Group selection and inclusive fitness are not equivalent; the Price equation vs. models and statistics

It is often suggested that any group selection model can be recast in terms of inclusive fitness. A standard reference to support that claim is "Quantitative genetics, inclusive fitness, and group selection" by Queller (1992) in the American Naturalist 139 (3), 540-558. In that paper the Price equation is used for the derivation of this claim. Instead of a general derivation, we try out a simple model. For this simple example, we find that the result does not hold. The non-equivalence of group selection and kin selection is therefore not only an important finding in itself, but also a case where the use of the Price equation leads to a claim that is not correct. If results that are arrived at with the Price equation are not correct, they can typically be repaired by adding extra assumptions, or explicitly stating implicit ones. We give examples with relatively mild and with less mild extra assumptions. we also discuss why the Price equation is often referred to as dynamically insufficient, and we try to find out what Price's theorem could be. (c) 2011 Elsevier Ltd. All rights reserved.