This thesis is a cross-disciplinary project, drawing on the techniques of behavioural ecology to reconstruct aspects of the socioecology of the early hominids. The modelling approach advocated in this project is an attempt to move towards the conceptual modelling approach forwarded by Tooby & DeVore (1986), moving away from referential and phylogenetic primate models to reconstruct hominid social behaviour. The project contributes to one current aim in palaeoanthropology; to develop a theoretical framework with which to reconstruct the behaviour of extinct taxa. The raw data for these models come from comparative analyses of behavioural ecology of extant primates. The fundamental basis of systems models of socioecology, is that the relationship between the environment and behaviour is characterised. The choice of environmental parameters has proved to be crucial, therefore I have characterised the key environmental variables that affect animals both directly (e.g. thermoregulatory stress), and indirectly (e.g. via habitat productivity). The quantitative relationships found in this chapter will serve as useful constants for further models. I first present a re-analysis of the systems models of baboons, refining previous models by R. Dunbar. The statistical techniques underlying these linear program models was further supported by the stability of the models when new data were included. Long-term climate data that were accurately sited were found to produce the most predictive equations. The results of the reanalysis of the baboon models gave support to their extension to another taxa. I then extend the range of the time-budget based models to the chimpanzees. The initial focus on chimpanzees is not solely because they are phylogenetic analogues. The emphasis of the models is that they are habitat specific. Relationships between behaviour and environment are used to develop functional equations to explore an animals flexibility of response to varying environmental conditions. Predictions could be made about: maximum ecologically tolerable group size, territory size, diet and extent of geographic range. The geographic distribution of chimpanzees predicted by the model was found to match very closely the current distribution. The models of chimpanzee, baboon and gelada socioecology were then compared. Dietary differences between the taxa accounted for their ecological niche separation. The models of extant primate systems models developed in this thesis provide a firm foundation for extending the models to extinct taxa. Preliminary models are presented, extending these analyses to the extinct australopithecines to forward the development of a conceptual model for the early hominids.

The aim of this dissertation is to analyse two notions which inform contemporary evolutionary psychology. In Part I Tooby and Cosmides' (1992) Standard Model thesis of the history of twentieth century social science is examined with regard to social psychology. In Part II the practical and theoretical fecundity of the Environment of Evolutionary Adaptedness is examined, again with regard to social psychology. The analysis of the Standard Model thesis yields the result that it is not reliable as an intellectual history of social psychology. A principal reason for this is the failure of the thesis to acknowledge the instinct debate of the late nineteenth and early twentieth century. Further consideration of the instinct debate leads to the conclusion that evolutionary psychology may be in the process of repeating the history of social psychology rather than making substantive advances. The analysis of the Environment of Evolutionary Adaptedness concept yields two results. Firstly, in use it fails to accommodate the findings of palaeontology. Secondly, it promotes a view of mental capacity and functioning that is at odds with that of modern humans. Further consideration of the natural history of the human lineage leads to the conclusion that the past was not, in some sense, ontogenetically prior to the present and that it will not furnish social psychology with an adaptation that functions in a predictable manner. In Part III it is recommended that an evolutionary approach to social psychology should dispense with the concept of adaptation as proposed by evolutionary psychology.

