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Byrne, R. W., & Russon, A. E. (1998). Learning by imitation: a hierachical approach. Behav. Brain Sci., 21, 667–721.
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Byrne, R. W. (1999). Imitation without intentionality. Using string parsing to copy the organization of behaviour. Anim. Cogn., 2(2), 63–72.
Abstract: A theory of imitation is proposed, string parsing, which separates the copying of behavioural organization by observation from an understanding of the cause of its effectiveness. In string parsing, recurring patterns in the visible stream of behaviour are detected and used to build a statistical sketch of the underlying hierarchical structure. This statistical sketch may in turn aid the subsequent comprehension of cause and effect. Three cases of social learning of relatively complex skills are examined, as potential cases of imitation by string parsing. Understanding the basic requirements for successful string parsing helps to resolve the conflict between mainly negative reports of imitation in experiments and more positive evidence from natural conditions. Since string parsing does not depend on comprehension of the intentions of other agents or the everyday physics of objects, separate tests of these abilities are needed even in animals shown to learn by imitation.
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Byrne, R. W. (2002). Imitation of novel complex actions: What does the evidence from animals mean? In C. T. Snowdon, T. J. Roper, & J. S. Rosenblatt (Eds.), Advances in the Study of Behavior (Vol. 31, pp. 77–105). San Diego: Academic Press.
Abstract: Summary Underlying the various behaviors that are classified as imitation, there may be several distinct mechanisms, differing in adaptive function, cognitive basis, and computational power. Experiments reporting “true motor imitation” in animals do not as yet give evidence of production learning by imitation; instead, contextual imitation can explain their data, and this can be explained by a simple mechanism (response facilitation) which matches known neural findings. When imitation serves a function in social mimicry, which applies to a wide range of phenomena from neonatal imitation in humans and great apes to pair-bonding in some bird species, the fidelity of the behavioral match is crucial. Learning of novel behavior can potentially be achieved by matching the outcome of a model's action, and it is argued that vocal imitation by birds is a clear example of this method (which is sometimes called emulation). Alternatively, the behavior itself may be perceived in terms of actions that the observer can perform, and thus it may be copied. If the imitation is linear and stringlike (action level), following the surface form rather than the underlying plan, then its utility for learning new instrumental methods is limited. However, the underlying plan of hierarchically organized behavior is visible in output behavior, in subtle but detectable ways, and imitation could instead be based on this organization (program level), extracted automatically by string parsing. Currently, the most likely candidates for such capacities are all great apes. It is argued that this ability to perceive the underlying plan of action, in addition to allowing highly flexible imitation of novel instrumental methods, may have resulted in the competence to understand the intentions (theory of mind) of others.
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Byrne, R. W. (2000). How monkeys find their way: leadership, coordination, and cognitive maps of African baboons. In S. Boinski, & P. A. Garber (Eds.), On the Move: How and Why Animals Travel in Groups (pp. 491–518). Chicago: Chicago University Press.
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Ruiz, A., Gómez, J., Roeder, J., & Byrne, R. (2009). Gaze following and gaze priming in lemurs. Anim. Cogn., 12(3), 427–434.
Abstract: Abstract  Although primates have often been found to co-orient visually with other individuals, members of these same species have usually failed to use co-orientation to find hidden food in object-choice experiments. This presents an evolutionary puzzle: what is the function of co-orientation if it is not used for a function as basic as locating resources? Co-orientation responses have not been systematically investigated in object-choice experiments, and requiring co-orientation with humans (as is typical in object-choice tasks) may underestimate other species’ abilities. Using an object-choice task with conspecific models depicted in photographs, we provide experimental evidence that two lemur species (Eulemur fulvus, n = 4, and Eulemur macaco, n = 2) co-orient with conspecifics. Secondly, by analysing together two measures that have traditionally been examined separately, we show that lemurs’ gaze following behaviour and ultimate choice are closely linked. Individuals were more likely to choose correctly after having looked in the same direction as the model, and thus chose objects correctly more often than chance. We propose a candidate system for the evolutionary origins of more complex gaze following: ‘gaze priming.’
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Held, S., Baumgartner, J., Kilbride, A., Byrne, R. W., & Mendl, M. (2005). Foraging behaviour in domestic pigs (Sus scrofa): remembering and prioritizing food sites of different value. Anim. Cogn., 8(2), 114–121.
