A dissertation submitted to the Faculty of Science, University of Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Master of Science. / Dinocephalians form an important part of the Upper Permian therapsid faunas of South Africa and Russia, and have also recently been reported from China and Brazil, and even more recently a diverse dinocephalian fauna has been described from Zimbabwe. This thesis reports a new primitive tapinocephalid dinocephalian, NHMB 1556, from the Upper Madumabisa Mudstone Formation in Zimbabwe. NHMB 1556 is considered to be closely related to Avenantia and it possesses primitive tapinocephalid characters, but is more derived than Tapinocaninus, the most primitive tapinocephalid dinocephalian known. NHMB 1556 is distinguished from other tapinocephalines by having a groove on the squamosal below its dorsal contact with the parietal, a low squamosal-parietal suture on the posterior border of the temporal fenestra, a vomer which extends posteroventrally and forms the anterior margin of the interpterygoidal vacuity and basisphenoid, which is semi-circular anteroventrally. / AC2017

A dissertation submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Master of Science. Johannesburg, 2014. / Anteosaurs (Therapsida: Dinocephalia: Anteosauria) were the dominant terrestrial predators during the late Middle Permian period and became extinct at the close of the Tapinocephalus Assemblage Zone. Only two genera of anteosaurs, Australosyodon and Anteosaurus, are recognised from the Karoo rocks of South Africa. A newly discovered small anteosaurid skull from the Abrahamskraal Formation is fully described. Because of its relatively large orbits, the unfused nature of its sutures, and the lack of replacement teeth in the dental alveoli, the specimen is considered to be a juvenile Anteosaurus magnificus. A full computer-aided 3-D reconstruction of the skull enabled cranial measurements to be taken for an allometric analysis which included twenty-three measurements and eleven specimens. Positive allometry was found for four of the measurements suggesting fast growing in the temporal region, and a significant difference in the development of the postorbital bar and suborbital bar between juveniles and adults. Phylogenetic research shows that the Russian anteosaurids to be forerunners to Anteosaurus, and because the juvenile Anteosaurus (BP/1/7074) manifests many features of both Syodon and Titanophoneus, it is suggested that ontogenetic growth of Anteosaurus follows Haeckel’s Law.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第22279号 / 理博第4593号 / 新制||理||1659(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 本川 雅治, 准教授 中野 隆文, 教授 曽田 貞滋 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

京都大学 / 新制・課程博士 / 博士(理学) / 甲第25143号 / 理博第5050号 / 新制||理||1719(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 本川 雅治, 准教授 中野 隆文, 教授 森 哲 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DGAM

A integração morfológica determina a quantidade de variação disponível em diferentes direções do espaço multivariado entre um conjunto de caracteres, e pode assim ter importantes consequências evolutivas, enviesando a direção e a taxa de evolução morfológica em determinadas direções. Assim, estudar a evolução da integração morfológica é importante para entender a evolução fenotípica de uma linhagem. Neste trabalho, utilizo abordagens de genética quantitativa comparativa para estudar a evolução da integração morfológica no crânio em carnívoros da subordem Feliformia, grupo com extensa diversidade filogenética, morfológica e ecológica para o qual análises anteriores de integração analisaram número reduzido de espécies. Para este fim, comparo matrizes de covariância fenotípicas intra-populacionais quantificando a associação entre 35 distâncias lineares no crânio de um conjunto de táxons incluindo todas as sete famílias e mais de um terço da diversidade filogenética atual da subordem. Investigo também a associação entre integração morfológica e a evolução da morfologia média, assim como a influência da história evolutiva (filogenia) e de dois aspectos da ecologia (dieta e socialidade) sobre a evolução da morfologia média e da integração. Os padrões de integração fenotípica no crânio (i.e. quais caracteres variam em conjunto com maior intensidade) são bastante estáveis na subordem, como observado para outros mamíferos, mas revelam alguma sub-estruturação, com padrões um pouco distintos em Felidae, Hyaenidae e Prionodontidae As magnitudes de integração são também relativamente estruturadas, com valores geralmente mais altos em Felidae e consistentemente mais baixos em Hyaenidae. Apesar disso, a magnitude geral de integração é evolutivamente plástica, como em outros mamíferos, havendo importantes variações internas a cada família. Encontrei uma associação geral