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Multi-level selection can be understood via the Price equation or contextual analysis, which offer incompatible statistical decompositions of evolutionary change into components of group and individual selection. Okasha (Evolution and the levels of selection, 2006) argued that each approach suffers from problem cases. I introduce further problem cases for the Price approach, arguing that it is appropriate for MLS 2 group selection but not MLS 1. I also show that the problem cases Okasha raises for contextual analysis can be resolved. For some such cases, however, it emerges that there is no determinate answer to the question of how much of the total selective effect was due to group selection compared to individual selection. This suggests that when there is interaction between the effect of group character and individual character, one cannot separate selection into distinct ‘levels’ at all.

DOI: 10.1007/s11229-014-0569-0

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