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.

Prosaurolophus maximus is a crested hadrosaurine known from numerous specimens from the Dinosaur Park Formation of Alberta. Therefore, it is an ideal taxon to reconstruct patterns of growth and variation in hadrosaurids, and improve our understanding of their evolutionary relationships. This study describes the cranial anatomy of P. maximus, quantitatively examines its range of variation, and provides the first ontogenetic series for this taxon. A second species, P. blackfeetensis, was named based on morphological differences in the characteristic nasal crest; however, morphometric results fail to quantitatively differentiate P. blackfeetensis from P. maximus. A species-level phylogenetic analysis of hadrosaurids recovers P. maximus and P. blackfeetensis as sister taxa. Based on both the morphometric and phylogenetic data, this study supports the previous hypothesis that P. blackfeetensis is a junior synonym of P. maximus thereby substantially increasing its temporal range, and suggests a long period of morphological stasis in this taxon.

Prosaurolophus maximus is a crested hadrosaurine known from numerous specimens from the Dinosaur Park Formation of Alberta. Therefore, it is an ideal taxon to reconstruct patterns of growth and variation in hadrosaurids, and improve our understanding of their evolutionary relationships. This study describes the cranial anatomy of P. maximus, quantitatively examines its range of variation, and provides the first ontogenetic series for this taxon. A second species, P. blackfeetensis, was named based on morphological differences in the characteristic nasal crest; however, morphometric results fail to quantitatively differentiate P. blackfeetensis from P. maximus. A species-level phylogenetic analysis of hadrosaurids recovers P. maximus and P. blackfeetensis as sister taxa. Based on both the morphometric and phylogenetic data, this study supports the previous hypothesis that P. blackfeetensis is a junior synonym of P. maximus thereby substantially increasing its temporal range, and suggests a long period of morphological stasis in this taxon.

No description available.