entre a evolução da morfologia média e modificações em padrões e magnitudes de integração para a subordem como um todo e em vários grupos menos inclusivos, o que em alguns casos se manifesta como uma semelhança da integração entre táxons morfologicamente convergentes, mas filogeneticamente distantes. A evolução de padrões de integração se correlaciona à história filogenética em Feliformia como um todo e na maior parte dos grupos acima, mas não abaixo do nível de família, corroborando padrões de covariação relativamente distintos apesar de sua alta similaridade, bem como integração mais restrita internamente a cada família. A dieta está associada à filogenia, e correlaciona-se à evolução da morfologia média e dos padrões de integração na subordem como um todo antes, mas não após correção levando em conta a proximidade filogenética, indicando que alterações na dieta são parte da história evolutiva compartilhada que levou às diferenças em morfologia média e integração existentes no grupo. Os caracteres cujos padrões de covariação estão mais associados à dieta na subordem incluem partes das regiões oral e zigomática, além de algumas partes do neurocrânio. A socialidade está relacionada à evolução da morfologia craniana média, mas não aos padrões de integração gerais em Feliformia, apesar de existirem associações entre a socialidade e os padrões de resposta a seleção de alguns caracteres específicos, incluindo partes do neurocrânio e distâncias ligadas ao arco zigomático. Os mangustos (família Herpestidae) mostraram influência significativa tanto da dieta quanto da socialidade na evolução de sua morfologia craniana média e de seus padrões de integração, ressaltando a importância desses dois aspectos ecológicos na evolução fenotípica do grupo e mostrando, pela primeira vez, uma associação direta entre a integração no crânio e o comportamento social, possivelmente intermediada por aspectos relacionados ao desenvolvimento do cérebro. O presente trabalho sugere, assim, uma complexa interação de fatores afetando a evolução da morfologia craniana em Feliformia, e indica direções para trabalhos futuros, incluindo análises dos padrões de modularidade craniana no grupo e, possivelmente, análises comparativas em diferentes morfotipos hipercarnívoros abordando a morfologia média e a integração morfológica no crânio, bem como a dieta e possivelmente outros fatores / Morphological integration determines the standing variation available within a lineage in different directions of the multivariate space of a trait set, and can therefore have important evolutionary consequences, biasing the direction and rate of morphological evolution in certain directions. Consequently, studying the evolution of morphological integration is important to understand a lineage\'s phenotypic evolution. Here I employ a comparative quantitative genetic framework to study the evolution of morphological integration in the skull for the suborder Feliformia (Carnivora), a group with remarkable phylogenetic, morphological and ecological diversity for which previous analysis of integration included a limited number of species. To this purpose, I compare within-population phenotypic covariance matrices quantifying the associations between 35 skull linear measurements in a set of taxa including all seven feliform families and over one third of the suborder\'s extant phylogenetic diversity. I also assess the association between morphological integration and the evolution of average skull morphology, as well as the influence of evolutionary history (phylogeny) and two ecological aspects (diet and sociality) on the evolution of average morphology and integration. Skull integration patterns (i.e. which characters covary the most) are quite stable in the suborder, as seen among other mammals, but also have some structuring between groups, with relatively distinct patterns in Felidae, Hyaenidae and Prionodontidae. Integration magnitudes are also relatively structured, with generally higher values in Felidae and consistently lower values in Hyaenidae. Nevertheless, the overall magnitude of integration is evolutionarily labile, as in other mammals, with important variations occurring within each family. I found a general association between the evolution of average morphology and changes in integration patterns and magnitudes for the suborder as a whole and also for many less inclusive groups, which in some cases leads to similar integration between morphologically convergent, but phylogenetically distant taxa. The evolution of integration patterns is correlated to phylogenetic history in Feliformia as a whole and in most groups above, but none below the family level, corroborating relatively distinct covariation patterns in spite of their high similarity, and also a greater constraint of integration