Abstract: BACKGROUND: Foraging in groups offers animals a number of advantages, such as increasing their likelihood of finding food or detecting and avoiding predators. In order for a group to remain together, there has to be some degree of coordination of behaviour and movement between its members (which may in some cases be initiated by a decision-making leader, and in other cases may emerge as an underlying property of the group). For example, behavioural synchronisation is a phenomenon where animals within a group initiate and then continue to conduct identical behaviours, and has been characterised for a wide range of species. We examine how a pair of animals should behave using a state-dependent approach, and ask what conditions are likely to lead to behavioural synchronisation occurring, and whether one of the individuals is more likely to act as a leader. RESULTS: The model we describe considers how the energetic gain, metabolic requirements and predation risks faced by the individuals affect measures of their energetic state and behaviour (such as the degree of behavioural synchronisation seen within the pair, and the value to an individual of knowing the energetic state of its colleague). We explore how predictable changes in these measures are in response to changes in physiological requirements and predation risk. We also consider how these measures should change when the members of the pair are not identical in their metabolic requirements or their susceptibility to predation. We find that many of the changes seen in these measures are complex, especially when asymmetries exist between the members of the pair. CONCLUSION: Analyses are presented that demonstrate that, although these general patterns are robust, care needs to be taken when considering the effects of individual differences, as the relationship between individual differences and the resulting qualitative changes in behaviour may be complex. We discuss how these results are related to experimental observations, and how the model and its predictions could be extended.

Abstract: The variability of most environments taxes foraging decisions by increasing the uncertainty of the information available. One solution to the problem is to use dynamic averaging, as do some granivores and carnivores. Arguably, the same strategy could be useful for grazing herbivores, even though their food renews and is more homogeneously distributed. Horses (Equus callabus) were given choices between variable patches after short or long delays. When patch information was current, horses returned to the patch that was recently best, whereas those without current information matched choices to the long-term average values of the patches. These results demonstrate that a grazing species uses dynamic averaging and indicate that, like granivores and carnivores, they can use temporal weighting to optimize foraging decisions.

Abstract: We tested for social learning and imitation in common marmosets using an artificial foraging task and trained conspecific demonstrators. We trained a demonstrator marmoset to open an artificial fruit, providing a full demonstration of the task to be learned. Another marmoset provided a partial demonstration, controlling for stimulus enhancement effects, by eating food from the outside of the apparatus. We thus compared three observer groups, each consisting of four animals: those that received the full demonstration, those that received the partial demonstration, and a control group that saw no demonstration prior to testing. Although none of the observer marmosets succeeded in opening the artificial fruit during the test periods, there were clear effects of demonstration type. Those that saw the full demonstration manipulated the apparatus more overall, whereas those from the control group manipulated it the least of the three groups. Those from the full-demonstration group also contacted the particular parts of the artificial fruit that they had seen touched (localised stimulus enhancement) to a greater extent than the other two groups. There was also an interaction between the number of hand and mouth touches made to the artificial fruit for the full- and partial-demonstration groups. Whether or not these data represent evidence for imitation is discussed. We also propose that the clear differences between the groups suggest that social learning mechanisms provide real benefits to these animals in terms of developing novel food-processing skills analogous to the one presented here.

Abstract: It is possible that non-specialised cues transmitted by conspecifics guide animals' food search provided they have the cognitive abilities needed to read these cues. Macaques often check the mouth of their group-mates by olfactory and/or visual inspection. We investigated whether Tonkean macaques ( Macaca tonkeana) can find the location of distant food on the basis of cues conveyed by group-mates. The subjects of the study were two 6-year-old males, who belonged to a social group of Tonkean macaques raised in semi-free-ranging conditions. In a first experiment, we tested whether the subject can choose between two sites after having sniffed a partner who has just eaten food corresponding to one of the sites. We found that both subjects were able to choose the matching site significantly above the chance level. This demonstrated that Tonkean macaques are capable of delayed olfactory matching. They could associate a food location with an odour conveyed by a partner. In a second experiment, the same subjects were allowed to see their partner through a Plexiglas window. Both subjects were still able to choose the matching site, demonstrating they could rely on visual cues alone. Passive recruitment of partners appears possible in macaques. They can improve their foraging performances by finding the location of environmental resources from olfactory or visual cues conveyed by group-mates.

Abstract: Japanese workers have studied social acquisition patterns of new feeding habits in Macaca fuscata which they have termed precultural. The present study investigates the same phenomenon in the chacma baboon and the vervet monkey in their natural habitat. The questions addressed are: (1) How a new feeding habit enters a troop and by which age and sex category, also how it is propagated? (2) When individuals are permitted with a choice between palatable and unpalatable food, can they learn by demonstration only or do they have to pass through a direct learning process? (3) Can the results from the above questions be explained by social parameters such as the social structure of the individual species? It was found that juvenile baboons discover new food and that after the discovery propagation is instantaneous. In vervets discovery is random among the age classes and propagation is slow and takes place through certain 'pivot' individuals. Both species fail to learn about palatability by demonstration but have to go through a direct learning process. This contrasts strongly with the forest baboon Mandrillus sphinx that have been shown to learn by demonstration. Socially, baboon juveniles stay closer to each other than the adults who force them to live at the periphery of the troop. Vervets again forage without precise sub-group formation. The link between social and cultural propagation and social structure is discussed on the basis of these findings.