Il existe chez les espèces du genre Homo une diversité morphologique crânienne et cérébrale importante, et les interactions de ces deux éléments sont complexes. De manière générale, au cours de l’évolution de ce taxon, le neurocrâne prend une importance croissante par rapport au bloc facial en raison d’une expansion cérébrale marquée. Cependant, les modalités de cette expansion sont multiples, et elle se met en place chez les différentes espèces via des modifications morphologiques qui leur sont propres. Mise à part l’augmentation du volume cérébral, l’endocrâne témoigne de réorganisations neuroanatomiques. Ces différents facteurs - volume et organisation – ainsi que les contraintes morpho-fonctionnelles diverses exercées sur la face externe du crâne, sont susceptibles de résulter en une variété de relations morphologiques et spatiales entre le neurocrâne et l’endocrâne. Il est donc pertinent de documenter ces relations afin de pouvoir par la suite mieux appréhender la variabilité et les mécanismes évolutifs à l’oeuvre chez les différents taxons du genre Homo. Nous explorons dans ce travail de thèse les variations jointes du neurocrâne et de l’endocrâne dans le genre Homo et chez Homo sapiens. Cette contribution est basée sur l’analyse de modèles virtuels de crânes et d’endocrânes à l’aide de méthodes géométriques et d’une méthode innovative de déformations de surfaces. Nous avons étudié des données morphologiques issues de populations actuelles afin d’éclaircir la nature des relations entre le neurocrâne et l’endocrâne chez Homo sapiens. Pour cela, nous avons comparé les asymétries des hémisphères de l’endocrâne – qui reflètent celles des hémisphères cérébraux – aux asymétries de la voûte crânienne. Les schémas d’asymétrie bilatérale relevés sont identiques sur le crâne et sur l’endocrâne. Cela s’explique par une morphologie de la voûte du crâne calquée sur celle de l’endocrâne, malgré un effet « tampon » de l’os qui n’enregistre pas sur sa face externe toutes les asymétries cérébrales. Les possibles corrélations entre le degré d’asymétrie et des facteurs tels que la conformation générale du crâne, la robustesse des superstructures osseuses, le sexe et le volume endocrânien ont également été explorées. Nous avons ensuite analysé les schémas de co-variation entre neurocrâne et endocrâne au sein du genre Homo. Nous avons ainsi pu mettre en évidence des éléments de co-variation qui concernent l’ensemble du genre Homo, et d’autres qui sont spécifiques à certains taxons, notamment aux Néandertaliens ou à Homo sapiens. Ainsi, si la conformation de la voûte crânienne est très semblable à la morphologie endocrânienne, les interactions crâne-endocrâne dans la zone occipitale et cérébelleuse apparaissent plus variables, et semblent inféodées au degré de globularisation de l’ensemble du cerveau et du neurocrâne. Ces résultats mettent en évidence certaines interactions entre réorganisations cérébrales et morphologie crânienne chez les différentes espèces du genre Homo, et soulignent le caractère crucial du croisement des données et des méthodes pour l’interprétation du registre fossile. / Species of the genus Homo display cranial and endocranial morphological variations, with complex interactions between these two elements. Generally speaking, throughout the evolution of this taxon the neurocranium becomes increasingly important by comparison with the facial skeleton, due to a marked cerebral expansion. The modalities of this expansion differ accross species and occur at least partly through species-specific morphological processes. Apart from the increase in cerebral volume, the endocast bears the traces of neuroanatomical reorganisations. These two factors – volume and organisation – as well as various morpho-functional constraints on the external face of the cranium, may result in a variety of morphological and spacial relationships between the neurocranium and the endocranium. It is therefore important to document these relations in order to better apprehend the variability and the evolutionary mechanisms behind the morphologies of the different Homo species.This doctoral thesis explores the joint morphological variations of the neurocranium and endocast in the genus Homo and within Homo sapiens through multiple approaches. We offer a contribution to this topic based on shape analyses of virtual crania and endocasts, using geometric morphometrics and an innovative technique of surface deformations. We analysed morphological data from extant populations in order to clarify the nature of the relationship between the neurocranium and the endocast in Homo sapiens. One of the lines of evidence investigated is the correspondence between neurocranial and endocranial (and therefore cerebral) gross asymmetries. Our results show that the patterns of bilateral asymmetries of the neurocranium are identical to those of the endocranium. There is a close correspondance between the morphologies of the endocranial and cranial vaults, despite the neurocranium not displaying the full extent of cerebral asymmetries on its external vault. Correlations between asymmetry and factors including sex, endocranial volume and importance of the bony superstructures were also tested. Co-variation patterns between neurocranial and endocranial morphologies in the genus Homo were analysed. Our results highlight elements of co-variation between the neuro and endocranium which are shared accross the genus Homo, and others which are species-specific. While the cranial vault closely follows endocranial morphology, interaction patterns between the endo and neurocranium in the occipito-cerebellar area appear more variable and linked to the overall degree of globularisation of the brain and neurocranium. These results highlight some of the interactions between cerebral reorganisations and cranial morphology in Homo species, and underline the importance of crossing data and methods in order to interpret the fossil record.

<p>Unfortunately I have had bit of difficulty with setting up the page numbers appropriately. I have been unable to figure out how to make the preliminary part of document in Roman numerals while at the same time separating this from the main body of the text using Arabic numbers. My apologies, but I sincerely do not know how to resolve this frustrating problem. I can resolve this problem if I send you the preliminary part and main body as separate documents, but just not as one continuous document. Please contact me if this latter method is the way to go.</p> / <p>This thesis provides an expository account and critical analysis of Friedrich Nietzsche’s genealogical position on cruelty. Its primary engagement is with Nietzsche’s <em>On The Genealogy of Morals</em>, however, other works by this author are discussed when relevant. The general import of this thesis is threefold. First, it demonstrates Nietzsche’s genealogical account of cruelty, detailing its complex evolutionary progression and its various facets of influence. Second, this work identifies some authors who are critical of Nietzsche position on cruelty. These criticisms are identified and are then largely refuted on various grounds. Third, this thesis argues that an appropriate critical analysis of Nietzsche’s genealogical theorizing will be based on a cross-examination of his positions with current palaeoanthropological findings. The conclusion drawn from this analysis is that there is insufficient empirical evidence to substantiate Nietzsche’s accounts and his methodical approach to genealogical theorizing is, furthermore, untenable.</p>

The complex demographic changes that underlie the expansion of anatomically modern humans out of Africa have important consequences on the dynamics of natural selection and our ability to detect it. In this thesis, I aimed to refine our knowledge on human population history using ancient genomes, and then used a climate-informed, spatially explicit framework to explore the interplay between complex demographies and selection. I first analysed a high-coverage genome from Upper Palaeolithic Romania from ~37.8 kya, and demonstrated an early diversification of multiple lineages shortly after the out-of-Africa expansion (Chapter 2). I then investigated Late Upper Palaeolithic (~13.3ky old) and Mesolithic (~9.7 ky old) samples from the Caucasus and a Late Upper Palaeolithic (~13.7ky old) sample from Western Europe, and found that these two groups belong to distinct lineages that also diverged shortly after the out of Africa, ~45-60 ky ago (Chapter 3). Finally, I used East Asian samples from ~7.7ky ago to show that there has been a greater degree of genetic continuity in this region compared to Europe (Chapter 4). In the second part of my thesis, I used a climate-informed, spatially explicit demographic model that captures the out-of-Africa expansion to explore natural selection. I first investigated whether the model can represent the confounding effect of demography on selection statistics, when applied to neutral part of the genome (Chapter 5). Whilst the overlap between different selection statistics was somewhat underestimated by the model, the relationship between signals from different populations is generally well-captured. I then modelled natural selection in the same framework and investigated the spatial distribution of two genetic variants associated with a protective effect against malaria, sickle-cell anaemia and β⁰ thalassemia (Chapter 6). I found that although this model can reproduce the disjoint ranges of different variants typical of the former, it is incompatible with overlapping distributions characteristic of the latter. Furthermore, our model is compatible with the inferred single origin of sickle-cell disease in most regions, but it can not reproduce the presence of this disorder in India without long-distance migrations.