Finding food in tropical forests poses a potentially major problem for chimpanzees, whose ranging is thought primarily to be directed at locating suitable food resources: (1) chimpanzees are frugivorous, large bodied and live in large home ranges; (2) they lack specialised sensory or locomotor abilities, and terrestrial travel is known to be costly; but (3) fruits are randomly distributed in space and time. Evidence from studies of captive individuals suggests chimpanzees are capable of remembering the locations of out of sight resources and can compute least distance routes to these resources, but whether this ability translates to the natural foraging behaviour of wild chimpanzees has never been investigated. My observational study was designed to assess how the chimpanzees (Pan troglodytes schweinfurthii) of Budongo Forest, Uganda, locate these patchy resources. I mapped the routes of 14 focal individuals over a 12-month period. I considered how these foraging routes were structured by breaking the path into segments of travel between resources. Consecutive segments of travel between resources were found not to be independent, but assembled into "super-segments" that take in a number of resources along one trajectory. These super-segments are not necessarily directed towards feeding resources, however: travel is not always food directed. Comparisons of actual chimpanzee routes with randomly generated simulations suggest most individuals do not attempt to minimise their travel distances. There is evidence to suggest energetically stressed individuals can remember the locations of recently visited food resources and return to these patches in order to minimise travel distances when necessary, but overall, food is not difficult to find for this community of chimpanzees. I propose this is because males defend a territory with super-abundant food resources, meaning availability is not a limiting factor of foraging. Male chimpanzees can be characterised as convenience feeders, taking food whilst satisfying other, social needs.

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.

Ability of human listeners to understand altered speech is argued as evidence of uniquely human processing abilities, but early auditory experience also may contribute to this capability. I tested the ability of Panzee, a language-trained chimpanzee (Pan troglodytes), reared and spoken to from infancy by humans, to recognize synthesized words. Training and testing was conducted with different sets of English words in natural, “harmonics-only” (resynthesized using only voiced components), or “noise-vocoded” (based on amplitude-modulated noise bands) forms, with Panzee choosing from “lexigram” symbols that represented words. In Experiment 1 performance was equivalent with words in natural and harmonics-only form. In Experiment 2 performance with noise-vocoded words was significantly higher than chance but lower than with natural words. Results suggest specialized processing mechanisms are not necessary to speech perception in the absence of traditional acoustic cues, with the more important factor for speech-processing abilities being early immersion in a speech-rich environment.

No description available.

In the wild, a variety of inter-group behavioural differences have been reported for chimpanzees (Pan troglodytes) and it has been suggested that these may have resulted from social learning. To determine whether chimpanzees show the necessary propensity for social learning, groups of captive chimpanzees were tested in a series of experiments involving the use of two-action and bidirectional apparatuses. For comparison, and to shed light on any contrasts between our own and chimpanzee learning strategies, similar tests were also conducted with children (Homo sapiens) to ascertain the nature of their observational learning when watching conspecifics. Through the use of open diffusion and diffusion chain techniques, it was shown that both species learnt how to operate different foraging devices from observing an expert conspecific and this learning was strong enough for the generation of behavioural traditions which passed along multiple test ‘generations’. Additionally, ghost conditions were used to distinguish imitative and emulative learning by both species. With one of the two test devices used (the Slide-box) the first evidence for emulation learning by chimpanzees, through the use of a ghost condition, was shown. Children in this condition also showed apparent emulation; a contrast to previous research which has concluded that children tend to rely on imitation. Additionally, to test its potential for use in future social learning experiments, the ability of chimpanzees to learn from video-footage of an unknown conspecific was tested. It was found that the chimpanzees not only learnt how to operate two devices from observing this footage but also used the same alternative method used by the model chimpanzee.

