The components of directional and disruptive selection in heterogeneous group-structured populations

We derive how directional and disruptive selection operate on scalar traits in a heterogeneous group-structured population for a general class of models. In particular, we assume that each group in the population can be in one of a finite number of states, where states can affect group size and/or other environmental variables, at a given time. Using up to second-order perturbation expansions of the invasion fitness of a mutant allele, we derive expressions for the directional and disruptive selection coefficients, which are sufficient to classify the singular strategies of adaptive dynamics. These expressions include first- and second-order perturbations of individual fitness (expected number of settled offspring produced by an individual, possibly including self through survival); the first-order perturbation of the stationary distribution of mutants (derived here explicitly for the first time); the first-order perturbation of pairwise relatedness; and reproductive values, pairwise and three-way relatedness, and stationary distribution of mutants, each evaluated under neutrality. We introduce the concept of individual k-fitness (defined as the expected number of settled offspring of an individual for which k-1 randomly chosen neighbors are lineage members) and show its usefulness for calculating relatedness and its perturbation. We then demonstrate that the directional and disruptive selection coefficients can be expressed in terms individual k-fitnesses with k=1,2,3 only. This representation has two important benefits. First, it allows for a significant reduction in the dimensions of the system of equations describing the mutant dynamics that needs to be solved to evaluate explicitly the two selection coefficients. Second, it leads to a biologically meaningful interpretation of their components. As an application of our methodology, we analyze directional and disruptive selection in a lottery model with either hard or soft selection and show that many previous results about selection in group-structured populations can be reproduced as special cases of our model.