Modes of cooperation during territorial defense by African lions.

Cooperation during territorial defense allows social groups of African lions to defend access to resources necessary for individual reproductive success. Some forms of cooperation will be dependent upon cognition: reciprocity places greater cognitive demands on participants than does kinship or mutualism. Lions have well-developed cognitive abilities that enable individuals to recognize and interact with others in ways that seem to enhance their inclusive fitness. Male lions appear to cooperate unconditionally, consistently responding to roaring intruders regardless of their male companions' kinship or behavior. Female lions, however, do keep track of the past behavior of their female companions, apparently using the reliability of a companion as one means of assessing the risks posed by approaching intruders. Some "laggard" females may exploit the cooperative tendencies of "leaders" during territorial encounters. Although leader females clearly recognize laggards as such, the costs of tolerating laggards may be less than the benefits leaders gain through territorial defense behavior. Thus, although lions clearly have the cognitive ability to base cooperation on reciprocity, territorial defense cooperation appears instead to be based primarily on mutual benefits to participants for both male and female lions.