Sex allocation and the emergence of helping in cooperatively breeding species

In cooperative breeding systems individuals invest in the reproductive success of others. In this paper, we study the emergence of cooperative breeding systems in which reproductively active breeders receive investment from reproductively non-active helpers. Our goal is to understand how the division of an investment between male and female components of breeder fitness (i.e. the helper sex-allocation strategy) influences the emergence of cooperative breeding itself. Using mathematical models, we arrive at expressions for the inclusive-fitness advantage of helpful behaviour that generalize previous work. These expressions assume an ecologically stable environment, and that breeders make evolutionarily stable sex-allocation decisions. We find that, when breeders are extremely resource limited, the sex-allocation strategy used by a helper can be a key determinant in the success of helpful alleles. This finding, however, is restricted to cases in which helpers have access to intermediate levels of resources. Surprisingly, when helpers can make only a small investment in a recipient the division of the investment matters only very little to advantage of help. By contrast when resources are extremely abundant, we obtain the unsurprising result that the manner in which resources are allocated has little influence on the emergence of help. When breeders have access to intermediate levels of resources we find increasing relatedness can, in certain cases, inhibit the emergence of help. We also find that increasing the amount of resources available to a breeder can impede help as well. Both of these counter-intuitive results are mediated by evolutionary responses in breeder sex allocation.