The emergence of a superorganism through intergroup competition:
Surveys of insect societies have revealed four key, recurring organizational trends: (i) The most elaborated cooperation occurs in groups of relatives. (ii) Cooperation is typically more elaborate in species with large colony sizes than in species with small colony sizes, the latter exhibiting greater internal reproductive conflict and lesser morphological and behavioral specialization. (iii) Within a species, per capita brood output typically declines as colony size increases. (iv). The ecological factors of resource patchiness and intergroup competition are associated with the most elaborated cooperation. Predictions of all four patterns emerge elegantly from a game-theoretic model in which within-group tug-of-wars are nested within a between-group tug-of-war. In this individual selection model, individuals are faced with the problem of how to partition their energy between investment in intercolony competition versus investment in intracolony competition, i.e., internal tugs-of-war over shares of the resources gained through ...