Ornithischian dinosaurs form a diverse, globally-distributed clade including the dominant large land herbivores in many Mesozoic faunas. However, because we lack a well-resolved species-level phylogeny including basal and derived members, our understanding of the initial ornithischian radiation in the Late Triassic is unclear, which further complicates our ability to test biogeographic scenarios. A new species-level phylogenetic analysis, based on comprehensive empirical assessments of a wide sample of ornithischians with an emphasis on basal taxa, helps resolve the relationships of historically labile taxa. Heterodontosauridae is a monophyletic group outside of Genasauria, corroborating recent studies, though the current analysis did not recover distinct Laurasian and Gondwanan clades as has been previously reported. Lesothosaurus, traditionally considered as one of the most basal ornithischians, is here recovered as the basalmost neornithischian more closely related to cerapodans than to thyreophorans. The resultant phylogeny shows good resolution among basal taxa; however, most members of Cerapoda collapse into a polytomy recovering only a monophyletic Iguanodontia, Marginocephalia, and a few other derived clades. The phylogenetic placement of these taxa is critical for assessments of character evolution for more derived clades and ancestral state reconstructions for Ornithischia in general, as well as determining areas of origination for clades across the tree. With a detailed species-level phylogeny, I was able to estimate the hypothetical ancestral body plan for all Ornithischia by reconstructing the character states in the present analysis. The results of this analysis suggest that the hypothetical ancestral ornithischian had similar skull morphology to Lesothosaurus, but that the postcranial skeleton was more similar to other basal ornithischians such as Heterodontosaurus and to basal saurischians such as Herrerasaurus and Eoraptor. Most analyses of derived clades of ornithischians rely on Lesothosaurus as a representative outgroup taxon; however, the postcranial skeleton of Lesothosaurus represents a fairly specialized, relatively abbreviated forelimb unlike what the ancestral ornithischian possessed. Building on the species-level phylogeny focused on basal taxa, I performed a detailed event-based quantitative biogeographic analysis based on a comprehensive composite phylogeny including more derived taxa from previously published studies. A few patterns can be explained by vicariance but numerous dispersal events are required to explain overall ornithischian biogeography, particularly in the Late Jurassic and Early and Late Cretaceous. Although the early record of ornithischians is sparse, it is clear that the clade originated from small, bipedal cursors in southern Gondwana and spread into Laurasia by the Early Jurassic and quickly diversified into the iconic groups that characterize Ornithischia. A well-resolved phylogenetic and biogeographic hypothesis of ornithischian diversity, particularly with basal taxa such as the heterodontosaurids and Lesothosaurus, provides a framework on which to test hypotheses of coevolution as well as the evolution of herbivory, ontogeny, physiology, and sexual dimorphism in the fossil record.

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Parmi les dinosaures, le clade des Ornithopodes est l’un de ceux qui a rencontré le plus de succès. Apparus au Jurassique supérieur, le groupe s’est dispersé et diversifié jusqu’à sa disparition à la fin du Crétacé. Je me suis attachée à étudier le cerveau des Ornithopodes d’Europe et d’Asie, à établir des comparaisons avec les autres archosaures et de nouveaux arbres phylogénétiques incluant des caractères issus de l’endocrâne. Pour ce faire, des moulages et des reconstitutions à partir de données CT-scan ont été réalisées pour étudier la cavité endocrânienne de divers membres de ce groupe. J’ai réalisé des moulages endocrâniens en silicone de 3 taxons et les reconstructions à partir de données CT-scan de 3 autres taxons de Dinosaures Ornithopodes. Une collection unique d’endocrânes de crocodiles et d’oiseaux récents étend les possibilités de comparaison. Certains endocrânes des taxons fossiles ont confirmé les caractéristiques décrites précédemment dans la littérature, tandis que de nouveaux éléments sont apparus. Les endocrânes n’ont en effet pas seulement livré la morphologie du cerveau, mais aussi des valleculae, le détail des nerfs crâniens et de la glande pituitaire. J’ai ainsi pu étendre la présence de valleculae, qui est un indice fort en faveur d’un télencéphale développé, chez un nouvel Hadrosauroidea, alors que cela n’était connu que chez les Hadrosauridae et les membres dérivés de deux autres groupes (Theropoda et Pachycephalosauria). Le cerveau des Ornithopodes dérivés était caractérisé par des hémisphères cérébraux très larges et de forme arrondie. Les flexions crâniale et pontine sont inexistantes, à l’opposé de ce qui est observé chez les Saurischiens. Les pédoncules olfactifs étaient larges. J’ai également fourni de nouveaux exemples de l’influence de la taille de la glande pituitaire sur la taille totale de l’individu. Le cerveau des Ornithopodes a subi des changements au cours de leur évolution :le plus marquant est l’augmentation du volume des hémisphères cérébraux par rapport au reste du cerveau. J’émets l’hypothèse que cette augmentation résulte de la complexification des comportements chez les Ornithopodes.