Abstract: Male ungulates engage in intense competition for access to females during the breeding season. Although fights are generally dyadic level encounters, they are on occasion disrupted by the intervention of third-party males. We investigated these third-party interventions using predictions derived from Dugatkin's model (Dugatkin 1998, Proceedings of the Royal Society of London, Series B, 265, 433-437) of intervention behaviour. The model argues that when an individual successfully defeats an opponent there is an increase in the probability of winning a subsequent contest: a winner effect. Third-party intervention behaviour is predicted to occur as it serves to prevent either member of a competing dyad from successfully defeating his opponent, achieving a winner effect and subsequently becoming a threat to the intervener. Consistent with model predictions, our results show that intervening males held significantly higher rank than males that did not intervene and were also more likely to be dominant to both of the competing males. Intervening males did not selectively target competitors based on rank, nor did they target males based on overall dyadic rates of aggression between the intervener and competing males. Furthermore, interveners were more likely to have won their interaction immediately prior to intervention and were also likely to win their interaction subsequent to intervention when compared with contest success of the two competing males. Our results are consistent with predictions that support a winner effect for intervention behaviour in fallow deer fights.

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: Controversy exists regarding the nature of primate social relationships. While individual primates are frequently hypothesized to form enduring social bonds with conspecifics, recent studies suggest that relationships are labile, with animals interacting only over short periods to satisfy their immediate needs. Here I use data collected over 10 years on a community of chimpanzees, Pan troglodytes, at Ngogo, Kibale National Park, Uganda, to investigate whether male chimpanzees establish long-term social relationships and to determine the factors that affect variation in relationship quality and the stability of social bonds. Kinship and dominance rank influenced the quality of relationships. Maternal brothers and males of the same dominance rank class groomed each other more equitably than did unrelated males and males that were dissimilar in rank. In addition, males that formed strong social bonds groomed more equitably than did males that displayed weaker bonds. Social bonds were stable over time, with relationships in one year predicting those in subsequent years. Kinship and the quality of social relationships affected bond stability. Maternal half siblings and males that groomed each other equitably maintained longer-lasting bonds than did nonkin and males that groomed each other unevenly. Virtually all of the males established at least one enduring relationship with another individual. The most enduring bonds formed between a few pairs of maternal brothers and dyads that maintained balanced grooming interactions. These results indicate that male chimpanzees maintain long-lasting and equitable social bonds whose formation is affected by maternal kinship and the quality of social relationships.

Abstract: One of the big challenges for group-living animals is to find out who in a group has pertinent information (regarding food or predators) at any moment in time, because informed individuals may not be obviously recognizable to other group members. We found that individuals in human groups were capable of identifying those with information, and this identification increased group performance: the speed and accuracy of groups in reaching a target. Using video analysis we found how informed individuals might have been identified by other group members by means of inadvertent social cues (such as starting order, time spent following and group position). Furthermore, we were able to show that at least one of these cues, the group position of informed individuals, was indeed correlated with group performance. Our final experiment confirmed that leadership was even more efficient when the group members were given the identity of the leader. We discuss the effect of information status regarding the presence and identity of leaders on collective animal behaviour.

Abstract: In animals, behavioural properties such as aggressive propensity are often consistent over a life span, and they may form part of a behavioural syndrome. We studied how aggressive propensity influences dominance, contest behaviour and growth in the cooperatively breeding cichlid fish Neolamprologus pulcher. We tested whether intrinsic aggressive propensity (1) influences dominance in paired contests, (2) causes different aggression levels in contests with partners matched for aggressive propensity compared to unmatched partners, and how it (3) affects growth rate in groups that were either matched or unmatched for aggressive propensity. Intrinsic aggressive propensity was first scored with a mirror test and classified as high, medium or low. Thereafter we tested fish with either high or low aggressive propensity with partners matched for size and either matched or unmatched for aggressive type in a paired contest for a shelter. We scored dominance, aggression and submission. As predicted, (1) dominance was more clearly established in unmatched than in matched contests and (2) individuals with high aggressive propensity launched more attacks overall than fish with low intrinsic aggressiveness, suggesting a higher propensity to escalate independently of winning or losing the paired contest. However, contrary to expectation, (3) individuals with low aggressiveness grew faster than aggressive ones in unmatched groups, whereas the opposite occurred in matched groups. This suggests that individuals with low aggressive propensity may benefit from conflict evasion, which might allow them to gain dominance in the future owing to larger body size.