Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? Samir Okasha provides a comprehensive analysis of the debate in evolutionary biology over the levels of selection, focusing on conceptual, philosophical and foundational questions. A systematic framework is developed for thinking about natural selection acting at multiple levels of the biological hierarchy; the framework is then used to help resolve outstanding issues. Considerable attention is paid to the concept of causality as it relates to the levels of selection, in particular the idea that natural selection at one hierarchical level can have effects that 'filter' up or down to other levels. Unlike previous work in this area by philosophers of science, full account is taken of the recent biological literature on 'major evolutionary transitions' and the recent resurgence of interest in multi-level selection theory among biologists. Other biological topics discussed include