Complex cooperative strategies in group-territorial African lions.

Female lions (Panthera leo) showed persistent individual differences in the extent to which they participated in group-territorial conflict. When intergroup encounters were simulated by playback of aggressive vocalizations, some individuals consistently led the approach to the recorded intruder, whereas others lagged behind and avoided the risks of fighting. The lead females recognized that certain companions were laggards but failed to punish them, which suggests that cooperation is not maintained by reciprocity. Modification of the "odds" in these encounters revealed that some females joined the group response when they were most needed, whereas other lagged even farther behind. The complexity of these responses emphasizes the great diversity of individual behavior in this species and the inadequacy of current theory to explain cooperation in large groups.