A separação histórica entre as ciências naturais e as ciências sociais fundamenta-se na distinção ontológica entre os domínios da natureza e da cultura, e na ideia moderna de que a condição (cultural) humana corresponde a um afastamento radical dos outros animais. Porém, somando-se a outros críticos insatisfeitos com essa visão dualista, muitos estudiosos do comportamento animal tem utilizado o termo cultura em referência a não humanos, provocando uma controvérsia que ainda parece longe de um consenso. Neste trabalho, investiguei o sentido da noção de cultura para os antropólogos e o uso etológico (limitando-nos aos primatas) do termo, com os objetivos de compreender melhor a controvérsia e identificar caminhos possíveis na busca por um consenso. Na Antropologia, a noção moderna de cultura se desenvolveu do século XIX até os anos 1950. Cultura passou a ser vista como um fenômeno emergente exclusivamente humano, dependente de nossa capacidade de utilizar símbolos e correspondendo aos padrões e normas comportamentais, artefatos, ideias e, principalmente, valores que os indivíduos adquirem no processo de socialização. Mais recentemente, essa concepção de cultura, e a epistemologia dualista que a sustenta, tem sido alvo de críticas e intenso debate. Ainda que não compartilhem um arcabouço teórico comum, virtualmente todos os antropólogos contemporâneos concordam que o comportamento cultural humano é fundamentalmente simbólico. A discussão recente em torno da atribuição de cultura a primatas não humanos remonta aos estudiosos japoneses que, na década de 1950, acompanharam a dispersão de uma nova técnica de manipulação de alimento em Macaca fuscata, e descreveram o fenômeno com os termos pré-cultura, subcultura e cultura infra-humana. A partir da década de 1960, as pesquisas de campo com populações selvagens e as evidências experimentais de aprendizagem em contexto social levaram ao estabelecimento da Primatologia Cultural e os prefixos foram abandonados. Entre primatólogos, o termo cultura se refere a padrões comportamentais que dependem de um contexto social para se desenvolver, e que podem atravessar gerações. Eu sugiro uma estratégia analítica que distingue os motivos de discordância entre descrições, explicações, teorias e visões de mundo, e argumento que a controvérsia é complexa e inclui discordâncias entre visões de mundo sem, no entanto, dividir os envolvidos em grupos homogêneos (digamos, primatólogos contra antropólogos). Por conta disso, a redefinição e o uso que os primatólogos fazem do termo acabam por manter ilesos os fundamentos da dicotomia natureza/cultura, o que pode explicar, parcialmente, a manutenção da controvérsia. Concluo que o diálogo entre os dois lados da fronteira será imprescindível para os pesquisadores que estiverem interessados em buscar uma abordagem consensual. É possível alcançar um consenso, mas a busca por uma abordagem sintética do comportamento animal que inclua os humanos deverá levar ao abandono ou reconstrução das dualidades natureza/cultura, inato/adquirido e gene/ambiente, e também da atribuição de primazia causal aos genes. Além disso, é necessário discutir a fundo sobre como incluir a questão do simbolismo e do significado em uma perspectiva comparativa / The Western ontological distinction between nature and culture, and the idea that the human (cultural) condition makes us radically different from other animals, are evident in the historical separation between the natural and social sciences. In parallel to other critics of this dualist view, some animal behaviorists have been using the term culture in relation to nonhumans, starting a controversy that is still far from cooling down. In this study, I investigated the meaning of the term culture as used by anthropologists, and also its recent use by ethologists (limiting myself to primatology), in order to better understand the controversy and identify possible paths that might lead to a consensus. In Anthropology, the modern concept of culture developed between the 19th century and the 1950s. It came to be seen as an emergent phenomenon exclusive to human social life. It was dependent on our capacity to use symbols and corresponded to behavioral patterns and norms, artifacts, ideas, and values that individuals acquire in the process of socialization. But this conception of culture, and the dualist epistemology supporting it, have since been largely criticized and intensely debated. Although contemporary anthropologists do not share a common ground or framework, virtually all of them agree that human cultural behavior is fundamentally symbolic. Recent attribution of culture to nonhuman primates started with Japanese scholars who, from the 1950s onward, have followed closely the spread of novel behaviors in Macaca fuscata, which they described with expressions such as preculture, subculture and infrahuman culture. Since the 1960s, field studies on wild populations and experimental research on learning in a social context, have led to the establishment of Cultural Primatology, and the prefixes were abandoned. Among primatologists, the term culture refers to behavioral patterns that depend on the social context to develop and that might be recurrent through generations. I suggest that it might be analytically useful to distinguish the matters of a disagreement between descriptions, explanations, theories and worldviews, and argue that this controversy goes all the way up to the highest reason of disagreement (worldviews). Still, one cannot divide those involved in it into a few homogeneous groups (say, primatologists contra anthropologists). Primatologists redefinition and use of the term do not alter the foundations of the criticized nature/culture dichotomy, and that might at least partially explain the maintenance of the controversy. It is possible to reach a consensus, but the search for a synthetic framework for animal behavior that includes humans might lead to the abandonment or reconstruction of the related dichotomies of nature/culture, innate/acquired and gene/environment, as well as of the causal primacy attributed to genes. It is also necessary to discuss how to include symbols and meanings in a comparative perspective