within each family. Diet is associated to phylogeny, and correlates to the evolution of average morphology and integration patterns in the suborder before, but not after correction for phylogenetic relatedness, indicating that dietary changes are part of the shared evolutionary history which shaped the differences in average morphology and integration existing in the group. The characters with covariation patterns more closely associated to diet include parts of the oral and zygomatic skull regions, as well as parts of the neurocranium. Sociality is related to the evolution of average skull morphology, but not to general integration patterns in Feliformia as a whole, although there are correlations between sociality and the selection response patterns of some specific characters, including parts of the neurocranium and measurements related to the zygomatic arch. Mongooses (family Herpestidae) had the evolution of their average skull morphology and integration patterns significantly influenced by both diet and sociality, stressing the importance of these two ecological factors for phenotypic evolution of this group and showing for the first time a direct association between morphological integration in the skull and social behaviour, possibly mediated by aspects related to brain development. The present work thus suggests a complex interaction of factors affecting the evolution of skull morphology in Feliformia, and opens up directions for future work, including analyses of skull modularity patterns in this group and, possibly, comparative analyses of different hypercarnivore morphotypes addressing average skull morphology and morphological integration, as well as diet and possibly other factors

Cette thèse porte sur l'anatomie et la morphologie du complexe crânien chez les murinés. Ses objectifs sont de démontrer et de décrire le lien morpho-fonctionnel entre le crâne et la force de morsure, représentant la performance, d'identifier les sources génétiques de la variation de la performance et de la morphologie, et de comprendre comment la variation morpho-fonctionnelle dépend du mode de vie d'une espèce.La première partie décrit l'anatomie de l'appareil masticateur dans le genre Mus. Des différences fonctionnelles et liées au régime alimentaire sont montrées. Puis, on s'intéresse au lien entre morphologie et fonction, et aux proxies morpho-anatomiques permettant d'estimer la force de morsure. Un modèle biomécanique des muscles masticateurs est d'abord utilisé. Les estimations obtenues sontproches des mesures in vivo au niveau interspécifique, mais moins précises au niveau intraspécifique.L'avantage mécanique et la forme de la mandibule sont ensuite utilisés comme proxies. La force de morsure estimée et réelle sont corrélées au niveau interspécifique, mais moins au niveau intraspécifique, avec des différences selon les espèces. Pour expliquer ces imprécisions, la variation ontogénétique de la force de morsure et de la forme de la mandibule sont décrites. Lorsque l'âge est contrôlé, le développement osseux ralentit plus tôt que celui de la force de morsure, ce qui peut expliquer les biais trouvés dans les estimations de la force de morsure.La deuxième partie se concentre sur les sources génétiques des variations morpho-fonctionnelles. Chez Mus minutoides, des changements sur les chromosomes sexuels produisent des différences de taille et de force de morsure. Les mâles féminisés de cette espèces sont plus agressifs que les autres individus, et montrent une force de morsure plus puissante, principalement grâce à unplus gros crâne. Le(s) gène(s) féminisant produis(ent) donc des changements à l'échelle de l'organisme tout entier. La section suivante s'intéresse aux liens entre consanguinité, asymétrie et performance chez la souris domestique. Les souris les plus consanguines ne montrent pas d'augmentation de l'asymétrie de leurs mandibules, et la performance des souris les plus consanguines ou les plus asymétriques n'est pas affectée. Enfin, l'héritabilité de la force de morsure et de la morphologie est estimée. La force de morsure in vivo n'est pas héritable, bien que la morphologie le soit. Etant donné le lien entre morpho-anatomie et force de morsure, ces résultats suggèrent que les changements morphologiques sont des voies de moindre résistance évolutive, et qu'ils peuvent être à l'origine de changements de performance.La dernière partie utilise la morphologie et la performance en tant que caractères liés à l'écologie des espèces. La première section s'intéresse aux différences de performance entre Apodemus sylvaticus et Mus spretus, qui partagent leur habitat et leurs ressources alimentaires, malgré une différence de taille marquée. Nos résultats montrent un recouvrement des distributions des forces de morsures, soutenant l'hypothèse d'un régime