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A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfillment of the requirements for the degree of Doctor of Philosophy / This thesis is a study of quantifiable variation in the geometric shape of the superior articular surface of the talus of higher primates, with special reference to fossil tali of Plio- Pleistocene hominids. (Abbreviation abstract ) / AC2017

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Studies of comparative morphology clearly distinguish the shoulder morphology of Homo from that of the other hominoids. While the shoulder morphology of non-human hominoids is thought to signal adaptations to arboreal locomotion, human shoulder morphology is understood to have lost this adaptation during hominin evolution. Ideas how non-human hominoid shoulder morphology is advantageous in an arboreal context suggest that the specific shoulder morphological traits enhance the arm-raising mechanism. However, this idea has not been biomechanically tested. This thesis constitutes the first analysis of the biomechanical consequences of two distinct shoulder morphologies within Hominoidea by comparing the glenohumeral muscle capabilities of Gorilla to Homo. The biomechanical capabilities are evaluated by constructing a computational musculoskeletal model of a gorilla thorax, shoulder girdle and upper arm, which is used to predict relevant biomechanical metrics such as muscle moments and moment arms. Muscle moments and moment arms are predicted for two important mechanisms, arm-raising and arm-lowering. The predictions are compared to those of an already existing human musculoskeletal model in order to evaluate differences in arm-raising and arm-lowering capability based on the two distinct thorax and shoulder girdle morphologies. The results of the biomechanical analyses show that the arm-lowering mechanism is enhanced in Gorilla compared to Homo, instead of the arm-raising mechanism. The enhanced arm-lowering mechanism is evident by greater moment capacities of two important arm-lowering muscles, pectoralis major and teres major. The greater moments are the result of greater muscle force capacities and greater moment arms, due to the beneficial musculoskeletal geometry of Gorilla. The results highlight that a more distal muscle insertion along the humerus has the greatest enhancing effect on the arm-lowering moment arms of teres major and pectoralis major. Furthermore, thorax and shoulder girdle morphological traits that are well known to distinguish non-human apes from humans were found to contribute to the enhancement of the arm-lowering mechanism. The more cranially oriented glenoid, obliquely oriented scapular spine and cranial scapula position on the thorax enabled certain muscles to act as arm-lowering muscles in Gorilla, contrary to the arm-raising action capability that is predicted for Homo. The enhanced arm-lowering capability is likely advantageous for the arboreal locomotion of apes. During hoisting behaviours that are known to occur during suspension and vertical climbing, arm-lowering is used to lift the heavy body of the apes upward. The results of this thesis in conjunction with earlier EMG studies suggest those muscles which are highly activated during these hoisting behaviours also have enhanced arm-lowering capacities in Gorilla and potentially other non-human hominoids compared to Homo. As such, the results highlight shoulder morphological traits that are biomechanically important for the arboreal locomotor behaviour of apes. By this, the thesis demonstrates a link between the conformation of shoulder morphological traits and their biomechanical capability, which will aid future functional interpretations of extant and extinct species.:Acknowledgements Bibliographische Darstellung Summary Zusammenfassung Chapter 1: Exploring the functional morphology of the Gorilla shoulder through musculoskeletal modelling Chapter 2: Comparison of the arm-lowering performance between Gorilla and Homo through musculoskeletal modeling Conclusion Appendix A: Supplementary Information for Chapter 1 Appendix B: Supplementary Information for Chapter 2 Appendix C: Curriculum Vitae Appendix D: Author Contributions

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The Trivers-Willard hypothesis predicts a sex ratio bias contingent on maternal condition in species characterized by variation in male reproductive success. A male-biased sex ratio among mothers in good condition, and a female-biased sex ratio among mothers in poor condition is expected. Studies in humans have thus far provided mixed answers to the question of whether or not sex ratio is affected by maternal condition. The present study assessed whether or not the introduction of a western cash economy influenced the observed secondary sex ratio in Nepal's Khumbu region. Because acculturated villages provided better access to the cash economy and to health facilities, residence in an acculturated village was used as a proxy for "good" maternal condition. I analyzed demographic data gathered by survey in 1971 and 1982. The sample included 734 children from the 1971 survey and 1598 children from the 1982 survey. Using Poisson regression I analyzed the extent to which the sex ratios in age-stratified groups differed between the acculturated and unacculturated villages. In the 1971 dataset, the younger women in the acculturated villages displayed a significantly higher (p=.014) proportion of male offspring. It is likely that older women were subjected to minimal acculturation effects during their child-bearing years and among these data there was a lack of significant deviation between acculturated and unacculturated post-menopausal women. The rapid overall increase in acculturation between 1971 and 1982 likely made conditions in the two sets of villages much more similar by 1982. The results of this study underscore the impact that the transition to a market economy had on women in Nepal's Khumbu region. / Anthropology

