From the Price equation to the selection gradient in class-structured populations: a quasi-equilibrium route

Recent studies in theoretical evolutionary ecology have emphasised two approaches to modelling evolution. On the one hand, models based on a separation of time scales rely on the concept of invasion fitness. On the other hand, models based on the Price equation track the dynamics of a trait average, coupled with a description of ecological dynamics. The aim of this article is to show that, in class-structured populations, both approaches yield the same expression for the selection gradient under weak selection. Although the result is not new, I propose an alternative route to its derivation using the dynamics of scaled measures of between-class phenotypic differentiation. Under weak selection, these measures of phenotypic differentiation can be treated as fast variables compared to the trait mean, which allows for a quasi-equilibrium approximation. This suggests a different approach to calculating weak selection approximations of evolutionary dynamics, and clarifies the links between short- and long-term perspectives on evolution in structured populations.