Abstract: Feral pony mares, Equus caballus, at Assateague Island, Maryland, formed linear dominance hierarchies within bands. Generally, older mares dominated younger mares, and larger mares dominated smaller mares. Large mares initiated aggression more often than small mares when age was controlled for but, surprisingly, older mares initiated aggression less often than younger mares when size was controlled for. Thus, mares peak in aggressiveness fairly soon after achieving full size and then, while maintaining or improving their rank in the domainance hierarchy, progressively reduce their involvement in aggression as they grow older, Involvement in aggression per mare increased as number of mares in the group increased; this effect was independent of nearest-mare distances. Frequency of involvement in aggression did not differ between mares that had changed bands within the year and mares whose band association had continued for a year or more. Aggression was directed more frequently than expected at subordinate mares who were nursing, and also occurred more frequently than expected at water holes. The proportion of aggressive encounters during grazing closely matched the total proportion of time spent grazing. Subordinate mares with foals received aggression more often than subordinate mares without foals. The high frequency of aggression associated with foals and nursing suggests that interference with reproduction of subordiantes is an important mode of competition between mares. Such interference may be common in animals that feed on dispersed resources and live in small, cohesive groups.

Abstract: Individuals in a foraging group of wild bronwn capuchin monkeys choose different spatial positions relative to the rest of the group. Markov analysis of sequencess of individual spatial positions demonstrated significant differnces between individuals, which coul be categorized a posteriori into four homogenous subgroups. An individual's spatial position was related primarily to the amount of aggression it received from the group's dominant male, but also varied with its sex. Spatial choice varied with changes in an individual's social status, but did not vary consistently with seasonal differences in food availability. These results support the hypothesis that individuals compete for preferred spatial positions within a foraging group.

Abstract: Research on reciprocity is impaired by confusing definitions and often wrongly used statistical tests. Here, two models of the mechanism on which reciprocity may be based are discussed and an initial step towards a new fremework for its analysis is presented. A distinction is made between reciprocity and interchange. In the case of reciprocity, for one kind of act the same kind is received in return. In interchange, however, two different kinds of acts are bartered. Three types of reciprocity/interchange in social actions among all pairs of group-members are distinguished ([`]qualitative', [`]relative' and [`]absolute') on the basis of the precision of the reciprocity/interchange. Permutation procedures for association between matrices (such as the Mantel Z and two other newly derived tests) are used as a statistical test for detecting reciprocity/interchange. A rough comparison of the power of the two new tests is included. The tests can be applied to all kinds of group-living animals and to all sorts of social behaviour. The distinction between the three types of reciprocity/interchange and the matching statistical methods are also useful for defining and detecting other patterns in social interactions, like unidirectionality and associations between different kinds of social behaviour. The influence on social interactions of variables like dominance rank, age and sex can be analysed in the three forms by testing correlations between invented matrices which represent the influence of these variables (the so-called hypothesis matrices) and social interaction matrices. These methods are extended for two categories of individuals, thus allowing the investigation of, for example, reciprocity between males and females. The methods are illustrated with examples of coalition formation and grooming behaviour among captive chimpanzees, Pan troglodytes.

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: Many forms of interaction within and between species appear to be based on `scrounger' individuals or species exploiting a limited resource provided `producers'. A mathematical model is presented which shows whether or not scroungers are maintained in a group, depending on their frequency and the group size. Some of the predictions of the model were tested in captive flocks of house sparrows Passer domesticus L. Here the scroungers obtained most of their food (mealworms) by interaction and the producers found most of their food by actively foraging: the pay-off to each type was measured as mealworm capture rate. Neither type changed strategy opportunistically in response to instantaneous flock composition but, not surprisingly, scroungers fared better when one of more producers were present. However, scrougers did much worse than expected when greatly outnumbered by producers, perhaps because producers then found the available food very quickly.