Abstract: In studying the success of foraging animals, studies of interference competition have put emphasis on effects of competitor density, whereas studies of resource defense have focused on the effects of the spatial distribution of food within patches. Very few studies have looked at both factors simultaneously, that is, determined whether the effects of competitor density on foraging success depend on the spatial distribution of food. We studied the behavior and the foraging success of ruddy turnstones (Arenaria interpres) using an experiment in which we varied both the presence of a competitor and the food distribution. Because turnstones may differ strongly in their relative dominance status, we also experimentally varied the foragers' relative dominance status. We found that the presence of a competitor only reduced the foraging success of subordinate birds foraging at the clumped food distribution. At this condition, dominant and subordinate birds differed markedly in their foraging success. Contrary to our expectations, we did not observe more agonistic behavior at the clumped food distribution. This indicates that the amount of agonistic behavior observed may be a bad indicator of interference effects. These findings have specific implications for models of interference competition. Most notably they show that the effects of competitor density on agonistic behavior and foraging success may well depend on the spatial distribution of food and the foragers' relative dominance status. Additionally, our results suggest that social dominance will not be fully understood without considering long-term processes such as the formation and maintenance of social dominance hierarchies.

Abstract: We present the results of a series of computer simulations that examined the impact of winner, loser, and bystander effects on hierarchy formation in fused groups. These effects and their implications for hierarchy structure and aggressive interactions were first examined in small four-member groups. Subsequent to this, the two small groups were fused into a single larger group. Further interactions took place in this fused group, generating a new hierarchy. Our models demonstrate clearly that winner, loser, and bystander effects strongly influence both the structure and types of interactions that emerge from the fusion of smaller groups. Four conditions produced results in which the same general patterns were uncovered in pre- and postfusion groups: (1) winner effects alone, (2) bystander loser effects alone, (3) winner and bystander winner effects operating simultaneously, and (4) all four effects in play simultaneously. Outside this parameter space, hierarchy structure and the nature of aggressive interactions differed in pre- and postfusion groups. When only loser effects were in play, one of the two clear alphas from the prefused groups dropped in rank in the eight-member fused group. When bystander winner effects were in play, it was difficult to rank any of the eight individuals in the fused group, and players interacted almost exclusively with those that were not in their original four-member group. When loser and bystander loser effects operated simultaneously, two top-ranking individuals emerged in the fused groups, but the relative rank of the other players was difficult to assign.

Abstract: Prior modeling work has found that pure winner and loser effects (i.e., changing the estimation of your own fighting ability as a function of direct prior experience) can have important consequences for hierarchy formation. Here these models are extended to incorporate “bystander effects.” When bystander effects are in operation, observers (i.e., bystanders) of aggressive interactions change their assessment of the protagonists' fighting abilities (depending on who wins and who loses). Computer simulations demonstrate that when bystander winner effects alone are at play, groups have a clear omega (bottom-ranking individual), while the relative position of other group members remains difficult to determine. When only bystander loser effects are in operation, wins and losses are randomly distributed throughout a group (i.e., no discernible hierarchy). When pure and bystander winner effects are jointly in place, a linear hierarchy, in which all positions (i.e., {alpha} to {delta} when N = 4) are clearly defined, emerges. Joint pure and bystander loser effects produce the same result. In principle one could test the predictions from the models developed here in a straightforward comparative study. Hopefully, the results of this model will spur on such studies in the future.

Abstract: In many cooperatively breeding species, group members form a dominance hierarchy or queue to inherit the position of breeder. Models aimed at understanding individual variation in helping behavior, however, rarely take into account the effect of dominance rank on expected future reproductive success and thus the potential direct fitness costs of helping. Here we develop a kin-selection model of helping behavior in multimember groups in which only the highest ranking individual breeds. Each group member can invest in the dominant's offspring at a cost to its own survivorship. The model predicts that lower ranked subordinates, who have a smaller probability of inheriting the group, should work harder than higher ranked subordinates. This prediction holds regardless of whether the intrinsic mortality rate of subordinates increases or decreases with rank. The prediction does not necessarily hold, however, where the costs of helping are higher for lower ranked individuals: a situation that may be common in vertebrates. The model makes two further testable predictions: that the helping effort of an individual of given rank should be lower in larger groups, and the reproductive success of dominants should be greater where group members are more closely related. Empirical evidence for these predictions is discussed. We argue that the effects of rank on stable helping effort may explain why attempts to correlate individual helping effort with relatedness in cooperatively breeding species have met with limited success.

Abstract: One of the most prominent behavioural features of many forest primates are the loud calls given by the adult males. Early observational studies repeatedly postulated that these calls function in intragroup spacing or intergroup avoidance. More recent field experiments with Diana monkeys (Cercopithecus diana) of Taï Forest, Ivory Coast, have clearly shown that loud male calls function as predator alarm calls because calls reliably (1) label different predator classes and (2) convey semantic information about the predator type present. Here, I test the alarm call hypothesis another primate, the Campbell's monkey (C. campbelli). Like Diana monkeys, male Campbell's monkeys produce conspicuous loud calls to crowned hawk eagles (Stephanoaetus coronatus) and leopards (Panthera pardus), two of their main predators. Playback experiments showed that monkeys responded to the predator category represented by the different playback stimuli, regardless of whether they consisted of (1) vocalisations of the actual predators (crowned hawk eagle shrieks or leopard growls), (2) alarm calls to crowned hawk eagles or leopards given by other male Campbell's monkeys or (3) alarm calls to crowned hawk eagles or leopards given by sympatric male Diana monkeys. These experiments provide further evidence that non-human primates have evolved the cognitive capacity to produce and respond to referential labels for external events.