A separação histórica entre as ciências naturais e as ciências sociais fundamenta-se na distinção ontológica entre os domínios da natureza e da cultura, e na ideia moderna de que a condição (cultural) humana corresponde a um afastamento radical dos outros animais. Porém, somando-se a outros críticos insatisfeitos com essa visão dualista, muitos estudiosos do comportamento animal tem utilizado o termo cultura em referência a não humanos, provocando uma controvérsia que ainda parece longe de um consenso. Neste trabalho, investiguei o sentido da noção de cultura para os antropólogos e o uso etológico (limitando-nos aos primatas) do termo, com os objetivos de compreender melhor a controvérsia e identificar caminhos possíveis na busca por um consenso. Na Antropologia, a noção moderna de cultura se desenvolveu do século XIX até os anos 1950. Cultura passou a ser vista como um fenômeno emergente exclusivamente humano, dependente de nossa capacidade de utilizar símbolos e correspondendo aos padrões e normas comportamentais, artefatos, ideias e, principalmente, valores que os indivíduos adquirem no processo de socialização. Mais recentemente, essa concepção de cultura, e a epistemologia dualista que a sustenta, tem sido alvo de críticas e intenso debate. Ainda que não compartilhem um arcabouço teórico comum, virtualmente todos os antropólogos contemporâneos concordam que o comportamento cultural humano é fundamentalmente simbólico. A discussão recente em torno da atribuição de cultura a primatas não humanos remonta aos estudiosos japoneses que, na década de 1950, acompanharam a dispersão de uma nova técnica de manipulação de alimento em Macaca fuscata, e descreveram o fenômeno com os termos pré-cultura, subcultura e cultura infra-humana. A partir da década de 1960, as pesquisas de campo com populações selvagens e as evidências experimentais de aprendizagem em contexto social levaram ao estabelecimento da Primatologia Cultural e os prefixos foram abandonados. Entre primatólogos, o termo cultura se refere a padrões comportamentais que dependem de um contexto social para se desenvolver, e que podem atravessar gerações. Eu sugiro uma estratégia analítica que distingue os motivos de discordância entre descrições, explicações, teorias e visões de mundo, e argumento que a controvérsia é complexa e inclui discordâncias entre visões de mundo sem, no entanto, dividir os envolvidos em grupos homogêneos (digamos, primatólogos contra antropólogos). Por conta disso, a redefinição e o uso que os primatólogos fazem do termo acabam por manter ilesos os fundamentos da dicotomia natureza/cultura, o que pode explicar, parcialmente, a manutenção da controvérsia. Concluo que o diálogo entre os dois lados da fronteira será imprescindível para os pesquisadores que estiverem interessados em buscar uma abordagem consensual. É possível alcançar um consenso, mas a busca por uma abordagem sintética do comportamento animal que inclua os humanos deverá levar ao abandono ou reconstrução das dualidades natureza/cultura, inato/adquirido e gene/ambiente, e também da atribuição de primazia causal aos genes. Além disso, é necessário discutir a fundo sobre como incluir a questão do simbolismo e do significado em uma perspectiva comparativa / The Western ontological distinction between nature and culture, and the idea that the human (cultural) condition makes us radically different from other animals, are evident in the historical separation between the natural and social sciences. In parallel to other critics of this dualist view, some animal behaviorists have been using the term culture in relation to nonhumans, starting a controversy that is still far from cooling down. In this study, I investigated the meaning of the term culture as used by anthropologists, and also its recent use by ethologists (limiting myself to primatology), in order to better understand the controversy and identify possible paths that might lead to a consensus. In Anthropology, the modern concept of culture developed between the 19th century and the 1950s. It came to be seen as an emergent phenomenon exclusive to human social life. It was dependent on our capacity to use symbols and corresponded to behavioral patterns and norms, artifacts, ideas, and values that individuals acquire in the process of socialization. But this conception of culture, and the dualist epistemology supporting it, have since been largely criticized and intensely debated. Although contemporary anthropologists do not share a common ground or framework, virtually all of them agree that human cultural behavior is fundamentally symbolic. Recent attribution of culture to nonhuman primates started with Japanese scholars who, from the 1950s onward, have followed closely the spread of novel behaviors in Macaca fuscata, which they described with expressions such as preculture, subculture and infrahuman culture. Since the 1960s, field studies on wild populations and experimental research on learning in a social context, have led to the establishment of Cultural Primatology, and the prefixes were abandoned. Among primatologists, the term culture refers to behavioral patterns that depend on the social context to develop and that might be recurrent through generations. I suggest that it might be analytically useful to distinguish the matters of a disagreement between descriptions, explanations, theories and worldviews, and argue that this controversy goes all the way up to the highest reason of disagreement (worldviews). Still, one cannot divide those involved in it into a few homogeneous groups (say, primatologists contra anthropologists). Primatologists redefinition and use of the term do not alter the foundations of the criticized nature/culture dichotomy, and that might at least partially explain the maintenance of the controversy. It is possible to reach a consensus, but the search for a synthetic framework for animal behavior that includes humans might lead to the abandonment or reconstruction of the related dichotomies of nature/culture, innate/acquired and gene/environment, as well as of the causal primacy attributed to genes. It is also necessary to discuss how to include symbols and meanings in a comparative perspective

