Thesis (doctoral)--Albert-Ludwigs-Universität zu Freiburg im Breisgau, 1890. / "Erster Teil einer Untersuchung, die unter dem Title 'Versuch einer Anatomie des Genus Hylobates' erscheinen wird in: Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien, hrsg. von Dr. Max Weber, Professor der Zologie in Amsterdam."

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While gorillas and orangutans have been shown to display considerable manual skill in obtaining certain plant foods, complex feeding skills in chimpanzees have only been described in the restricted context of tool use. This thesis provides the first study of plant-processing skills in a non-tool using community of chimpanzees in Budongo Forest, Uganda. Furthermore, this community contains over 20% of individuals with upper or lower limb injuries. The strategies used by injured individuals in compensating for injury were investigated through a comparison of feeding skill between the able-bodied and injured population. A cognitive approach to feeding behaviour in chimpanzees was adopted, with respect to the implications this may have for overcoming the effects of injury. Chimpanzees were found to employ a broad range of skills in feeding, reflecting variation in their environment and in their diet. Three food types were examined, each illustrating a particular aspect of feeding skill. In processing leaves of Broussonettia papyrifera, chimpanzees use complex multi-stage feeding techniques, employ bimanual co-ordination at several stages and elicit behaviour that is hierarchical in overall organisation. Able-bodied individuals show considerable standardisation in their feeding with a preference for two techniques. In contrast, when feeding on figs, chimpanzees rely upon simple processing techniques but at the same time employ strategies that serve to minimize the effects of feeding competition. In the case of Ficus mucuso chimpanzees co-ordinate several handfuls of food simultaneously between limbs, and with Ficus sur, chimpanzees display a range of dynamic feeding postures and positions in order to access food patches and increase relative food availability. No significant hand preferences were found in any of the three feeding tasks. Even the most severe of injuries does not result in a decline in feeding efficiency, and the possible mechanisms contributing to this were addressed. Injured individuals were found not to invent novel solutions to familiar tasks, but instead to modify their existing repertoire in order to work around their injuries, thus sharing the program-level organisation observed in able-bodied individuals and compensating at the level of individual actions. However, the physical limitations imposed by the injured limb considerably reduce bimanual coordination and manoeuvrability in the tree, which may have long-term negative implications.

Gestural communication in a group of zoo-living lowland gorillas Videotaped observations of a group of zoo-living lowland gorillas collected over a seven- year period were used to study aspects of non-vocal communication. I discerned three classes of gesture: 1) tactile gestures, that depict motion paths iconically; 2) non-tactile silent gestures, some of which appear to be iconic and others deictic; and 3) audible gestures, that, unlike the other two classes, are species-typical gorilla behaviour. The iconic gestures appear to represent activities desired of another gorilla. In addition, one gorilla developed a gesture that was regularly used to suppress the playface, a facial expression that was presumably involuntary. Certain social and environmental conditions, such as the presence of competing males and a physical environment that permits female choice as to proximity with males, may promote the development of such forms of visual communication. I trace the development of gestures throughout the gorilla lifetime, and approach the acquisition of gestures from several viewpoints. "Ontogenic ritualization" and imitation are both found to have a limited explanatory value. One gorilla imitated human gestures, but there was no concrete evidence that these gorillas imitated each other. Repeated strings of gestures or other actions showed, however, that memory capacity exists in gorillas for reproducing complex sequences. Finally, I compare the gestural inventions of my zoo subjects with those of a gorilla taught American Sign Language, finding continuity in styles of depiction from portrayal of pure action to description of stationary objects. Gesture, in portraying action as well as in its ability to depict object shapes, can be seen as a necessary foundation for die eventual development of language in die hominid line.

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Thesis (M.A.)--York University, 2007. Graduate Programme in Psychology. / Typescript. Includes bibliographical references (leaves 40-49). Also available on the Internet. MODE OF ACCESS via web browser by entering the following URL: http://gateway.proquest.com/openurl?url_ver=Z39.88-2004&res_dat=xri:pqdiss&rft_val_fmt=info:ofi/fmt:kev:mtx:dissertation&rft_dat=xri:pqdiss:MR38790

Understanding the success of the human species is central to evolutionary anthropology. While we share many traits with our relatives the great apes, only humans migrated to all corners of the earth and domesticated other species, leading to the emergence of complex societies. Investigations into human genomes have shown that they are a rich source of information for insights into our past. However, for a complete understanding of human evolution it is necessary to look beyond our own genomes. This thesis is about using comparative genomics to place human evolution within a wider context by studying adaptation in our closest living relatives and in the species that we domesticated. In the first study, I investigate the genetic changes involved in the earliest stages of dog domestication. Using a global sample of dog and wolf genomes I identify regions that are highly diverged between these species. I find that selection in the initial stages of dog domestication likely involved genes involved in the fight-or-flight response, advancing our understanding of this process. In the second study, I look for commonalities in the genetic changes that occurred during animal domestication across species. I compare genome sequences from experimentally and historically domesticated species. I identify genes and variants that may underlie the phenotypic changes that occurred during domestication. I find evidence of biological pathways that appear to always be involved in the domestication process. In the third study, I characterise the signatures of natural selection in all major Hominidae lineages using population genomic data. I find that most signatures of positive selection are species specific, although some loci appear to be selected across several lineages. I determine that the efficacy of selection varies between species and is significantly correlated with long-term effective population size. These results contribute to a more complete understanding of human evolution. i This thesis is based on the following manuscripts: 1. Cagan A & Blass T. (2016) Identification of genomic variants putatively targeted by selection during dog domestication. BMC Evolutionary Biology,16:1. 2. Cagan A, Albert FW, Plyusnina I, Trut L, Renaud G, Romagné F, Wiebe V, Kozhemjakina R, Gulevich R, Trapezov O, Yudin N, Alekhina T, Aitnazarov R, Trapezova L, Herbeck Y, Schöneberg T, Pääbo S. Genes and pathways selected during animal domestication. Submitted to eLife. 3. Cagan A, Theunert C, Laayouni H, Santpere G, Pybus M, Casals F, Prüfer K, Navarro A, Marques-Bonet T, Bertranpetit J, Andrés AM. (2016). Natural Selection in the Great Apes. Molecular Biology and Evolution, 33:3268-3283.:Summary 1 Zusammenfassung 8 Chapter 1 Identification of genomic variants putatively targeted by selection during dog domestication 16 Chapter 2 Genes and pathways selected during animal domestication 38 Chapter 3 Natural Selection in the Great Apes 73 References 149 Acknowledgments 152 Curriculum Vitae 153 Declaration of Independence 156 Author Contribution Statements 157

