Abstract: It was formerly argued that alternative evolutionarily stable strategies (ESSs) are possible for animal contests characterized by some asymmetry that can be perceived with perfect accuracy. Where roles A and B refer to the asymmetry between opponents, ESSs are: [`]fight when A, retreat when B', and vice versa. Either can be an ESS, but only if the [`]reserve strategy' (=what an animal does when it fights) is sufficiently damaging. We examine the [`]war of attrition' (winner = opponent that persists longer). In a population at either ESS, reserve strategy is never normally shown; it is therefore subject to drift unless the selective action of rare individuals which break the convention is considered. These could arise either by mutation or by mistakes in role assessment. When mutations and mistakes simply specify that occasionally an animal fights when it [`]should' retreat, selection adjusts reserve strategy to a level where only one ESS (the [`]commonsense' ESS) is possible, if the asymmetry is relevant to payoff. Thus for asymmetries in fighting ability or resource value, the individual with the lower score will retreat. However, we are particularly concerned with cases where both payoff-relevant aspects (fighting ability and resource value) are asymmetric. If opponents sustain contest costs at rates KA and KB, and their resource values are VA and VB, an [`]optimal assessor' strategy defined by the interaction between the two asymmetries, is a unique ESS. It obeys the rule [`]fight on estimating role A, where VA/KA>VB/KB; retreat in B'. If mistakes can occur in both roles, but are very rate, the ESS is not fundamentally altered though there will be infinitesimal tendencies for persisting in role B. Selection to improve assessment abilities intensifies as abilities improve, but is weak if roles A and B are rather similar. Over a range of similarity between roles, an [`]owner wins' convention may be adopted if ownership correlates positively with role A and an individual cannot tell when it would otherwise pay him to break the convention. We also examine a contest in which information about roles can be acquired only during a contest itself, and at a cost. Much depends on the rate at which information is acquired relative to the rate at which costs are expended, and on whether contests normally escalate in intensity, remain at the same level, or de-escalate. Selection favours short contests when costs are high relative to resource value, where the outcome of a round contains much information about fighting ability, and where the actual disparity in fighting ability is large.

Abstract: For species in which group membership frequently changes, it has been a challenge to characterize variation in individual interactions and social structure. Quantifying this variation is necessary to test hypotheses about ecological determinants of social patterns and to make predictions about how group dynamics affect the development of cooperative relationships and transmission processes. Network models have recently become popular for analyzing individual contacts within a population context. We use network metrics to compare populations of Grevy's zebra (Equus grevyi) and onagers (Equus hemionus khur). These closely related equids, previously described as having the same social system, inhabit environments differing in the distribution of food, water, and predators. Grevy's zebra and onagers are one example of many sets of coarsely similar fission-fusion species and populations, observed elsewhere in other ungulates, primates, and cetaceans. Our analysis of the population association networks reveals contrasts consistent with their distinctive environments. Grevy's zebra individuals are more selective in their association choices. Grevy's zebra form stable cliques, while onager associations are more fluid. We find evidence that females associate assortatively by reproductive state in Grevy's zebra but not in onagers. The current approach demonstrates the utility of network metrics for identifying fine-grained variation among individuals and populations in association patterns. From our analysis, we can make testable predictions about behavioral mechanisms underlying social structure and its effects on transmission processes.

Abstract: Leadership is not an inherent quality of animal groups that show directional locomotion. However, there are other factors that may be responsible for the occurrence of leadership in fish shoals, such as individual differences in nutritional state between group members. It appears that front fish have a strong influence on directional shoal movements and that individuals that occupy such positions are often characterised by larger body lengths and lower nutritional state. Potential interactions between the two factors and their importance for positioning within shoals need further attention. Initiation of directional movement in stationary shoals and position preferences in mobile shoals need to be addressed separately because they are potentially subject to different constraints. Individuals that initiate a swimming direction may not necessarily be capable of the sustained high swimming performance required to keep the front position or have the motivation to do so, for that matter. More empirical and theoretical work is necessary to look at the factors controlling positioning behaviour within shoals, as well as overall shoal shape and structure. Tracking of marked individuals whose positioning behaviour is monitored over extended time periods of hours or days would be useful. There is an indication that shoal positions are rotated by individuals according to their nutritional needs, with hungry fish occupying front positions only for as long as necessary to regain their nutritional balance. This suggests that shoal members effectively take turns at being leaders. There is a need for three-dimensional recordings of shoaling behaviour using high-speed video systems that allow a detailed analysis of information transfer in shoals of different size. The relationship between leadership and shoal size might provide an interesting field for future research. Most studies to date have been restricted to shoals of small and medium size and more information on larger shoals would be useful.