title: The emergence of leaders and followers in foraging pairs when the qualities of individuals differ.
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.
  
  
  CONCLUSIONS
  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.
authors.affiliation: Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK. sean.rands@bristol.ac.uk
authors.forename: Sean A
authors.initials: SA
authors.lastname: Rands
authors.forename: Guy
authors.initials: G
authors.lastname: Cowlishaw
authors.forename: Richard A
authors.initials: RA
authors.lastname: Pettifor
authors.forename: J Marcus
authors.initials: JM
authors.lastname: Rowcliffe
authors.forename: Rufus A
authors.initials: RA
authors.lastname: Johnstone
mesh_terms: D000818:Animals; D005247:Feeding Behavior; D020387:Food Chain; D008954:Models, Biological; D012930:Social Dominance