Abstract: In most cooperatively breeding birds and mammals, reproductive rates are lower for social subordinates than for dominants, and it is common for reproduction in subordinates to be completely suppressed. Early research conducted in captivity showed that losing fights can increase glucocorticoid (GC) secretion, a general response to stress. Because GCs can suppress reproduction, it has been widely argued that chronic stress might underlie reproductive suppression of social subordinates in cooperative breeders. Contradicting this hypothesis, recent studies of cooperative breeders in the wild show that dominant individuals have elevated GCs more often than do subordinates. The findings that elevated GCs can be a consequence of subordination or a cost of dominance complicate the conventional view of social stress, with broad ramifications for the evolution of dominance and reproductive suppression.

Abstract: Achieving a high social rank may be advantageous for individuals at high population densities, because dominance status may determine the priority of access to limited resources and reduce individual loss of body mass. The establishment of dominance relationships between individuals involves variable levels of aggressiveness that can be influenced by resource availability. The relationship between social rank and aggressiveness and the impacts of resource abundance on aggressiveness are, however, poorly understood, but may be relevant to understand the mechanisms determining dominance relationships between individuals. We experimentally simulated, in seminatural enclosures, a deterioration of winter forage quality induced by a high-density deer population and examined the effects of (1) social dominance and (2) diet quality on aggressiveness, forage intake and body mass loss of white-tailed deer, Odocoileus virginianus, fawns during two winters. Within diet-quality treatments, fawns were consistently organized into linear hierarchies and showed clear dominance relationships. Dominants initiated more interactions and showed higher aggressiveness than subordinates, but subordinates had higher forage intake than dominants throughout winter. Social rank did not influence cumulative body mass loss of fawns. During both winters, fawns fed the control diet maintained their aggressiveness level, whereas fawns fed the poor-quality diet decreased it. Our experimental approach revealed that white-tailed deer responded to a reduction in winter forage quality by modifying their aggressiveness, indicating that ungulates may show plasticity not only in their foraging behaviour in response to decreased resources but also in their social behaviour.

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.