joined two Harvard mathematicians and theoretical biologists, Martin Nowak and Corina Tarnita, for a top-to-bottom analysis of inclusive fitness. Nowak and Tarnita had independently discovered that the foundational assumptions of inclusive fitness theory were unsound, while I had demonstrated that the field data used to support the theory could be explained equally well, or better, with direct natural selection—as in the sex-allocation case of ants just described.

