Inclusive fitness models with two sexes (Q1105509)

Much recent work has focused on the transition from \textit{G. R. Price}'s [Nature 227, 520-521 (1970)] formula for allele frequency change to an inclusive fitness condition for the selective advantage of a certain behaviour. In case there is any kind of asymmetry between the sexes, the analysis must keep track of the two sexes separately, and this leads to a number of different forms of the expression for inclusive fitness. In this paper I gather these forms together and note the assumptions needed to make each valid. I also show how inclusive fitness should be formulated when the behaviour of the actor is controlled by another individual. I illustrate the inclusive fitness approach with a sex allocation example in a haplodiploid population with a local breeding structure.