Os depósitos triássicos continentais do sul do Brasil abrigam uma grande diversidade de tetrápodes terrestres, incluindo terápsidos, rincossauros, rincocefálios e arcossauros, como pseudosúquios e dinossauromorfos. Inserida neste contexto, a Formação Santa Maria, de porção superior datada do Carniano superior, tem papel fundamental no entendimento da origem e irradiação inicial dos dinossauromorfos, pois abriga alguns dos mais antigos registros do grupo em todo mundo, incluindo vários fósseis de dinossauros. Atualmente, a fauna de dinossauromorfos desta unidade é representada por Ixalerpeton polesinensis, Teyuwasu barberenai, Staurikosaurus pricei, Saturnalia tupiniquim, Pampadromaeus barberenai, Buriolestes schultzi e Bagualosaurus agudoensis, enquanto para o Noriano da Formação Caturrita são conhecidos Guaibasaurus candelariensis, Unaysaurus tolentinoi e Sacisaurus agudoensis. Visando o melhor entendimento da diversidade de dinossauromorfos oriundos destes depósitos, foram descritos, no contexto dessa tese, diversos novos fósseis do grupo: ULBRA-PVT059, 280, LPRP/USP 0651, MCN PV 10007-8, 10026, 10027 e 10049. Adicionalmente, foi considerado o recente histórico de pesquisas sobre a origens dos dinossauros para examinar o impacto de novas descobertas e das diferentes hipóteses filogenéticas no entendimento dos padrões biogeográficos da irradiação dos dinossauros. ULBRA-PVT059 e 280 representam os holótipos de duas espécies de dinossauromorfos: Ixalerpeton polesinensis e Buriolestes schultzi. I. polesinensis é o primeiro lagerpetídeo descrito para o Brasil e o único no mundo que preserva elementos do crânio e do membro escapular. O material revela que algumas características antes inferidas como sinapomórficas para Dinosauria já estavam presentes em outros dinossauromorfos. B. schultzi é um sauropodomorfo, provável grupo-irmão dos demais representantes do grupo. Além disso, sua anatomia dentária e relações filogenéticas sugerem que os primeiros dinossauros, incluindo os sauropodomorfos, eram adaptados a faunivoria. LPRP/USP 0651 é o holótipo de uma nova espécie de dinossauro, Nhandumirim waldsangae, da Formação Santa Maria. Apesar de incompleto, as partes preservadas mostram que este se tratava de um indivíduo juvenil, mas que difere em vários aspectos dos demais dinossauros do Carniano, em especial daqueles provenientes dos mesmos níveis estratigráficos. As relações filogenéticas de N. waldsangae indicam que o novo táxon se trata de um dinossauro saurísquio não-sauropodomorfo, possivelmente afim aos terópodos. MCN PV 10007-8, 10026, 10027 e 10049 se tratam de materiais de dinossauros provenientes da localidade tipo de Sacisaurus agudoensis. Estes representam um sauropodomorfo morfologicamente mais semelhante a membros mais recentes do grupo do que aqueles do Carniano. Assim, correlações bioestratigráficas sugeridas pela presença destes sauropodomorfos indicam uma idade mais nova para a localidade tipo de S. agudoensis do que a das biozonas carnianas. As análises biogeográficas consistentemente otimizaram a porção sul do Gondwana como a área ancestral de Dinosauria, o mesmo se dando para clados mais inclusivos. Estes resultados mostram que a hipótese em questão é robusta mesmo com maior amostragem taxonômica e geográfica, e independentemente das hipóteses filogenéticas. Desta forma, é demonstrado que não há suporte para a hipótese da Laurásia representar a área ancestral dos dinossauros. / The Triassic deposits of southern Brazil harbor a great diversity of terrestrial tetrapods, including therapsids, rhynchocephalians, rhynchosaurs, and archosaurs like pseudosuchians and dinosauromorphs. In this context, the Carnian Santa Maria Formation is important for the understanding of the origins and early diversifications of Dinosauromorpha, as it bears one of the oldest records for the group worldwide, including some of the oldest dinosaurs. Its dinosauromorph fauna is currently represented by Ixalerpeton polesinensis, Staurikosaurus pricei, Saturnalia tupiniquim, Pampadromaeus barberenai, Buriolestes schultzi, Bagualosaurus agudoensis, and Teyuwasu barberenai. In comparison, the Norian Caturrita Formation have yielded Guaibasaurus candelariensis, Unaysaurus tolentinoi, and Sacisaurus agudoensis. In order to better understand the dinosauromorph diversity from these deposits, several new fossil remains were described as parts of this thesis: ULBRA-PVT 059, 280, LPRP/USP 0651, MCN PV 10007-8, 10026, 10027, and 10049. In addition, the last 20 years of research efforts on the origins of dinosaurs were compiled to investigate the impact of new discoveries and conflicting phylogenetic hypotheses on the biogeographic history of early dinosauromorphs. ULBRA-PVT 059 and 280 represent the holotypes of a lagerpetid dinosauromorph, Ixalerpeton polesinensis, and a sauropodomorph dinosaur, Buriolestes schultzi. I. polesinensis is the first lagerpetid described from Brazil and only worldwide that preserves skull and scapular limb remains, showing that some previously inferred dinosaur synapomorphies were already present in other early diverging dinosauromorphs. B. schultzi is found as the sister-group to all other sauropodomorphs. In addition, its tooth anatomy and phylogenetic position suggest that early dinosaurs, including sauropodomorphs, were adapted to faunivory. LPRP/USP 0651 is the holotype of a new dinosaur, Nhandumirim waldsangae, from the Santa Maria Formation. Although incomplete, the preserved parts show that it was a juvenile individual, but differing in several respects from other Carnian dinosaurs, especially those from the same stratigraphic levels. The phylogenetic relations of N. waldsangae suggest that the new taxon is a nonsauropodomorph saurischian dinosaur, possibly related to theropods. Dinosaur materials from the type-locality of Sacisaurus agudoensis (MCN PV 10007-8, 10026, 10027, and 10049) represent a sauropodomorph, more similar morphologically to later members of the group than to those of Carnian age. Hence, biostratigraphic correlations suggested by these sauropodomorphs indicate an age for the type-site of S. agudoensis younger than that of the Carnian biozones. Biogeographic analyzes consistently optimize southern Gondwana as the ancestral area for Dinosauria, and this is also the case for more inclusive clades. The results show that the South Gondwanan hypothesis for the origin of dinosaurs is robust even with increased taxonomic and geographic sampling, and independent of phylogenetic uncertainties. It is, therefore, demonstrated that there is no support for Laurassia as the ancestral area of dinosaurs.

