Life history, habitat saturation and the evolution of fecundity and survival altruism.

Hamilton's rule provides a general description of the conditions for the evolution of altruism. But altruism can take different forms depending on which life-history trait is affected by the helping behavior (fecundity vs. survival helping). In particular, these different forms of helping may have very different demographic consequences, which may feed back on evolution. We examine the interplay between various forms of helping and demography in viscous populations with empty sites. A key component of our analysis is the local density of empty sites experienced by a focal individual, which provides a measure of habitat saturation. Habitat saturation is shown to have contrasting effects depending on (1) whether the physiological costs and benefits of helping affect fecundity, survival or both; and (2) whether the costs of helping are paid in a density-dependent or density-independent manner. For a given level of habitat saturation and with density-dependent reproduction, we find that the conditions for the evolution of helping should be more favorable in the survival altruism life cycle with a cost on fecundity, and more stringent in the fecundity altruism life cycle with a cost on survival. More generally, our analysis stresses the importance of taking into account the feedback between population demography, life history, and kin selection when investigating the selective pressures on altruism.