Chance events began with the Big Bang and influenced all subsequent evolutionary processes, including the genesis and evolution of biological life. The complex and versatile human brain has come to dominate the environment. Self-awareness generated the idea of spiritual identity and survival beyond death. The assumption of demi-god status gave sanctity and dignity to humans, a moral separation that justified the exploitation of other animals. But evolution has a common origin, fleshed out in the biological similarities of all mammals, including humans. Neurological and biochemical correlates suggest the omnipresence of consciousness and self-consciousness at levels that vary according to nervous system complexity. Accepting a more inclusive morality does not diminish the value of human life. It acknowledges their power to adapt to changing conditions. Human survival depends on change and human lives can only benefit from a more inclusive worldview that considers the welfare of all sentient beings and the sustainability of a shared environment. / Philosophy and Systematic Theology / M.A. (Philosophy)

Reconstructing the origins and evolutionary journey of humans is a central piece of biology. Complementary to archeology, population genetics studying genetic variation among individuals in extant populations has made considerable progress in understanding the evolution of our species. Particularly, studies in indigenous humans provide valuable insights on the prehistory of humans because their life history closely resembles that of our ancestors. Despite these efforts, it can be difficult to disentangle population genetic inferences because of the interplay among evolutionary forces, including mutation, recombination, selection, and demographic processes. To date, few studies have adopted a comprehensive framework to jointly account for these confounding effects. The shortage of such an approach inspired this dissertation work, which centered on the development of model-based analysis and demonstrated its importance in population genetic inferences. Indigenous African Pygmy hunter-gatherers have been long studied because of interest in their short stature, foraging subsistence strategy in rainforests, and long-term socio-economic relationship with nearby farmers. I proposed detailed demographic models using genomes from seven Western African Pygmies and nine Western African farmers (Appendix A). Statistical evidence was shown for a much deeper divergence than previously thought and for asymmetric migrations with a larger contribution from the farmers to Pygmies. The model-based analyses revealed significant adaption signals in the Pygmies for genes involved in muscle development, bone synthesis, immunity, reproduction, etc. I also showed that the proposed model-based approach is robust to the confounding effects of evolutionary forces (Appendix A). Contrary to the low-latitude African homeland of humans, the indigenous Siberians are long-term survivors inhabiting one of the coldest places on Earth. Leveraging whole exome sequencing data from two Siberian populations, I presented demographic models for these North Asian dwellers that include divergence, isolation, and gene flow (Appendix B). The best-fit models suggested a closer genetic affinity of these Siberians to East Asians than to Europeans. Using the model-based framework, seven NCBI BioSystems gene sets showed significance for polygenic selection in these Siberians. Interestingly, many of these candidate gene sets are heavily related to diet, indicating possible adaptations to special dietary requirements in these populations in cold, resource-limited environments. Finally, I moved beyond studying the history of extant humans to explore the origins of our species in Africa (Appendix C). Specifically, with statistical analyses using genomes only from extant Africans, I rejected the null model of no archaic admixture in Africa and in turn gave the first whole-genome evidence for interbreeding among human species in Africa. Using extensive simulation analyses under various archaic admixture models, the results suggest recurrent admixture between the ancestors of archaic and modern Africans, with evidence that at least one such event occurred in the last 30,000 years in Africa.

The chimpanzee (Pan troglodytes, Pongidae) among all other living species, is our closest relation, with whom we last shared a common ancestor less than five million years ago. These African apes make and use a rich and varied kit of tools. Of the primates, and even of the other Great Apes, they are the only consistent and habitual tool-users. Chimpanzees meet the criteria of working definitions of culture as originally devised for human beings in socio-cultural anthropology. They show sex differences in using tools to obtain and to process a variety of plant and animal foods. The technological gap between chimpanzees and human societies living by foraging (hunter-gatherers) is surprisingly narrow, at least for food-getting. Different communities of chimpanzees have different tool-kits, and not all of this regional and local variation can be explained by the varied physical and biotic environments in which they live. Some differences are likely customs based on non-functionally derived and symbolically encoded traditions. Chimpanzees serve as heuristic, referential models for the reconstruction of cultural evolution in apes and humans from an ancestral hominoid. However, chimpanzees are not humans, and key differences exist between them, though many of these apparent contrasts remain to be explored empirically and theoretically.