Multilevel social structure predicts individual helping responses in a songbird.

Multilevel societies are formed when stable groups of individuals spatially overlap and associate preferentially with other groups, producing a hierarchical social structure. Once thought to be exclusive to humans and large mammals, these complex societies have recently been described in birds. However, it remains largely unclear what benefits individuals gain by forming multilevel societies. One hypothesis-based on food sharing in hunter-gatherers-is that multilevel societies facilitate access to a range of cooperative relationships, with individual investment varying across the hierarchical levels of the society. We tested experimentally whether such graded cooperation occurs in the multilevel society of a songbird, the superb fairy-wren (Malurus cyaneus). Specifically, we measured whether responses to playbacks of distress calls-used to recruit help when in extreme danger-varied according to the social level at which the focal individual is connected with the caller. We predicted that anti-predator responses should be highest within breeding groups (the core social unit), intermediate between groups from the same community, and lowest across groups from different communities. Our results confirm that birds exhibit the predicted hierarchical pattern of helping and that, within breeding groups, this pattern is independent of kinship. This pattern of graded helping responses supports the hypothesis that multilevel social structures can sustain stratified cooperative relationships and reveals similarity in cooperation in qualitatively different behaviors-anti-predator behavior and food sharing-in the multilevel societies of songbirds and humans.