Abstract: In a wide range of animal species, males coerce females to mate with them, either by physically forcing them to mate, by harassing them until they mate or by punishing persistent refusal to mate. The first section of this paper argues that the possibility of forced copulation can generate arms races between males and females that may have substantial costs to both sexes. In the second section, it is suggested that sexual harassment commonly represents a `war of attrition' between the sexes; existing game theory models that may apply to sexual conflict over mating decisions are reviewed. The third section develops a simple prospective model for the evolution of intimidation by punishment in situations where males can raise the probability that females will accept their advances in future by punishing them for refusal to mate. Where the benefits of sexual coercion to males are high, all three male strategies may develop to a point where they have substantial costs to females. In the final section, evidence that female behaviour is adapted to minimizing these costs is reviewed.

Abstract: In nature foragers must exploit resources that vary randomly in both the energy acquired per item (reward) and the time required to pursue, capture and process an item (delay). Furthermore, rewards and delays associated with particular resources may often covary significantly. An analytical model asks how variance-covariance levels for rewards and delays could influence choice of resources when lack of information or cognitive limitation implies that a consumer attempts to maximize its short-term rate of energy gain. Both greater expected reward and reduced expected delay clearly should enhance preference for a resource. The model predicts that increased delay variance and reduced reward-delay covariance should increase a forager's preference for a resource. A forager should be risk-averse towards reward variance when the reward-delay covariance is positive, but should become risk-prone towards reward variance when the reward-delay covariance is negative.

Abstract: Theory predicts that individuals living in fission-fusion societies, in which group members frequently change subgroups, should modify grouping patterns in response to varying social and environmental conditions. Spotted hyaenas, Crocuta crocuta, are long-lived carnivores that reside in permanent social groups called clans. Clans are complex, fission-fusion societies in which individual members travel, rest and forage in subgroups that frequently change composition. We studied two clans in Kenya to provide the first detailed description of fission-fusion dynamics in this species. Because social and ecological circumstances can influence the cohesiveness of animal societies, we evaluated the extent to which specific circumstances promote the formation of subgroups of various sizes. We found that cooperative defence of shared resources during interclan competition and protection from lions were cohesive forces that promoted formation of large subgroups. We also tested hypotheses suggesting factors limiting subgroup size. Mothers with small cubs avoided conspecifics, thereby reducing infanticide risk. Victims of aggression either reconciled fights or separated from former opponents to reduce the immediate costs of escalated aggression in the absence of food. As predicted by the ecological constraints hypothesis, hyaenas adjusted their grouping patterns over both short and long time scales in response to feeding competition. Crocuta were most gregarious during periods of abundant prey, joined clanmates at ephemeral kills in numbers that correlated with the energetic value of the prey and gained the most energy when foraging alone because cooperative hunting attracted numerous competitors. Overall, our findings indicate that resource limitation constrains grouping in this species.