Class Structure, Demography, and Selection: Reproductive-Value Weighting in Nonequilibrium, Polymorphic Populations

In natural populations, individuals of a given genotype may belong to different classes. Such classes can represent different age groups, developmental stages, or habitats. Class structure has important evolutionary consequences because the fitness of individuals with the same genetic background may vary depending on their class. As a result, demographic transitions between classes can cause fluctuations in the trait mean that need to be removed when estimating selection on a trait. Intrinsic differences between classes are classically taken into account by weighting individuals by class-specific reproductive values, defined as the relative contribution of individuals in a given class to the future of the population. These reproductive values are generally constant weights calculated from a constant projection matrix. Here, I show for large populations and clonal reproduction that reproductive values can be defined as time-dependent weights satisfying dynamical demographic equations that depend only on the average between-class transition rates over all genotypes. Using these time-dependent demographic reproductive values yields a simple Price equation where the nonselective effects of between-class transitions are removed from the dynamics of the trait. This generalizes previous theory to a large class of ecological scenarios, taking into account density dependence, ecological feedbacks, arbitrary strength of selection, and arbitrary trait distributions. I discuss the role of reproductive values for prospective and retrospective analyses of the dynamics of phenotypic traits.