alimentaire commun. Cette coexistence pourrait être due à une abondance de ressources là où ces espèces sont syntopiques. Enfin, la variation morphologique est quantifiée chez plusieurs murinés d'Asie du Sud-Est. Les espèces de rats commensaux et généralistes sont moins variables que les autres espèces. Les espèces synanthropiques montrent des réponses morpho-fonctionnelles convergentes entre localités qui suggèrent qu'elles partagent des patrons communs de réponse et donc qu’elles peuvent s'adapter aux variations environnementales régionales induites par l’homme. L'approche utilisée dans cette thèse nous a permis de montrer le lien entre variation génétique,phénotypique et écologique. Ce lien parfois difficile à décrire, est cependant à la base de l'apparition de nouvelles formes et espèces, et constitue un aspect crucial de la biologie de l'évolution. / The theme of this thesis is the anatomy and morphology of the cranio-mandibular complexin various species of murine rodents. The main objectives are to describe the morpho-functional link between the skull and bite force as a measure of performance, to identify the genetic sources of morphological and performance variation, and finally to understand how morpho-functional variation depends on a species diet and lifestyle. The first part describes the anatomy of the masticatory apparatus in the genus Mus. Differences were found, which could be interpreted functionally and linked to variation in diets. The first part also investigates the links between morpho-anatomy and function, and various morphological proxies used for estimating bite force. This is done by building a biomechanical model of masticatory muscles. The bite force estimates obtained match the in vivo measurements at the inter-specific level, but are less precise at the intra-specific level. Then, two osseous mandibular proxies of bite force are compared (lever arms and their mechanical advantage, and its shape data). In vivo and estimated bite force werewell related at the inter-specific level, but less at the intra-specific level, depending on the species. To explain these imprecisions, the ontogenetic variation of bite force and mandibular morphology isdescribed. Under controlled age, the bony development is slowing down earlier than bite force, whichcan partly explain the inconsistencies of estimated bite force.The second part focuses on the genetics of morphological and functional variation. In Musminutoides, changes in the sexual chromosomes entail size and performance changes. The feminized males found in this species are known to be more aggressive than other individuals, and they produce ahigher bite force, mainly due to an increase in skull size. The feminizing gene(s) therefore drive whole-organism-scale changes. Then, the links between inbreeding, asymmetry and performance areinvestigated in the house mouse. The most inbred mice do not experience an increase in the asymmetry of their mandibles. Contrary to expectations, the performance of the most inbred or most asymmetric mice do not decrease and differences in asymmetry levels have no influence on biting performance.The last section estimates the heritabilities of bite force and morphology. In vivo bite force is notheritable, but some morphological characters are. Given the functional link between morpho-anatomyand bite force shown in the first part, these results suggest that morphological changes represente volutionary pathways of least resistance, and drive changes in performance rather than behavioral or related traits.The last part took morphology and performance as linked to a species' niche. The first sectionexplores the differences between Apodemus sylvaticus and Mus spretus. Both share their habitat and food resources, in spite of a marked size difference. Results show great overlap between their bite force distributions supporting the hypothesis of a shared diet. The absence of shift in a trait related toresource use may be due to a large abundance of the food resources where both species are found insyntopy. In the final section the morphological variation in several rodents from Southeast Asia wasquantified. Less morphological variability is found in generalist and commensal rats by comparison toother species. At the community level, synanthropic species show frequent convergent responsesbetween localities in terms of bite force and morphology. These common patterns in response suggestthat synanthropic species tend to be very adaptable to regional environmental differences.The approaches used in this thesis enable us to show the link between genetic, phenotypic andecological variation. This link, sometimes difficult to describe, is nevertheless at the root of theappearance of new forms and species, and constitutes a crucial aspect of evolutionary biology.