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A unique pattern of pedal loading from heel-strike at touchdown to hallucal propulsion at toe-off is a distinct feature of mature human bipedalism, however, its first appearance in the fossil record is debated. The main goal of this dissertation is to identify anatomical correlates to a modern human heel-strike, rigid foot, and propulsive hallucal toe-off. First, a biomechanical analysis of toddler walking is used as a 'natural experiment' to investigate the influence of non heel-strike (NHS, n = 11) and immature heel-strike (IHS, n = 7) on the location of the center of pressure and orientation of the ground reaction force resultant in relation to specific foot bones during stance phase. With an expanded knowledge of foot bone loading in toddlers, a microarchitectural approach is used to test the influence of a heel-strike, rigid foot, and propulsive hallucal toe-off on trabecular bone fabric properties in an ontogenetic series of human and African ape (chimpanzee, bonobo, and gorilla) calcanei, tali, first metatarsal heads and hallucal distal phalanges. This dissertation presents the first ontogenetic analysis of pedal trabecular bone in primates. Heel-strike and toe-off are developmentally independent from one another. Although most toddlers lack a hallucal toe-off, NHS and IHS apply equally high propulsive forces when the entire width of their forefoot is in contact with the ground. Biomechanical and fossil evidence suggest that a generalized active propulsion may have preceded the evolution of a propulsive hallucal toe-off. Although pedal trabecular fabric properties are more complex than predicted, trabecular correlates to heel-strike and hallucal toe-off are identified within adult human foot bones. Compared to toddlers and African apes, adult humans have a unique combination of relatively thick trabecular struts and an anteroplantar to posterodorsal primary trabecular orientation in the plantar aspect of the calcaneal tuber. In the calcaneal tendon volume of interest, adult humans have a unique anteroplantar to posterodorsal primary trabecular orientation associated with a propulsive hallucal toe-off. This dissertation provides the comparative context necessary to begin assessing the evolution and developmental timing of foot function and specific bipedal gait events in juvenile and adult fossil hominins. / text

This thesis has the overall aims of creating a welfare-sensitive additive index of captive great ape husbandry using the expert opinion of relevant stakeholders, highlighting the aspects of husbandry requiring most improvement and allowing enclosures in different zoos and wildlife parks to be ranked in a uniform way. I investigated the attitudes of experts on the management of captive great apes to gain a greater understanding of the features of their environment that may be critical in maintaining a high standard of welfare. From the experts ranking of the different Levels of facilities offered to the great apes I constructed an additive index with 17 key attributes with between two and four Levels for each. The Great Ape Welfare Index (GAWI) derived from expert opinion has Social structure, Enclosure appearance, Group size, Enclosure furnishings and Avoidance provision as the most important attributes of a management system. The GAWI was then validated by recording the behaviour of the captive great apes housed in Australian and overseas zoos, and comparing their activity budgets to wild individuals. Using a Penrose Distance Statistic a significant negative correlation was found between Index score and the departure from wild activity budgets. Additionally I found a significant negative correlation between the change in the proportion of time spent resting and the welfare Index score achieved indicating that those great apes housed in enclosures achieving higher Index scores are more active in general. Finally the husbandry and management systems for great apes currently used by zoos globally were investigated and their ranking in the Index were calculated. This allowed the particular aspects of great ape management requiring improvement to be identified for each enclosure. Comparisons of regional means showed that Australian zoos achieved significantly higher Index scores than those in other parts of the world. The proportion of great apes housed in New Zealand zoos reported as displaying an abnormal behaviour was significantly higher than all other regions with the proportion of great apes housed in Australia displaying significantly less abnormal behaviour when compared to the US. The highest index achieved by any enclosure was 97.5 for an enclosure that housed a group of gorillas, with the lowest index allocated was 52.2 for an orangutan enclosure. The presence of abnormal behaviours was also investigated with the most common abnormal behaviours reported being regurgitation and re-ingestion (15.8%), hair plucking (10.2%) and begging (7.9%) with some animals reported as exhibiting four different abnormal behaviours. The GAWI has the potential to improve the captive environments provided to great apes through highlighting the most critical aspects of their husbandry requiring improvement. This should encourage zoos to aim higher than just complying with minimum standards set by their regional zoological associations. As the index correlates with the behaviour of the great apes housed, their behaviour may more closely resemble wild individuals if improvements are made. The GAWI has been compared to two alternative models to investigate whether the methods used to create the index could be simplified while still allowing adequate discrimination between enclosures and species. This index can be adapted for use in other institutions housing great apes and can act as a model to create indices for other species. The index has been submitted to the primate Taxon Advisory Group of the Australian Regional Association of Zoological Parks and Aquaria for their consideration of its use in their accreditation scheme or husbandry guidelines.