Group augmentation and the evolution of cooperation

The group augmentation (GA) hypothesis states that if helpers in cooperatively breeding animals raise the reproductive success of the group, the benefits of living in a resulting larger group--improved survival or future reproductive success--favour the evolution of seemingly altruistic helping behaviour. The applicability of the GA hypothesis remains debatable, however, partly owing to the lack of a clear conceptual framework and a shortage of appropriate empirical studies. We conceptualise here the GA hypothesis and illustrate that benefits of GA can accrue via different evolutionary mechanisms that relate closely to well-supported general concepts of group living and cooperation. These benefits reflect several plausible explanations for the evolutionary maintenance of helping behaviour in cooperatively breeding animals.