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Smith, J. E., Kolowski, J. M., Graham, K. E., Dawes, S. E., & Holekamp, K. E. (2008). Social and ecological determinants of fission-fusion dynamics in the spotted hyaena. Anim. Behav., 76(3), 619–636.
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.
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Smith, J. E., Van Horn, R. C., Powning, K. S., Cole, A. R., Graham, K. E., Memenis, S. K., et al. (2010). Evolutionary forces favoring intragroup coalitions among spotted hyenas and other animals. Behav. Ecol., 21(2), 284–303.
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.
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