Abstract: Behavioral ecologists have long been interested in understanding the adaptive value of antipredator behavior (Sih 1987Go; Lima and Dill 1990Go; Lima 1998Go). A recent review by Lind and Cresswell (2005)Go, however, noted some important difficulties with quantifying the fitness consequences of antipredator behaviors. In essence, Lind and Cresswell suggest that most studies do not provide strong evidence on the adaptive value of antipredator behavior because they do not consider 1) trade-offs between antipredator and reproductive performance, 2) the abilities of organisms to avoid fitness losses associated with constraints on focal traits by employing behavioral alternatives (behavioral compensation), and 3) the effects of behavioral defenses at different stages of the predation sequence. The authors rightfully assert that an understanding of these issues can only be accomplished by measuring multiple traits and fitness components (i.e., survival and reproduction). Nevertheless, the question of how to integrate such data into

Abstract: Coalitionary support in agonistic interactions represents cooperation because intervening in a fight is potentially costly to the donor of support but benefits the recipient. Here, we first review the characteristics of, and evolutionary forces favoring, intragroup coalitions in 49 species and find that patterns of intragroup coalition formation are remarkably similar between primates and nonprimates. We then test hypotheses suggesting kin selection, reciprocal altruism, and direct benefits as adaptive explanations for coalitionary interventions among adult female spotted hyenas (Crocuta crocuta) belonging to a large social group in Kenya. As predicted by kin selection theory, females supported close kin most often, and the density (connectedness) of cooperation networks increased with genetic relatedness. Nevertheless, kinship failed to protect females from coalitionary attacks. We found no evidence of enduring alliances based on reciprocal support among unrelated adult females. Instead, donors generally minimized costs to themselves, intervening most often during low-intensity fights and when feeding opportunities were unavailable. Females also gained direct benefits from directing coalitionary attacks toward subordinates. Finally, females monitored the number of dominant bystanders in the “audience” at fights and modified their level of cooperation based on this knowledge. Overall, hyenas made flexible decisions regarding whether or not to intervene in fights, modifying their tendency to cooperate based on multiple types of information about their immediate social and ecological environments. Taken together, these findings indicate that the combined evolutionary forces of kin selection and direct benefits derived from reinforcing the status quo drive coalitionary interventions among adult female spotted hyenas.

Abstract: Considerable interspeci®c variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current ``ecological model'', predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social elationships among gregarious females are here characterized as DispersalEgalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained i€erences in the occurrence of formal submission signals, decided dominance relation ships, coalitions and female philopatry. Group size and female rank generally a€ect female reproduction success as the model predicts, and studies of closely related species in di€erent ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the e€ects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment.
Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains
several phenomena that the ecological model does not, is a ``socioecological model'' based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail.

Abstract: Summary The introduction of game-theoretical thinking into evolutionary biology has laid the groundwork for a heuristic view of animal behaviour in which individuals employ “strategies” – rules that instruct them how to behave in a given circumstance to maximize relative fitness. Axelrod and Hamilton (1981) found that a strategy called Tit-For-Tat (TFT) is one robust cooperative solution to the iterated Prisoner's Dilemma game. There exists, however, little empirical evidence that animals employ TFT. Predator inspection in fish provides one ecological context in which to examine the use of the TFT strategy.