Using the Price equation to detect inclusive fitness in class-structured populations

Inclusive fitness theory has transformed the study of adaptive evolution since 1964, contributing to significant empirical findings. However, its status as a theory has been challenged by the proposals of several alternative frameworks. Those challenges have been countered by analyses that use the Price equation and the regression method. The Price equation is a universal description of evolutionary change, and the partitioning of the Price equation using the regression method immediately yields Hamiltons rule, which embodies the main tenets of inclusive fitness. Hamiltons rule captures the intensity and direction of selection acting on social behaviour and its underlying causal structure. Recent work, however, has suggested that there is an anomaly in this approach: in some cases, the regression method fails to estimate the correct values of the variables in Hamiltons rule and the causal structure of the behaviour. Here, I address this apparent anomaly. I argue that the failure of the simple regression method occurs because social players vary in baseline fecundity. I reformulate the Price equation and regression method to recover Hamiltons rule and I show that the method correctly estimates its key variables. I show that games where baseline fecundity varies among individuals represent a more general set of games that unfold in class-structured populations. This framework supports the robustness and validity of inclusive fitness.