<p>Une nouvelle phylogénie a été établie, bénéficiant de l’apport de nouveaux caractères basés sur l’endocrâne. Elle apporte des éléments de réflexion intéressants quant à la position de plusieurs taxons d’Iguanodontia basaux. La résolution est cependant faible et d’autres études devront être menées dans le futur. Les relations de parenté ne sont pas stables et de faibles changements entraînent des différences notables dans les résultats des analyses phylogénétiques./Among the Dinosauria, Ornithopoda were one of the most successful clade. Since the Late Jurassic, they spread and diversified until the end of the Cretaceous. I studied the brain of Ornithopoda from Europe and Asia, established comparisons with other archosaurs and new phylogenetic analyses including endocranial characters. In order to do this, I made silicone endocasts of 3 taxa and virtual reconstructions from CT-scan dataset of 3 other taxa of ornithopod dinosaurs. A collection of extant crocodiles and birds allows more points of comparison. Some endocasts made on the fossil specimens confirmed previously described characteristics, while new ones were brought to our attention. The endocasts opened up not only the morphology of the brain, but also the anatomy of the cranial nerves, the pituitary gland and the presence of valleculae. This last element, evidence of a developped telencephalon, has been established in a new Hadrosauroidea species as his oldest occurrence. The brain of more derived Ornithopods was characterized by very large cerebral hemispheres. The pontine and cranial flexures disappeared, to the contrary to what is observed in Saurischians. The olfactory peduncles were large. New evidences about the correlation between the size of the pituitary gland and the size of the individual. The Ornithopod’s brain changed throughout their evolution :the most striking is the increase of the cerebral hemispheres. The complexity of behaviors exhibited by Ornithopods is suggested as the trigger of the increase of the size of the cerebral hemispheres.<p>A new phylogenetic analysis was established, including new characters from the endocranial cavity. It brings interesting perspectives about the position of several basal Iguanodontia. Unfortunately the resolution is weak and new studies will be needed. The relationships are not stable and small changes lead to instabilities in the result of the phylogenetic analysis. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

For 130 years dinosaurs have been divided into two distinct clades – Ornithischia and Saurischia. This dissertation looks at the earliest evolution of the clade Dinosauria by focusing upon the interrelationships of the major subsidiary clades within it. It does this following examination, comparison and description of early dinosaur material, and by utilising modern phylogenetic analysis techniques, to rigorously and objectively test the fundamental groupings within the clade Dinosauria using a newly compiled dataset of early dinosaurs and other dinosauromorphs (= close dinosaur relatives). The current consensus on how the principal clades within Dinosauria (Theropoda, Sauropodomorpha and Ornithischia) are related to one another is challenged by the results of these analyses. This study finds, for the first time, a sister-group relationship between Ornithischia and Theropoda, here termed Ornithoscelida. Consequently, a new definition for Dinosauria is presented, as the historic definition would exclude all members of Sauropodomorpha from the clade. As well as this, I propose revisions to the definitions of each of the principal dinosaurian sub-divisions and propose a new timeframe and geographic setting for the origin of Dinosauria. These new hypotheses force re-evaluations of early dinosaur cladogenesis and character evolution, suggest the independent acquisition of hypercarnivory in multiple dinosaur groups and offers an explanation for many of the anatomical features previously regarded as striking convergences between theropods and early ornithischians. As well as presenting new anatomical data on many early dinosaurs and dinosauromorphs, including a comprehensive re-description of the postcranial anatomy of Lesothosaurus diagnosticus (Chapter 2), and a new anatomical dataset of early dinosaurs (the largest ever compiled), this thesis goes on to implement the new dataset to investigate a number of important outstanding questions about early dinosaur evolution and provides new lines of enquiry for future workers to pursue. The results of this thesis reveal the oldest known members of the dinosaurian clades Theropoda and Sauropodomorpha (Chapter 6), as well as a new clade within Ornithischia; a taxon previously thought to represent a derived theropod has been recovered as a potential ‘missing link’ between theropods and ornithischians using the new dataset. This work now provides a unique tool for the assessment of the phylogenetic affinities of early dinosaurs and dinosauromorphs and, once published, will hopefully become the benchmark dataset for palaeontologists working in this area.