Zoological institutions emphasise the importance of excelling in the areas of animal welfare, conservation, education, and research, not only to better the lives of the animals under their care, but to also influence the general population in the pursuit to conserve the natural world. As a result, zoo life is anything but simple. This research project monitored the lives of a captive group of chimpanzees over a two-and-a-half-year period, during which time we explored four research topics while assessing the development of a cognitive research programme and chimpanzee (Pan troglodytes) introductions in a zoo: welfare, cognition, public engagement with science, and animal management. The project’s use of touchscreen technology and on-exhibit research was the first of its kind for the Royal Zoological Society of Scotland’s Edinburgh Zoo. As a result, the researchers placed a great deal of importance not only on assessing the welfare of the chimpanzees throughout training and testing phases, but also assessing the public’s perception of cognitive research being conducted through an internationally broadcast documentary about the project. In the short duration of the project, these research naïve chimpanzees did not fully grasp the concept of video selection in our free-choice activity, but overall, the introduction of a cognitive research programme did not compromise welfare, and the chimpanzees’ repeated interest suggests that chimpanzees found the research to be reinforcing. Partly funded by the BBC, the Chimpcam Project was shown in the UK (broadcast January 2010) and in a variety of other countries, including the United States and Canada (on Animal Planet in 2011). The broadcast allowed us to gather information over the internet on the wider public’s perception of conducting research with great apes in zoos, to complement data collected on visitors to the exhibit itself. Our assessment of the documentary’s impact on public perception showed that it had a positive influence on perceptions of zoo research, scientists, welfare, and the importance of choice for animals. During this research project, a new group of chimpanzees arrived in Edinburgh as part of the international breeding programme for western chimpanzees (Pan troglodytes verus). As the zoo’s focus switched to helping the two chimpanzee groups merge into one, we took the opportunity to apply psychological research to this context, namely the use of video as a research tool and the recognition of the importance of individual differences in response to challenge. The project maintained the cognition and welfare focus by using video introductions (allowing the chimpanzees to watch video footage of the individuals they were about to meet and track the formation of other sub-groups). In addition, personality ratings and chimpanzee behaviour during the visual access period (an animal management technique used prior to physical introductions where the groups could see each other without physical contact) were collected to examine the efficacy of these measures in guiding introductions in order to reduce risk. Personality ratings and behaviours observed during the video introductions could predict the chimpanzees’ behaviour during the physical introductions, however, the visual access period had no predictive power. The welfare implications of the introduction process were also assessed and suggested that: the choice of location (i.e. options of where to be) was more important than the total amount of available space; having individuals removed from your group was more stressful than having individuals added; self-directed behaviour (SDB) performance was context-specific where rubbing significantly increased during periods of uncertainty that were not necessarily negatively valenced; regurgitation and reingestion (R/R) decreased over time; and both in-group members and those of high ranks spent more time grooming others. Overall our data indicate that the chimpanzees coped well with both cognitive challenges and social upheaval during introductions. Despite being regularly studied in captivity and in the wild, chimpanzees have a great deal more to teach us about their world. In order to provide the best welfare for the chimpanzees in our care, we need to understand how research and management practices affect their lives and how the public interpret what we do as researchers. By understanding these aspects of their world, we can better serve those in captivity and influence public opinion on the importance of conserving those in the wild.