Tesis entregada a la Universidad de Chile en cumplimiento parcial de los requisitos para optar al grado de Magíster en Ciencias Biológicas. / El origen del vuelo en Aves está marcado tanto por disminuciones significativas en tamaño corporal como por el aumento en tamaño relativo de las extremidades anteriores. No obstante, previo al origen del vuelo, la evolución de ambos caracteres pudo estar asociada mediante de un patrón alométrico evolutivo negativo para las extremidades anteriores de terópodos. Esta relación ha sido descrita para la mayor parte de Theropoda, perdiéndose en la base de Aves (desde Archaeopteryx lithographica), clado donde se describen en su lugar isometrías o alometrías positivas tanto evolutivas como ontogenéticas. No obstante, las publicaciones donde estas alometrías han sido descritas no corrigen apropiadamente la no-independencia estadística presente entre especies emparentadas. Además, a pesar de que se ha sugerido, ningún estudio a la fecha ha evaluado formalmente la posibilidad de la pérdida del patrón de alometría evolutiva negativa para extremidades anteriores en otros taxa pertenecientes a Theropoda. Del mismo modo la posible relación entre el patrón alométrico evolutivo negativo para extremidades anteriores y un patrón equivalente en la ontogenia no ha sido discutida. En el presente estudio se procedió a re-evaluar el patrón alométrico evolutivo negativo para extremidades anteriores en Theropoda, además de posibles desviaciones respecto a este, utilizando datos de longitudes de húmero y fémur correspondientes a 163 especímenes distribuidos en 108 géneros, incorporando variabilidad intraespecífica, incerteza filogenética y dos modelos evolutivos alternativos a las correcciones filogenéticas utilizadas en los análisis realizados. Mediante estos análisis se recuperaron valores de alometría negativa para el conjunto de los terópodos no-avianos encontrándose además que Coelophysoidea, Ornithomimosauria y Oviraptorosauria presentan diferencias significativas respecto al patrón alométrico general bajo ambos modelos evolutivos, exhibiendo isometría. Se encontró alometría evolutiva negativa para extremidades anteriores en Dromaeosauridae, corroborando que, probablemente, este patrón alométrico evolutivo estaba presente en formas cercanas a Aves. Al revisar la evidencia publicada respecto a alometrías negativas ontogenéticas para extremidades anteriores en Theropoda se encontraron patrones similares a los patrones alométricos evolutivos obtenidos, del mismo modo esta correspondencia se encontró en Aves y otros clados cercanos a Theropoda permitiendo proponer que, además de una órbita y caja craneana expandidas, la paedomorfosis observada en el origen de Aves incluyó extremidades anteriores proporcionalmente mayores como otro rasgo juvenil. / The origin of flight in Aves is marked both by significant decreases in body size and by the increase in relative forelimb size. However, before the origin of flight, the evolution of both characteristics may have been coupled through a pattern of negative forelimb evolutive allometry in Theropoda. This pattern was lost at the origin of Aves (since Archaeopteryx lithographica), clade where isometry or positive allometry has been described instead for both: the phylogeny and ontogeny. However, the publications where these allometries have been described do not properly correct for the statistical non-independence present among related species. Furthermore, even though it has been suggested, no study has formally tested if other taxa in Theropoda show loss of the negative allometric pattern as well. Similarly, the possible relation between the negative forelimb evolutive allometric pattern and an equivalent ontogenetic one has not been discussed. In the present study we re-evaluated the negative forelimb allometric pattern found in the evolution of Theropoda and tested for significate deviations from it using femur and humerus lengths from 163 specimens representing 108 genera, we also incorporated intraspecific variation, phylogenetic uncertainty and two alternative evolutionary models to the phylogenetic corrections used in the analyses. We recovered an evolutionary negative forelimb allometry for all non-avian theropods; we also identified that Coelophysoidea, Ornithomimosauria and Oviraptorosauria show significative differences from the general allometric pattern presenting isometry. We found evolutionary negative forelimb evolutive allometry in Dromaeosauridae showing that this pattern was, most likely, still present in forms close to Aves. Upon reviewing published evidence for ontogenetic negative forelimb allometry in Theropoda we found similar patterns to the ones obtained through our analysis, we also observed similarities between ontogenetic and evolutive forelimb allometric patterns described for Aves and other taxa close to Theropoda suggesting that, in addition to an expanded orbit and braincase, the paedomorphosis observed at the origin of birds probably included proportionally larger arms as another juvenile trait. / Proyecto Anillo ACT172099 y Fondecyt 1150906. / Julio 2019

No description available.

