Abstract: According to the socioecological framework, transitivity (or linearity) in dominance relationships is related to competition over critical resources. When a population is structured into groups, the intensity of between- versus within-group competition influences the form and function of its social organization. Few studies have compared the type and relative intensity of competition at these two levels. African elephants have well-structured social relations, providing an exemplary system for such a study. We report on dominance hierarchies among free-ranging elephants and evaluate the factors that drive their socioecological structure to lie in a region of the three-dimensional nepotism/despotism/tolerance space rarely observed among social species; namely, where non-nepotistic, transitive dominance hierarchies within groups emerge despite kin-based philopatry and infrequent agonistic interactions over widely distributed resources. We found significant transitivity in dominance hierarchies between groups. Dominance relations among the matriarchs of different social groups were primarily age based, rather than driven by physical or group size, and group matriarch rank influenced the dominance relationships among nonmatriarchal females in the population. Our results suggest that between-group dominance relationships induce tolerance among group members, which in combination with high group relatedness, reduces the benefits of nepotism. We postulate that cognitive abilities and high risk of injury in contests enhance winner and loser effects, facilitating the formation of transitive dominance relationships, despite widely distributed resources over which infrequent competition occurs. The interplay of cognitive abilities, winner and loser effects, resource distribution, and within- and between-group dominance relationships may produce behaviour in other strongly social mammals that differs from that predicted by a superficial application of current socioecological models.

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: 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: 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: The role of dominant individuals in leading groups of animals is not yet well understood. We investigated whether dominant beef cows, Bos taurus, have more influence on herd movement on pasture than more subordinate cows. A herd of 15 Gasconne cows was observed for a 3-week period between dawn and dusk. The positions of all adult cows were recorded with GPS collars at 1 min intervals and the behaviour of each cow was recorded in 5 min scans. The dominance hierarchy was recorded by ad libitum sampling. Through cluster analysis of the recorded data, we distinguished three herd behaviour patterns: resting, foraging and travelling. Dominant cows were closer to the front of the herd during both travelling and foraging. During travelling, more dominant cows also had more direct trajectories and were more aligned both with their nearest neighbours and with the whole herd. During foraging, the trajectories of dominant cows were shorter than those of subordinate cows. The results indicate that foraging and short-distance travelling movements by female beef cattle are not led by any particular individual but rather are influenced by a graded type of leadership; that is, the more dominant a cow is, the stronger the influence it may have on the movements of the herd.