Tropidolaemus wagleri is a species of Asian pitviper with a geographic range including Thailand, Vietnam, Malaysia, Singapore, Bruniei, parts of Indonesia, and the hilippines. Tropidolaemus is a member of the Crotalinae subfamily, within Viperidae. The genus Tropidolaemus includes five species, and was once included within the genus Trimeresurus. While some osteologic characteristics have been noted a comprehensive description of cranial elements has not been produced for T. wagleri. An in-depth description of the cranial skeleton of Tropidolaemus wagleri lays the foundation for future projects to compare and contrast other taxa within Crotalinae and Viperidae. The chosen reference specimen was compared to the presumed younger specimens to note any variation in ontogeny. The study here provides a comprehensive description of isolated cranial elements as well as a description of ontogenetic change within the specimens observed. This study contributes to the knowledge of osteological characters in T. wagleri and provides a foundation for a long term project to identify isolated elements in the fossil record.

A integração morfológica determina a quantidade de variação disponível em diferentes direções do espaço multivariado entre um conjunto de caracteres, e pode assim ter importantes consequências evolutivas, enviesando a direção e a taxa de evolução morfológica em determinadas direções. Assim, estudar a evolução da integração morfológica é importante para entender a evolução fenotípica de uma linhagem. Neste trabalho, utilizo abordagens de genética quantitativa comparativa para estudar a evolução da integração morfológica no crânio em carnívoros da subordem Feliformia, grupo com extensa diversidade filogenética, morfológica e ecológica para o qual análises anteriores de integração analisaram número reduzido de espécies. Para este fim, comparo matrizes de covariância fenotípicas intra-populacionais quantificando a associação entre 35 distâncias lineares no crânio de um conjunto de táxons incluindo todas as sete famílias e mais de um terço da diversidade filogenética atual da subordem. Investigo também a associação entre integração morfológica e a evolução da morfologia média, assim como a influência da história evolutiva (filogenia) e de dois aspectos da ecologia (dieta e socialidade) sobre a evolução da morfologia média e da integração. Os padrões de integração fenotípica no crânio (i.e. quais caracteres variam em conjunto com maior intensidade) são bastante estáveis na subordem, como observado para outros mamíferos, mas revelam alguma sub-estruturação, com padrões um pouco distintos em Felidae, Hyaenidae e Prionodontidae As magnitudes de integração são também relativamente estruturadas, com valores geralmente mais altos em Felidae e consistentemente mais baixos em Hyaenidae. Apesar disso, a magnitude geral de integração é evolutivamente plástica, como em outros mamíferos, havendo importantes variações internas a cada família. Encontrei uma associação geral entre a evolução da morfologia média e modificações em padrões e magnitudes de integração para a subordem como um todo e em vários grupos menos inclusivos, o que em alguns casos se manifesta como uma semelhança da integração entre táxons morfologicamente convergentes, mas filogeneticamente distantes. A evolução de padrões de integração se correlaciona à história filogenética em Feliformia como um todo e na maior parte dos grupos acima, mas não abaixo do nível de família, corroborando padrões de covariação relativamente distintos apesar de sua alta similaridade, bem como integração mais restrita internamente a cada família. A dieta está associada à filogenia, e correlaciona-se à evolução da morfologia média e dos padrões de integração na subordem como um todo antes, mas não após correção levando em conta a proximidade filogenética, indicando que alterações na dieta são parte da história evolutiva compartilhada que levou às diferenças em morfologia média e integração existentes no grupo. Os caracteres cujos padrões de covariação estão mais associados à dieta na subordem incluem partes das regiões oral e zigomática, além de algumas partes do neurocrânio. A socialidade está relacionada à evolução da morfologia craniana média, mas não aos padrões de integração gerais em Feliformia, apesar de existirem associações entre a socialidade e os padrões de resposta a seleção de alguns caracteres específicos, incluindo partes do neurocrânio e distâncias ligadas ao arco zigomático. Os mangustos (família Herpestidae) mostraram influência significativa tanto da dieta quanto da socialidade na evolução de sua morfologia craniana média e de seus padrões de integração, ressaltando a importância desses dois aspectos ecológicos na evolução fenotípica do grupo e