Abstract: Interventions occur when animals interfere in competitive interactions between two or more individuals. Interveners can alter the nature of the ongoing interaction by targeting one party (attacking, biting) and supporting the other. Three theories have been proposed to account for intervention behaviour: kin selection, reciprocity and direct benefits. The kin selection hypothesis predicts that interveners will selectively support relatives over nonrelatives; the reciprocity hypothesis predicts that when intervener [`]A' supports individual [`]B', later [`]B' will intervene and support [`]A'; and the direct benefits hypothesis predicts that target/support patterns should serve the immediate interests of the intervener. We tested the reciprocity and direct benefits hypotheses by exploring third-party interventions in play fighting among littermates of domestic dogs. Interveners in dyadic play did not preferentially target or support preferred playmates of the intervener. Interveners targeted the dog in the losing role at the time of the intervention, and they did not show reciprocity in support. Taken together, these last two findings suggest that littermates benefit directly and use interventions opportunistically to practise offence behaviours directed at littermates already behaving subordinately. Opportunities to practise targeting in a playful setting may help structure dominance relationships among littermates. Additionally, the tendency for puppies to do what the other is doing (target the dog in the losing role) may pave the way for synchronizing cooperative behaviours during group hunting and territorial defence. The types of behaviours used to intervene changed over development, but the outcome following an intervention remained stable.

Abstract: Personality in animal behaviour describes the observation that behavioural differences between individuals are consistent over time and context. Studies of group-living animals show that movement order among individuals is also consistent over time and context, suggesting that some individuals lead and others follow. However, the relationship between leadership and personality traits is poorly studied. We measured several personality traits and leadership of individual barnacle geese, Branta leucopsis. We measured body size and scored the dominance of individuals living in a stable group situation before subjecting them to an open-field test, an activity test, a novel-object test, and a leadership test in which the order of the movement of individuals in pairs towards a feeding patch was scored. We found high repeatability for activity and novel-object scores over time. Leadership was strongly correlated with novel-object score but not with dominance rank, activity or exploration in an open field. These results provide evidence that leadership is closely related to some aspects of personality. Interestingly, an individual's arrival at the food patch was affected not only by the novel-object score of the focal individual, but also by the novel-object score of the companion individual, indicating that movement patterns of individuals living in groups are affected by the personality traits of other group members and suggesting that movement patterns of a group may be shaped by the mix of personality types present in the group.

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: Cerebral lateralization, the preferential use of one hemisphere of the brain to perform certain cognitive functions, is a widespread and evolutionarily ancient adaptation. Lateralization appears to enhance cognitive capacity, yet substantial individual variation in the strength cerebral lateralization is apparent in all species studied so far. It is puzzling that cerebral lateralization, a seemingly advantageous trait, has not been driven to fixation. It has been suggested that variation in lateralization may be linked to individual variation in behaviour, which itself may be subject to disruptive selection. We examined the relation between cerebral lateralization and individual variation in boldness in the convict cichlid, Archocentrus nigrofasciatus. We show that convict cichlids that are more strongly lateralized when exploring a familiar environment, but not a novel one, are quicker to emerge from a shelter in a test for boldness. The possibility that cerebral lateralization is linked to life history strategy is discussed.

Abstract: Play behaviour is widespread in mammals, but benefits to play have been difficult to demonstrate. Physical training is one of the many proposed hypotheses, suggesting that males and females should play differently, that increased maternal investment should lead to increases in play, and that increases in play should result in physical advantages. In a population of feral horses, Equus caballus, males and females did not differ in their play behaviour except that males initiated more of their play bouts than females. Maternal condition influenced play behaviour only in males, with sons of mothers in good condition playing more. However, when we controlled for maternal effects by comparing a son and a daughter of the same mother, daughters played more when their mother was in poor condition and sons played more when their mother was in good condition. Mothers of foals that played more lost more condition. Therefore, the difference in play behaviour could not be explained by offspring sex or maternal condition alone, but play behaviour mirrored variation in maternal investment. In addition, those individuals that played more survived better and had better body condition as yearlings despite weaning earlier. Since increased activity has been linked to enhanced musculoskeletal development in domestic horses, we suggest that play provides a link between increased maternal investment, increased body condition and future reproductive success in feral horses, and probably in other species.