Elephants' complex societies, well-developed communication systems, evolutionary history and close working relationship with humans make them an important species for studies of cognition but research on elephant cognition is sparse. In this thesis I aim to illuminate the social cognition involved in the interpretation and production of visual signals by African elephants (Loxodonta africana). My results are intended to contribute to the cross-species literature on social cognition and help to elucidate wild elephant social behaviour. I studied captive elephants, housed at an elephant-back safari company in Victoria Falls, and wild elephants in Hwange National Park, both in Zimbabwe. Wild elephants display a vast array of postures, actions and signals. I found that elephants recognise visual attentiveness in others when they signal silently, producing more signals when their audience can see them, and using the body and face orientation of an audience to judge their attention. When responding to typically human visual signals, elephants immediately responded correctly to deictic gestures, including variants of pointing that they were unlikely to have already experienced. These results indicate elephants' astonishing sensitivity to even subtle social cues. I found no indication that elephants reason about mental states such as false beliefs, or rationality; however, limitations of the experimental design meant I was unlikely to find such an ability even if it is present in elephants. Furthermore, I discovered that elephants have a form of referential indication in their natural communication in the wild. Elephants match their direction of attention with a type of trunk action produced by a group member. Attending to human-like signals, and interpreting them as communicative is an advantage for any animal working with humans and that ability might explain the choice of species that are ancestors of today's domestic animals.

Thesis (M.A.)--Georgia State University, 2007. / Title from file title page. David A. Washburn, committee chair; Eric Vanman, Michael J. Beran, Heather Kleider, committee members. Electronic text (76 p. : col. ill.) : digital, PDF file. Description based on contents viewed Mar. 25, 2008. Includes bibliographical references (p. 71-76).

Thesis (Ph. D.)--Ludwig-Maximilians-Universität München, 2006. / Title from PDF title page (viewed on June 4, 2006). Includes bibliographical references.

A full understanding of any biological trait requires investigation of its evolutionary origin. Primates inspire great curiosity amongst researchers due to the remarkable diversity across species in both anatomical and behavioural traits, including sociality, sexual behaviour, life histories, neuro-anatomy, cognitive abilities and behavioural repertoires. The study of primates has involved comparative approaches since its inception, however, the necessary tools for statistically investigating the macro-evolutionary processes responsible for current diversity in biological traits have been developed only in the last 30 years or so, namely phylogenetic reconstruction and phylogenetic comparative methods. Amongst a multitude of evolutionary questions that can be addressed by phylogenetic comparative analyses, this thesis attempts to address two in particular, concerning primates. First, chapters 3 and 4 use meta-analysis and phylogenetic comparative analyses to investigate the evolution of large, brightly coloured ‘exaggerated sexual swellings' in female Catarrhine (‘Old World') primates. Together, chapters 3 and 4 show that such swellings are signals of temporal fertility, and present evidence to suggest that swellings co-evolved with conditions favouring male mate choice and cryptic female choice, therefore shedding light on the general conditions under which female signals of temporal fertility should evolve. Second, chapters 5 and 6 use phylogenetic comparative analyses investigate the evolution of enlarged brain size in the primate order. Together, chapters 5 and 6 suggest that multiple selection pressures have contributed to diversity in brain size and cognitive traits across primates, including sociality, intra-sexual competition and extended life history. Further, analyses presented in chapter 6 suggest that reliance on learned behaviour is a self-reinforcing evolutionary process, favouring ‘runaway' increases in cognitive abilities and reliance on culture in some primate lineages, which parallels increases in brain size, cognitive ability and reliance on culture in human evolution.

By investigating the cognitive capacities of non-human primates, we can begin to understand the cognitive capacities of the evolutionary ancestors we share with these species. While there is a great deal of research exploring the socio-cognitive abilities of simian primates, prosimians have not been sufficiently studied. Without data from these species, our knowledge about the evolution of the primate mind is limited to the common ancestor shared between simian primates only, precluding understanding of the phylogenetic origins of certain phenomena. I explored the socio-cognitive capacities of lemurs, a type of prosimian primate. I studied several areas of social cognition related to social referencing, defined as the ability to use and seek out social information when appraising objects or events. As social referencing is a popular subject in both human developmental and non-human primate literature, I aimed to determine how prosimians’ capacities compare. My research was conducted with captive lemurs of three species: Eulemur fulvus fulvus, Eulemur macaco macaco, and Eulemur fulvus rufus. I found that lemurs use social cues regarding food palatability to modify their own feeding behaviour and that they visually attend to conspecifics differently when presented with novel, as compared to familiar, foods. Lemurs also visually referred to a human experimenter’s face when presented with an anomalous interaction and went on to engage in gaze alternation. Lemurs failed to use information about the experimenter’s attentional state, however, when modifying their use of a trained gesture. Finally, I found that lemurs are able to visually co-orient with conspecifics, correctly prioritising information from the head over that from the body, and that they go on to use conspecific gaze to locate hidden resources. These results show that lemurs are more cognitively advanced than previously thought and the origins of some social referencing skills may be phylogenetically older than previously hypothesised.

