Darwinian fitness, its directional derivative, and Hamilton's rule for limited dispersal with class structure under within and between generation environmental stochasticity
This paper formalizes Darwinian invasion fitness and its phenotypic derivative for group-structured populations under limited dispersal and environmental stochasticity, demonstrating how Hamilton's marginal rule can be derived as an actor-centered inclusive-fitness effect that integrates class-specific fitness differentials, relatedness, and reproductive values.