted uncertainties regarding the interrelationships of the main dinosaur lineages (Sauropodomorpha, Theropoda and Ornithischia), and questioned the traditional hypothesis that the group originated in South Gondwana and gradually dispersed over Pangaea. Here, we use an historical approach to examine the impact of new fossil discoveries and changing phylogenetic hypotheses on biogeographical scenarios for dinosaur origins over 20 years of research time, and analyse the results in the light of different fossil record sampling regimes. Our results consistently optimize South Gondwana as the ancestral area for Dinosauria, as well as for more inclusive clades including Dinosauromorpha, and show that this hypothesis is robust to increased taxonomic and geographic sampling and divergent phylogenetic results. Our results do not find any support for the recently proposed Laurasian origin of dinosaurs and suggest that a southern Gondwanan origin is by far the most plausible given our current knowledge of the diversity of early dinosaurs and non‐dinosaurian dinosauromorphs.