Multi-level selection and the issue of environmental homogeneity

In this paper, I identify two general positions with respect to the relationship between environment and natural selection. These positions consist in claiming that selective claims need and, respectively, need not be relativized to homogenous environments. I then show that adopting one or the other position makes a difference with respect to the way in which the effects of selection are to be measured in certain cases in which the focal population is distributed over heterogeneous environments. Moreover, I show that these two positions lead to two different interpretations-the Pricean and contextualist ones-of a type of selection scenarios in which multiple groups varying in properties affect the change in the metapopulation mean of individual-level traits. Showing that these two interpretations stem from different attitudes towards environmental homogeneity allows me to argue: (a) that, unlike the Pricean interpretation, the contextualist interpretation can only claim that drift or selection is responsible for the change in frequency of the focal trait in a given metapopulation if details about whether or not group formation is random are specified; (b) that the traditional main objection against the Pricean interpretation-consisting in arguing that the latter takes certain side-effects of individual selection to be effects of group selection-is unconvincing. This leads me to suggest that the ongoing debate about which of the two interpretations is preferable should concentrate on different issues than previously thought.