Ecological Transitions in Grouping Benefits Explain the Paradox of Environmental Quality and Sociality.

Both benign and harsh environments promote the evolution of sociality. This paradox-societies occur in environments of such contrasting quality-may be explained by the different types of benefits that individuals receive from grouping: resource defense benefits that derive from group-defended critical resources versus collective action benefits that result from social cooperation among group members. Here, we investigate cooperative behavior in the burying beetle Nicrophorus nepalensis along an elevational gradient where environmental quality (climate and competition) varies with altitude. We show that climate (temperature) and competition (both intra- and interspecific) independently and synergistically influence sociality via different grouping benefits that vary along the gradient. At low elevations where interspecific competition for resources is intense, groups gain from the collective action benefit of increased interspecific competitive ability. In contrast, pairs have higher fitness at intermediate elevations where intraspecific competition for resources is greatest because resource defense is the key grouping benefit. However, groups and pairs have similar fitness at high elevations, suggesting that there is no grouping benefit in such physiologically challenging environments. Our results demonstrate that sociality is favored for different reasons under a range of environmental conditions, perhaps explaining why animal societies occur in environments of such contrasting quality.

Resolving the Paradox of Environmental Quality and Sociality: The Ecological Causes and Consequences of Cooperative Breeding in Two Lineages of Birds.

Cooperatively breeding animals occur in virtually every ecosystem on earth. Comparative and biogeographic studies suggest that both benign and harsh-as well as stable and fluctuating-environments can favor the evolution of cooperative breeding behavior. The fact that cooperative societies occur in environments of such contrasting quality creates a paradox of environmental quality and sociality. The dual benefits framework-which leads to the prediction that the ecological consequences of sociality (e.g., range size) vary depending on the benefits that individuals of each species receive by forming social groups-offers a potential resolution to this paradox. Here we use a case study of two avian lineages, starlings (Sturnidae) and hornbills (Bucerotidae), in which environmental unpredictability appears to have opposite effects on the evolution of cooperation to test the dual benefits framework. Consistent with previous work, harsh and unpredictable environments promote cooperative breeding behavior in starlings, which in turn leads to larger geographic ranges. However, cooperatively breeding hornbills occur in benign and stable environments, but sociality does not influence range size. Our study suggests that the paradox of environmental quality and sociality arises largely because cooperative breeding is an umbrella term encompassing social species that form groups for different reasons. We demonstrate that differentiating among the functional causes of social group formation is critical for developing a predictive framework for understanding the evolution of cooperative breeding behavior.