Inclusive-fitness logic of cooperative breeding with benefits of natal philopatry

In cooperatively breeding species, individuals help to raise offspring that are not their own. We use two inclusive-fitness models to study the advantage of this kind of helpful behaviour in social groups with high reproductive skew. Our first model does not allow for competition among relatives to occur but our second model does. Specifically, our second model assumes a competitive hierarchy among nest-mates, with non-breeding helpers ranked higher than their newborn siblings. For each model, we obtain an expression for the change in inclusive fitness experienced by a helpful individual in a selfish population. The prediction suggested by each expression is confirmed with computer simulation. When model predictions are compared to one another, we find that helping emerges under a broader range of conditions in the second model. Although competition among kin occurs in our second model, we conclude that the life-history features associated with this competition also act to promote the evolutionary transition from solitary to cooperative breeding.