Abstract: This experiment investigated whether domestic pigs can remember the locations of food sites of different relative value, and how a restricted retrieval choice affects their foraging behaviour. Nine juvenile female pigs were trained to relocate two food sites out of a possible eight in a spatial memory task. The two baited sites contained different amounts of food and an obstacle was added to the smaller amount to increase handling time. On each trial, a pig searched for the two baited sites (search visit). Once it had found and eaten the bait, it returned for a second (relocation) visit, in which the two same sites were baited. Baited sites were changed between trials. All subjects learnt the task. When allowed to retrieve both baits, the subjects showed no preference for retrieving a particular one first (experiment 1). When they were allowed to retrieve only one bait, a significant overall preference for retrieving the larger amount emerged across subjects (experiment 2). To test whether this preference reflected an avoidance of the obstacle with the smaller bait, 15 choice-restricted control trials were conducted. In control trials obstacles were present with both baits. Pigs continued to retrieve the larger bait, indicating they had discriminated between the two food sites on the basis of quantity or profitability and adjusted their behaviour accordingly when the relocation choice was restricted. This suggests for the first time that domestic pigs have the ability to discriminate between food sites of different relative value and to remember their respective locations.
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Cochet, H., & Byrne, R. W. (2013). Evolutionary origins of human handedness: evaluating contrasting hypotheses. Animal Cognition, 16(4), 531–542.
Abstract: Variation in methods and measures, resulting in past dispute over the existence of population handedness in nonhuman great apes, has impeded progress into the origins of human right-handedness and how it relates to the human hallmark of language. Pooling evidence from behavioral studies, neuroimaging and neuroanatomy, we evaluate data on manual and cerebral laterality in humans and other apes engaged in a range of manipulative tasks and in gestural communication. A simplistic human/animal partition is no longer tenable, and we review four (nonexclusive) possible drivers for the origin of population-level right-handedness: skilled manipulative activity, as in tool use; communicative gestures; organizational complexity of action, in particular hierarchical structure; and the role of intentionality in goal-directed action. Fully testing these hypotheses will require developmental and evolutionary evidence as well as modern neuroimaging data.
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Barton, R. A., Byrne, R. W., & Whiten, A. (1996). Ecology, feeding competition and social structure in baboons. Behav. Ecol. Sociobiol., 38(5), 321–329.
Abstract: Predictions of the model of van Schaik (1989) of female-bonding in primates are tested by systematically comparing the ecology, level of within-group contest competition for food (WGC), and patterns of social behaviour found in two contrasting baboon populations. Significant differences were found in food distribution (percentage of the diet from clumped sources), feeding supplant rates and grooming patterns. In accord with the model, the tendencies of females to affiliate and form coalitions with one another, and to be philopatric, were strongest where ecological conditions promoted WGC. Group fission in the population with strong WGC was “horizontal” with respect to female dominance rank, and associated with female-female aggression during a period of elevated feeding competition. In contrast, where WGC was low, females' grooming was focused on adult males rather than other females. Recent evidence suggests that group fission here is initiated by males, tends to result in the formation of one-male groups, and is not related to feeding competition but to male-male competition for mates. An ecological model of baboon social structure is presented which incorporates the effects of female-female competition, male-male competition, and predation pressure. The model potentially accounts for wide variability in group size, group structure and social relationships within the genus Papio. Socio-ecological convergence between common baboons and hamadryas baboons, however, may be limited in some respects by phylogenetic inertia.
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Riley, J. L., Noble, D. W. A., Byrne, R. W., & Whiting, M. J. (2017). Does social environment influence learning ability in a family-living lizard? Anim. Cogn., 20(3), 449–458.
Abstract: Early developmental environment can have profound effects on individual physiology, behaviour, and learning. In birds and mammals, social isolation during development is known to negatively affect learning ability; yet in other taxa, like reptiles, the effect of social isolation during development on learning ability is unknown. We investigated how social environment affects learning ability in the family-living tree skink (Egernia striolata). We hypothesized that early social environment shapes cognitive development in skinks and predicted that skinks raised in social isolation would have reduced learning ability compared to skinks raised socially. Offspring were separated at birth into two rearing treatments: (1) raised alone or (2) in a pair. After 1 year, we quantified spatial learning ability of skinks in these rearing treatments (N = 14 solitary, 14 social). We found no effect of rearing treatment on learning ability. The number of skinks to successfully learn the task, the number of trials taken to learn the task, the latency to perform the task, and the number of errors in each trial did not differ between isolated and socially reared skinks. Our results were unexpected, yet the facultative nature of this species' social system may result in a reduced effect of social isolation on behaviour when compared to species with obligate sociality. Overall, our findings do not provide evidence that social environment affects development of spatial learning ability in this family-living lizard.
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Byrne, R. W. (1993). Do larger brains mean greater intelligence? Behav. Brain Sci., 16(4), 696–697.
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