Selective advantage of redirected helping in a viscous population.

When a brood fails, the failed parent can help a neighbor rear its offspring. This behavior is known as redirected helping and occurs in various species. The advantage of redirected helping may seem obvious, provided the individual whose brood fails helps a related neighbor: the helper at least gains indirect fitness by redirecting its parental effort. However, complications arise when considering a viscous population, where individuals remain on or close to their natal site. In such a population, individuals compete with relatives, which dilutes the advantage of helping and may counteract it altogether. This raises a question: when can we expect redirected helping to evolve in a viscous population? We address this question with inclusive fitness models. We find that redirected helping can always be favored in a viscous population, provided the cost is sufficiently low. We also identify life-history features - like survival, dispersal, and brood-failure rate - that promote redirected helping. The effect of these life-history features, in general, depends on which component of fitness (survival or fecundity) benefits from help and how brood failure varies among demes. Unlike previous authors, we find that helping can be more strongly promoted when it provides survival rather than fecundity benefits.