付記する学位プログラム名: 霊長類学・ワイルドライフサイエンス・リーディング大学院 / 京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第22721号 / 理博第4630号 / 新制||理||1665(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)准教授 足立 幾磨, 准教授 後藤 幸織, 教授 高田 昌彦 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

付記する学位プログラム名: 霊長類学・ワイルドライフサイエンス・リーディング大学院 / 京都大学 / 新制・課程博士 / 博士(理学) / 甲第23054号 / 理博第4731号 / 新制||理||1678(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)准教授 足立 幾磨, 准教授 後藤 幸織, 教授 高田 昌彦 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DFAM

京都大学 / 新制・課程博士 / 博士(理学) / 甲第24183号 / 理博第4874号 / 新制||理||1697(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)准教授 足立 幾磨, 准教授 Huffman Michael Alan, 教授 今井 啓雄 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DFAM

Decision making is a framework we impose on a vast universe of possible behaviors to better understand the advantages and disadvantages of acting in different ways. Decisions under uncertainty are of particular interest because stochasticity is a feature of environments both today and throughout evolutionary history. As a result, we might expect decision makers (DMs) to have evolved mechanisms to handle variability. In this dissertation, I examine common decision processes in several model species: European starlings (Sturnus vulgaris), Banded tetras (Astyanax fasciatus) and humans (Homo sapiens). The broad range of approaches discussed include currencies DMs are expected to maximize (Risk Sensitivity Theory, Expected Utility), the currencies DMs do maximize (e.g. long versus short term rate maximizing models), the representation of outcomes in memory (Scalar Expectancy Theory) as well as explicit choice mechanisms (Sequential Choice Model). The first section of this thesis discusses responses to risk, offering humans and starlings choices between options which deliver certain or variable outcomes. Starlings demonstrate sensitivity to changes in the probability of variable outcomes and strong support for local rate maximization. Humans appear similarly sensitive to their own accuracy when task difficulty is varied. When the DM’s affective state was manipulated, neither humans nor starlings exhibit changes in risk preferences, but the effectiveness of these manipulations used could not be confirmed. Another topic of inquiry is the effect of the DM’s state at the time of valuation learning. State dependent valuation learning is demonstrated for the first time in a fish species, but Within Trial Contrast is not observed in starlings. Lastly, two experiments find strong support for the Sequential Choice Model, a promising new model of the mechanism of choice. Taken together, these experiments offer a glimpse into shared decision processes, but leave open questions about the mechanisms through which value is acquired.

This dissertation explored the social cognitive abilities of the Clark’s nutcracker (Nucifraga columbiana), a relatively non-social, food-caching corvid. Corvids are a family of large-brained birds, which are capable of remarkable cognitive feats (e.g., future planning, tool use). These cognitive abilities have been revealed predominantly by testing social species, supporting popular theories that living in social groups drove the evolution of complex cognition. However, few studies have investigated the social cognitive abilities of corvid species that do not live in large groups. Here, I developed novel procedures using the food-caching behaviour of Clark’s nutcrackers as a tool to explore two cognitive abilities predicted to be limited to social species: mirror self-recognition (Chapter 2) and cooperation (Chapter 4). In Chapter 2, birds cached food when alone, with a conspecific present, and with a regular or blurry mirror. The nutcrackers suppressed caching with a regular mirror (as done with a conspecific), but not with the blurry mirror. When integrated with the traditional ‘mark test’, the birds also showed evidence of self-recognition with the blurry mirror by attempting to remove a coloured mark placed on their body with the blurry mirror, but not with an opaque barrier. In Chapter 3, I discuss the importance of self-recognition as a precursor for complex and flexible social cognitive abilities such as cooperation. To investigate cooperation, in Chapter 4 the birds experienced having their caches exchanged with another bird over multiple trials. This procedure assessed whether the normal response of cache suppression with a conspecific could be over-ridden if the experimental contingencies made cache sharing beneficial. The nutcrackers continued to cache in this context, and male birds increased caching when cooperation from the conspecific was exaggerated artificially by the experimenter. Combined, the results indicate the non- social Clark’s nutcracker is capable of mirror self-recognition, and the ability to distinguish one’s ‘self’ from others may facilitate flexible caching decisions, contrary to the predictions of the social living hypotheses. The findings indicate social living alone does not strongly predict complex cognitive abilities and, instead, that multiple evolutionary paths exist for the development of complex cognition. / October 2016

No description available.