xvii, 149 p. : ill. (some col.), maps / The ranging behavior of an animal can reveal much about the social and ecological conditions it faces. Food availability, feeding competition, population pressures, metabolic requirements and human influences can all influence the ranging behavior of individuals. For modern humans, the manner in which we move about our world is limited only by access to technology and other cultural factors. Of course, it has not always been that way. Based on recent fossil discoveries, our earliest bipedal ancestors more closely resembled the living great apes in morphology. Consequently, studies of great ape behavior have been used to reconstruct scenarios of early hominin behavior. And while much has been written about chimpanzee ( Pan troglodytes ) ranging in this regard, less is known about bonobos (Pan paniscus ). Along with chimpanzees, bonobos are our closest phylogenetic relative, existing today as a descendant of a common ancestor the Homo and Pan genera shared sometime around six million years ago. Despite their close taxonomic relationship, however, there are a variety of behavioral differences between bonobos and chimpanzees. The aim of this dissertation is to better understand these differences within the context of ranging and social behavior and apply the results to models of early hominin behavior. More specifically, I used a Geographic Information Systems (GIS) approach to examine general bonobo ranging data, the differences in ranging behavior between males and females, and the manner in which neighboring groups interact. Compared with chimpanzees, bonobos at Lomako range over a much smaller total area, are not territorial, and differ in the composition of social parties. In general, female bonobos are more gregarious and cohesive, moving in semi-stable groups I call "cliques", while males are less aggressive and more likely to move independently. These results likely reflect the high levels of food availability for bonobos at Lomako, reducing the level of feeding competition, and emphasizing social and mating strategies in group formation. Consequently, the spectrum of potential early hominin ranging behavior must be expanded from the current chimp-centric perspective. / Committee in charge: Frances White, Chairperson; Stephen Frost, Member; Larry Sugiyama, Member; James Schombert, Outside Member

Les cinq études détaillées au cours de cette thèse interrogent divers aspects de la théorie de l’esprit, de la conscience de soi ainsi que du lien (multiple ou unique) qui unit ces deux capacités sociocognitives si particulières. Les deux premières études mettent en évidence différentes variables qui peuvent – ou non- influencer l’attribution d’intention et de fausse croyance à autrui. La troisième étude porte plus précisément sur les différents composants de la conscience de soi et sur leur trajectoire développementale. La quatrième étude interroge le lien développemental entre la théorie de l’esprit et la conscience de soi, alors que la dernière étude questionne la présence de ce lien chez les chimpanzés (Pan Troglodytes). Nous démontrerons, tout au long de cette thèse, que la théorie de l’esprit et la conscience de soi sont composées de différents éléments et que leur acquisition est graduelle. / Doctorat en Sciences Psychologiques et de l'éducation / info:eu-repo/semantics/nonPublished