Abstract: We spent two winters studying the social behaviour of wild woodland caribou (Rangifer tarandus caribou) at a time when their main food (ground lichens; Cladina sp.) is available only at snow craters dug by the animals. The competition for access to such craters was severe, the animals constantly trying to take over the craters of others. During a two-month period when a group maintained a constant size (20) and composition (all age-sex classes represented), we could rank the animals in a rather linear dominance hierarchy (Landau's index = 0.87). Rank was correlated with access to resources, percent of time spent active, and percent of time feeding in craters. It was also correlated with age and antler size. However, rank is not an attribute of individuals, but of a relationship between individuals. As such it is only an intervening variable between physical attributes and access to resources, a variable whose value has meaning only within a given group. Among the three attributes studied (age, sex, antler size), the latter was by far the best predictor of the occurrence and outcome of interactions. Between two individuals within any of the three age-sex classes studied (adult and yearling males and adult females), the one with larger antlers initiated significantly more often, escalated its aggression (to the point of hitting the target) less often, and enjoyed a higher success rate in obtaining resources. When their antlers were larger than those of an adult male target (i.e. males that had shed their antlers), adult females won almost all their interactions with adult males even though they escalated only one fourth of them. This clarifies the long-standing speculation that female caribou have antlers and shed them later than males, in order to overcome their sexual handicap in competition for food in the winter. We conclude that the link between rank and dominance of an individual on one hand, and some of its attributes on the other (e.g. sex, age, weight, antler size) is fundamentally realized by the animal itself through its active preference for targets it is likely to beat, i.e. targets with smaller antlers.

Abstract: This simulation study explores some conditions leading to transitivity within dominance orders. Combinations of three parameters were varied to study their consequences upon hierarchy formation and upon the degree of linearity of resultant structures. The factors studied were: (1) the importance of initial resource holding potentials (RHPs); (2) changes brought in RHPs by successive victories and defeats; and (3) accuracy of RHP assessment made by opponents. Results show that initial differences in RHP always lead to perfectly transitive chains whose rank order reflects the importance of initial differences. Even when simulated animals make important errors while assessing each other during round robin tournaments, emerging dominance structures are perfectly linear and ranks obtained in the structure are highly correlated with initial values in RHPs. Moreover, accumulated experiences of victory and/or defeat alone always lead to perfectly linear hierarchies. Their combination with initial individual differences in RHP led to the same conclusion. Even when assessment was far from being perfect, not only perfect chains were formed but initial values in RHPs significantly influenced rank order when the contribution of victory and defeat to RHP was relatively unimportant. The higher the importance of victory and defeat to RHP as compared to that of initial RHP values, the lower was the correlation between initial RHP values and the ranks order reached by individuals in the resultant hierarchies. In general also, the lower the variation within initial RHPs, the lower was the correlation between initial RHPs and ranks in the hierarchy. At a given level of initial RHP dispersion, increasing the contribution of victory and defeat to RHP diminished the correlation between initial RHP values and obtained ranks. In addition, inaccurate assessment reduced the overall correlation, especially when dispersion of initial RHP values was low and the contribution of victory and defeat relatively unimportant. These results shed some light on the controversy about the respective roles of initial individual attributes and that of patterns of resolution in the formation of animal hierarchies. We present the emergence of social order within closed systems as those simulated here as a case of self-organization.

Abstract: Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10-20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks.

Abstract: The function of olfactory signalling in social species is less well understood than in asocial species. Consequently, we examined olfactory communication in the ringtailed lemur, a socially complex primate that retains a functional vomeronasal organ, has well-developed scent glands and shows a suite of scent-marking behaviour. To assess the information content of different types of scent gland secretions, we decoupled olfactory cues from the visual and behavioural modalities with which scent marking is normally associated. We presented male and female subjects (signal receivers) with a series of choice tests between odours derived from conspecific donors (signal senders) varying by sex, age, social status and reproductive condition. We additionally examined the influence of the receivers' reproductive state and familiarity with the signaller. The reproductive condition, social status and familiarity of senders and receivers affected signal transmission; specifically, male receivers attended most to the odours of conspecifics in breeding condition and to the odours of familiar, dominant animals. By contrast, females varied their responses according to both their own reproductive state and that of the sender. Based on male and female patterns of countermarking, we suggest that scent marking serves a function in intergroup spacing and intrasexual competition for both sexes, as might be expected in a female-dominant species. By contrast, minimal female interest in male odours counters a female mate choice function for scent marking in this species. Nevertheless, scent marks are critical to male-male competition and, therefore, may be subject to sexual selection.

Abstract: A social hierarchy is generally assumed to exist in those mammalian societies in which the costs and benefits of group living are distributed unevenly among group members. We analysed infrared closed-circuit television footage, collected over 3 years in Wytham Woods, Oxfordshire, to test whether social groups of European badgers have dominance hierarchies. Analysis of directed aggression between dyads revealed linear dominance hierarchies in three social-group-years, but patterns within social groups were not consistent across years. Dominance hierarchies were significantly steeper than random in five out of six social-group-years. In those social-group-years where a linear hierarchy was determined, there was an effect of sex on dominance rank, with females gaining significantly higher rank than males in two social-group-years. Overall, rank was not related to age, nor did it appear to affect the likelihood of an individual being wounded, or an individual's breeding status. The latter resulted from nonorthogonality between sex and breeding status, as there were only two breeding males. Overall, hierarchies were primarily dominated by breeding females, and may occur when breeding competition arises. Relatedness, unreciprocated allogrooming and sequential allomarking were not consistently related to levels of directed aggression across social-group-years. We suggest that dominance structures within European badger groups may be context dependent, with future study required to complete our understanding of where, and when, they arise.