L’évolution des dinosaures sauropodomorphes et la radiation des grands sauropodes sont encore loin d'être comprises, ni même datées avec précision. Si les premiers sauropodes semblent apparaître à la fin du Trias, ils ne se diversifient qu'à partir du Jurassique moyen. L’incertitude entourant la définition à donner au clade Sauropoda est une des principales causes de l'absence de consensus quant à la transition prosauropodes – sauropodes. C’est ce problème du « stem-group » des sauropodes qui est abordé dans cette thèse. Certains prosauropodes sont ainsi aujourd’hui considérés comme des sauropodes à part entière par certains auteurs. Ces taxons basaux diffèrent cependant beaucoup selon les auteurs. La multiplication des analyses cladistiques n’a rien résolu puisque celles-ci n’ont jamais été comparées entre elles.La découverte en 2008, par une équipe du MNHN, dans le Jurassique inférieur du Lesotho, d'un squelette de prosauropode articulé sert ici de base à une étude anatomique et systématique des sauropodomorphes basaux. De nombreux autres restes de dinosaures inédits du Lesotho appartenant aux collections du MNHN viennent compléter le matériel d’étude. Parmi ceux-ci, du matériel rapporté au prosauropode de Maphutseng et des fossiles attribués au genre Meroktenos sont entièrement décrits en détail pour la première fois.Une comparaison critique et détaillée des analyses phylogénétiques antérieures a été réalisée en utilisant la méthode de cladistique comparée telle qu’elle a été formalisée par Sereno en 2009. Elle a permis de démontrer l’impact très important du choix des caractères et de leur codage sur la topologie des arbres. Suite à cela, un premier retour sur une liste compilée de 449 caractères a été effectué. Il a nécessité la vérification d’une matrice de 15 000 à 20 000 cellules, et plus de 1300 mesures. Ce retour aux caractères ont entraîné la suppression de plus d’une centaine de caractères, et la création d’une matrice inédite de 49 unités taxonomiques et 308 caractères. Les résultats de l’analyse phylogénétique de cette matrice, bien que préliminaires, soulèvent plusieurs points intéressants. Plusieurs clades, généralement retrouvés dans les analyses les plus récentes publiées par d’autres auteurs, n’apparaissent pas. En revanche, des clades qui n’avaient jusque-là jamais été retrouvés font leur apparition. Les différentes analyses pointent également du doigt le problème de l’origine des Sauropodomorpha, et relancent le débat sur l’origine du clade Sauropoda. / The evolution of sauropodomorph dinosaurs and the radiation of the large sauropods are still far from being understood and well-dated. If the first sauropods seem to appear at the end of the Triassic, they don’t start to diversify before the Middle Jurassic. The uncertainty surrounding the definition of the clade Sauropoda is one of the main causes of the lack of consensus regarding the prosauropod – sauropod transition. It is this issue of the “stem-group” of sauropods that is dealt with in this thesis. Some prosauropods are now considered to be sauropods by some authors. These basal taxa differ a lot depending on the authors. The multiplication of the cladistic analyses didn’t solve the problem as they never were compared.The discovery in 2008, by a MNHN team, in the Early Jurassic of Lesotho, of a prosauropod articulated skeleton is used herein as the basis for anatomical and systematic study of basal sauropodomorphs. Many other new dinosaur remains from Lesotho, housed in the MNHN collections, complement the study material. Among these, material referred to the Maphutseng prosauropod and fossils referred to the genus Meroktenos are described in details for the first time.A critical and detailed comparison of previous phylogenetic analyses is performed, using the comparative cladistics method as formalized by Sereno in 2009. This work allowed to highlight the significant impact of the choice of characters and their scoring on the topology of the trees. After that, a first reappraisal of a list of 449 compiled characters was carried out. It required the reexamination of a matrix consisting in 15 000 to 20 000 cells, and more than 1300 measures. This work led to remove more than one hundred morphological characters, and the building of a new matrix including 49 taxonomic units and 308 characters. The results of the phylogenetic analysis, although preliminary, raise some interesting points. Several clades, often recovered in the most recent analyses published by other authors, are not recognized here. However, clades that had, until then, never been found, are recovered. The different analyses also point out the hurdle of the origin of Sauropodomorpha, and reopen the debate on the origin of Sauropoda.