mostrando, pela primeira vez, uma associação direta entre a integração no crânio e o comportamento social, possivelmente intermediada por aspectos relacionados ao desenvolvimento do cérebro. O presente trabalho sugere, assim, uma complexa interação de fatores afetando a evolução da morfologia craniana em Feliformia, e indica direções para trabalhos futuros, incluindo análises dos padrões de modularidade craniana no grupo e, possivelmente, análises comparativas em diferentes morfotipos hipercarnívoros abordando a morfologia média e a integração morfológica no crânio, bem como a dieta e possivelmente outros fatores / Morphological integration determines the standing variation available within a lineage in different directions of the multivariate space of a trait set, and can therefore have important evolutionary consequences, biasing the direction and rate of morphological evolution in certain directions. Consequently, studying the evolution of morphological integration is important to understand a lineage\'s phenotypic evolution. Here I employ a comparative quantitative genetic framework to study the evolution of morphological integration in the skull for the suborder Feliformia (Carnivora), a group with remarkable phylogenetic, morphological and ecological diversity for which previous analysis of integration included a limited number of species. To this purpose, I compare within-population phenotypic covariance matrices quantifying the associations between 35 skull linear measurements in a set of taxa including all seven feliform families and over one third of the suborder\'s extant phylogenetic diversity. I also assess the association between morphological integration and the evolution of average skull morphology, as well as the influence of evolutionary history (phylogeny) and two ecological aspects (diet and sociality) on the evolution of average morphology and integration. Skull integration patterns (i.e. which characters covary the most) are quite stable in the suborder, as seen among other mammals, but also have some structuring between groups, with relatively distinct patterns in Felidae, Hyaenidae and Prionodontidae. Integration magnitudes are also relatively structured, with generally higher values in Felidae and consistently lower values in Hyaenidae. Nevertheless, the overall magnitude of integration is evolutionarily labile, as in other mammals, with important variations occurring within each family. I found a general association between the evolution of average morphology and changes in integration patterns and magnitudes for the suborder as a whole and also for many less inclusive groups, which in some cases leads to similar integration between morphologically convergent, but phylogenetically distant taxa. The evolution of integration patterns is correlated to phylogenetic history in Feliformia as a whole and in most groups above, but none below the family level, corroborating relatively distinct covariation patterns in spite of their high similarity, and also a greater constraint of integration within each family. Diet is associated to phylogeny, and correlates to the evolution of average morphology and integration patterns in the suborder before, but not after correction for phylogenetic relatedness, indicating that dietary changes are part of the shared evolutionary history which shaped the differences in average morphology and integration existing in the group. The characters with covariation patterns more closely associated to diet include parts of the oral and zygomatic skull regions, as well as parts of the neurocranium. Sociality is related to the evolution of average skull morphology, but not to general integration patterns in Feliformia as a whole, although there are correlations between sociality and the selection response patterns of some specific characters, including parts of the neurocranium and measurements related to the zygomatic arch. Mongooses (family Herpestidae) had the evolution of their average skull morphology and integration patterns significantly influenced by both diet and sociality, stressing the importance of these two ecological factors for phenotypic evolution of this group and showing for the first time a direct association between morphological integration in the skull and social behaviour, possibly mediated by aspects related to brain development. The present work thus suggests a complex interaction of factors affecting the evolution of skull morphology in Feliformia, and opens up directions for future work, including analyses of skull modularity patterns in this group and, possibly, comparative analyses of different hypercarnivore morphotypes addressing average skull morphology and morphological integration, as well as diet and possibly other factors