In the recent philosophical literature, two questions have arisen concerning the status of natural selection: 1) Is it a population-level phenomenon, or is it an organism-level phenomenon? 2) Is it a causal process, or is it a purely statistical summary of lower-level processes? In an earlier work (Millstein 2006), I argue that natural selection should be understood as a population-level causal process, rather than a purely statistical population-level summation of lower-level processes or as an organism-level causal process. In a 2009 essay entitled “Productivity, relevance, and natural selection,” Stuart Glennan argues in reply that natural selection is produced by causal processes operating at the level of individual organisms, but that ...