This study examined object permanence in Sumatran orangutans (Pongo abelii), Western lowland gorillas (Gorilla gorilla gorilla), and black-and-white-ruffed lemurs (Varecia variegata) at Zoo Atlanta. A literature review reveals two main issues with object permanence research in non-human primates. One of the issues is that it is difficult to make valid comparisons between different species because very few studies have been conducted using appropriate controls. Thus, one of the goals of this study was to conduct control trials for all tasks in the traditional object permanence test battery, in order to reliably assess and compare performance in the species under study. The second issue is concerned with the finding that all of the non-human primate species tested so far have failed one of the more difficult tasks in the test battery, namely the non-adjacent double invisible displacement task. It has been hypothesized that this performance limitation is a result of the manner in which the task is presented. Thus, the second goal of this study was to modify the existing methodology and present the task to gorillas and orangutans in locomotive space to see if performance improves. This is the first study to present this task to non-human primate species in locomotive space. This study found that orangutans were the only species to reliably pass most tasks in the traditional object permanence test battery. Black-and-white ruffed lemurs failed most visible and invisible displacement tasks. Owing to the small sample size of gorillas in this study, further research is required before any firm conclusions can be made about the ability of this species to solve visible and invisible displacement tasks in the traditional object permanence test battery. Presenting the boxes in locomotive space to gorillas and orangutans did not improve performance on the non-adjacent double invisible displacement task. Further research is required to resolve the question of whether this performance limitation is a result of the manner in which the task was presented.

In nature, pollinators must navigate fields of resources presenting a variety of features, differing in shape, size, colour, etc. Foraging on a flower by flower basis is slow and maladaptive. Instead foragers must be able to differentiate between rewarding and unrewarding floral species while also generalizing learned information between flowers of the same species. The ability to categorize stimuli occurs on several levels of abstraction, laid out by Herrnstein (1990). In order to categorize objects, animals must first be able differentiate between them without memorizing each stimulus separately. Consequently, objects can be grouped by physical characteristics through perceptual categorization, or, on a more abstract scale, by the function they serve to the animal. This thesis explores the bees’ ability to categorize flowers, following the levels of categorization to answer two questions: Can bees form categories? And how abstract can their categorization become? There has been limited investigation previously in categorization in bees, and no research done on invertebrates addresses categorization beyond physical features. The bees’ ability to form categories was evaluated with four experiments of preference: 1) similar-but-different judgements; 2) perceptual categorization; 3) simple and mediated generalization; and 4) functional relevance. Results show firstly that bees can generalize characteristics within flower species, but also differentiate individual flowers. Secondly, bees can form perceptual categories, and while they rely on physical floral features for categorization, bees also demonstrated preliminary abilities for functional generalization as well. These results provide an explanation to natural foraging techniques adopted by the bees. The adaptive nature of categorization allows foragers to find resources more efficiently and better prepare in changing environments.

This diploma thesis is focused on research of individual recognition in African grey parrots based on visual cues from stimulus card made from photographs of familiar conspecifics, specifically on testing the significance of selected visual features. Theoretical part of the thesis deals with the general individual recognition ability followed by specifics of this ability in humans, non-human primates and birds including African grey parrot. Great attention is paid specifically to the individual recognition ability in humans, since it is the aim of this thesis to interpret the results of hypotheses testing comparatively. In order to test the hypotheses seven modifications of conspecifics' photographs were created. Experiment employed matching-to- sample as method, subjects were three African grey parrots. Findings were to a large extent in accordance with findings of analogical studies that used human subjects. Results imply, that visual information in African grey parrot is processed holistically, with structure and pigmentation of feathers on the abdominal part of the parrot having most likely the highest significance for successful individual recognition of a familiar conspecific. Key words: individual recognition, African grey parrot, comparative cognition, visual recognition

Cumulative culture is rare, if not altogether absent in non-human species. At the foundation of cumulative learning is the ability to flexibly modify, relinquish or build upon prior behaviours to make them more productive or efficient. Within the primate literature, a failure to optimise solutions in this way is often proposed to derive from low-fidelity copying of witnessed behaviours, sub-optimal social learning heuristics, or a lack of relevant socio-cognitive adaptations. However, humans can also be markedly inflexible in their behaviours, perseverating with, or becoming fixated on outdated or inappropriate responses. Humans show differential patterns of flexibility as a function of cognitive load, exhibiting difficulties with inhibiting suboptimal behaviours when there are high demands on working memory. Here I present a series of studies on captive chimpanzees which show that not only is inhibitory control compromised in chimpanzees, but indicate ape behavioural conservatism may be underlain by similar constraints as in humans; chimpanzees show relatively little conservatism when behavioural optimisation involves the inhibition of a well-established but simple solution, or the addition of a simple modification to a well-established but complex solution. In contrast, when behavioural optimisation involves the inhibition of a well-established but complex solution, and especially when the alternative solution is also complex, chimpanzees show evidence of behavioural conservatism. I propose that conservatism is linked to behavioural complexity, potentially mediated by cognitive resource availability, and may be an important factor in the evolution of cumulative culture.