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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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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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Topál, J., Byrne, R. W., Miklósi, Á., & Csányi, V. (2006). Reproducing human actions and action sequences: “Do as I Do!” in a dog. Anim. Cogn., 9(4), 355–367.
Abstract: We present evidence that a dog (Philip, a 4-year-old tervueren) was able to use different human actions as samples against which to match his own behaviour. First, Philip was trained to repeat nine human-demonstrated actions on command ('Do it!'). When his performance was markedly over chance in response to demonstration by one person, testing with untrained action sequences and other demonstrators showed some ability to generalise his understanding of copying. In a second study, we presented Philip with a sequence of human actions, again using the 'Do as I do' paradigm. All demonstrated actions had basically the same structure: the owner picked up a bottle from one of six places; transferred it to one of the five other places and then commanded the dog ('Do it!'). We found that Philip duplicated the entire sequence of moving a specific object from one particular place to another more often than expected by chance. Although results point to significant limitations in his imitative abilities, it seems that the dog could have recognized the action sequence, on the basis of observation alone, in terms of the initial state, the means, and the goal. This suggests that dogs might acquire abilities by observation that enhance their success in complex socio-behavioural situations.
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Janson, C., & Byrne, R. (2007). What wild primates know about resources: opening up the black box. Anim. Cogn., 10(3), 357–367.
Abstract: Abstract We present the theoretical and practical difficulties of inferring the cognitive processes involved in spatial movement decisions of primates and other animals based on studies of their foraging behavior in the wild. Because the possible cognitive processes involved in foraging are not known a priori for a given species, some observed spatial movements could be consistent with a large number of processes ranging from simple undirected search processes to strategic goal-oriented travel. Two basic approaches can help to reveal the cognitive processes: (1) experiments designed to test specific mechanisms; (2) comparison of observed movements with predicted ones based on models of hypothesized foraging modes (ideally, quantitative ones). We describe how these two approaches have been applied to evidence for spatial knowledge of resources in primates, and for various hypothesized goals of spatial decisions in primates, reviewing what is now established. We conclude with a synthesis emphasizing what kinds of spatial movement data on unmanipulated primate populations in the wild are most useful in deciphering goal-oriented processes from random processes. Basic to all of these is an estimate of the animals ability to detect resources during search. Given knowledge of the animals detection ability, there are several observable patterns of resource use incompatible with a pure search process. These patterns include increasing movement speed when approaching versus leaving a resource, increasingly directed movement toward more valuable resources, and directed travel to distant resources from many starting locations. Thus, it should be possible to assess and compare spatial cognition across a variety of primate species and thus trace its ecological and evolutionary correlates.
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Bates, L. A., Sayialel, K. N., Njiraini, N. W., Poole, J. H., Moss, C. J., & Byrne, R. W. (2008). African elephants have expectations about the locations of out-of-sight family members. Biol Lett, 4(1), 34–36.
Abstract: Monitoring the location of conspecifics may be important to social mammals. Here, we use an expectancy-violation paradigm to test the ability of African elephants (Loxodonta africana) to keep track of their social companions from olfactory cues. We presented elephants with samples of earth mixed with urine from female conspecifics that were either kin or unrelated to them, and either unexpected or highly predictable at that location. From behavioural measurements of the elephants' reactions, we show that African elephants can recognize up to 17 females and possibly up to 30 family members from cues present in the urine-earth mix, and that they keep track of the location of these individuals in relation to themselves.
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Byrne, R. W., & Bates, L. A. (2006). Why are animals cognitive? Curr Biol, 16(12), R445–8.
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Byrne, R. W. (2009). Animal imitation. Current Biology, 19(3), R111–R114.
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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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Farmer, K., Krueger, K., & Byrne, R. (2010). Visual laterality in the domestic horse (Equus caballus) interacting with humans. Anim. Cogn., 13, 229–238.
Abstract: Most horses have a side on which they are easier to handle and a direction they favour when working on a circle, and recent studies have suggested a correlation between emotion and visual laterality when horses observe inanimate objects. As such lateralisation could provide important clues regarding the horse’s cognitive processes, we investigated whether horses also show laterality in association with people. We gave horses the choice of entering a chute to left or right, with and without the passive, non-interactive presence of a person unknown to them. The left eye was preferred for scanning under both conditions, but significantly more so when a person was present. Traditionally, riders handle horses only from the left, so we repeated the experiment with horses specifically trained on both sides. Again, there was a consistent preference for left eye scanning in the presence of a person, whether known to the horses or not. We also examined horses interacting with a person, using both traditionally and bilaterally trained horses. Both groups showed left eye preference for viewing the person, regardless of training and test procedure. For those horses tested under both passive and interactive conditions, the left eye was preferred significantly more during interaction. We suggest that most horses prefer to use their left eye for assessment and evaluation, and that there is an emotional aspect to the choice which may be positive or negative, depending on the circumstances. We believe these results have important practical implications and that emotional laterality should be taken into account in training methods.
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Bates, L. A., Lee, P. C., Njiraini, N., Poole, J. H., Sayialel, K., Sayialel, S., et al. (2008). Do Elephants Show Empathy? J Conscious Stud, 15(10-11), 204–225.
Abstract: Elephants show a rich social organization and display a number of unusual traits. In this paper, we analyse reports collected over a thirty-five year period, describing behaviour that has the potential to reveal signs of empathic understanding. These include coalition formation, the offering of protection and comfort to others, retrieving and 'babysitting' calves, aiding individuals that would otherwise have difficulty in moving, and removing foreign objects attached to others. These records demonstrate that an elephant is capable of diagnosing animacy and goal directedness, and is able to understand the physical competence, emotional state and intentions of others, when they differ from its own. We argue that an empathic understanding of others is the simplest explanation of these abilities, and discuss reasons why elephants appear to show empathy more than other non-primate species.
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