The ability to learn spatiotemporal characteristics of biologically significant events is advantageous for an animal and is known as time-place learning (TPL). Gallistel (1990) proposed an influential theory positing that whenever a biologically significant event occurred, a memory code was automatically formed, encoding the nature of the event, and the time and place in which it occurred. When the animal is later faced with a biological need it could consult these memory codes and determine when and where that need had been met in the past. This information could be used to guide current behaviour. Importantly, Gallistel theorized that the encoding of the spatiotemporal characteristics of an event into a tripartite code was an automatic process. Despite the appealing power and simplicity of Gallistel's theory, I have provided arguments suggesting that it has serious limitations. Perhaps the most damaging evidence against this theory is the reluctance of rats to demonstrate daily TPL (i.e., events that vary in location depending on time of day). Widman, Gordon, and Timberlake (2000) argue that for TPL to occur the response cost for incorrect decisions must be high. While this hypothesis is unable to explain the inconsistencies in TPL, it does highlight the fact that animals do not automatically store time-place-event information as a tripartite code. If they did, it would not make sense for them to ignore such information in some tasks. I have provided an alternative hypothesis that states that whenever a biologically significant event occurs two bipartite memory codes (time-event and place-event) are automatically formed. Only under some conditions, perhaps those with high response cost, do animals form tripartite codes. For this reason, rats often have difficulty learning a TPL task; although rats easily learn a place preference for those places that provide reinforcement (place-event), and easily learn a go/no-go discrimination (time-event). This thesis provides data from both the daily and interval TPL realms supporting the proposed theory of bipartite codes. Although rats do not readily learn daily TPL tasks, they do demonstrate knowledge of interval TPL under a variety of conditions designed to enhance the ecological validity of the task. The properties of interval TPL are discussed. / Arts, Faculty of / Psychology, Department of / Graduate

While most human language is expressed verbally, the gestures produced concurrent to speech provide additional information, help listeners interpret meaning, and provide insight into the cognitive processes of the speaker. Several theories have suggested that gesture played an important, possibly central, role in the evolution of language. Great apes have been shown to use gestures flexibly in different situations and to modify their gestures in response to changing contexts. However, it has not previously been determined whether ape gestures are defined by structural variables, carry meaning, are used to intentionally communicate specific information to others, or can be used strategically to overcome miscommunication. To investigate these questions, I studied three captive populations of orangutans (Pongo pygmaeus and P. abelii) in European zoos for 10 months. Sixty-four different gestures, defined through similarities in structure and use, were included in the study after meeting strict criteria for intentional usage. More than half of the gesture types were found to coincide frequently with specific goals of signallers, and were accordingly identified as having meanings. Both structural and social variables were found to determine gesture meaning. The recipient’s gaze in both the present and the past, and the recipient’s apparent understanding of the signaller’s gestures, affected the strategies orangutans employed in their attempts to communicate when confronted with different types of communicative failure (e.g. not seeing, ignoring, misunderstanding, or rejecting a gesture). Maternal influence affected the object-directed behaviour and gestures of infants, who shared more gestures with their mothers than with other females. These findings demonstrate that gesture can be used as a medium to investigate not only the communication but also the cognition of great apes, and indicate that orangutans are more sensitive to the perceptions and knowledge states of others than previously thought.

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 Cognitive Evaluation Theory explains the outcomes of studies employing Deci's paradigm, but only when used post hoc. A basic assumption is that extrinsic rewards always increase intrinsic motivation for nonhuman subjects. Deci's paradigm was modified for use with 22 rats to test this assumption. Running in an exercise wheel was the intrinsically motivated activity studied. ANCOVA revealed that external rewards increased intrinsic interest on the first day following the cessation of reinforcement (F = 8.32), but on two subsequent days and again a week later, no significant differences between the reward and control groups were evident (F = .29; F = .33; F = 3.70). The assumption was not supported. It was demonstrated that repeated posttest measures are necessary to avoid basing conclusions upon one point along the extinction continuum.

Understanding intentionality, i.e. coding the object directedness of agents towards objects, is a fundamental component of Theory of Mind abilities. Yet it is unclear how it is perceived and coded in different species. In this thesis, we present a series of comparative studies to explore human adults' and Capuchin monkeys' ability to infer intentional objects from actions. First we studied whether capuchin monkeys and adult humans infer a potential object from observing an object-directed action. With no direct information about the goal-object, neither species inferred the object from the action. However, when the object was revealed, the monkeys retrospectively encoded the directedness of the object-directed action; unexpectedly, in an adapted version of the task adult humans did not show a similar ability. We then adapted another paradigm, originally designed by Kovács et al (2010), to examine whether the two species implicitly register the intentional relation between an agent and an object. We manipulated an animated agent and the participants' belief about a ball's presence behind a hiding screen. We found no evidence showing that humans or monkeys coded object-directedness or belief. More importantly, we failed to replicate the original results from Kovács et al's study, and through a series of follow up studies, we questioned their conclusions regarding implicit ToM understanding. We suggested that, instead of implicit ToM, results like Kovacs et al's might be interpreted as driven by “sub-mentalizing” processes, as suggested by Heyes (2014). We conclude that so called ‘implicit ToM' may be based upon the computation of intentional relations between perceived agents and objects. But, these computations might present limitations, and some results attributed to implicit ToM may in fact reflect “sub-mentalizing” processes.