How to measure inclusive fitness.

Although inclusive fitness (Hamilton 1964) is regarded as the basic currency of natural selection, difficulty in applying inclusive fitness theory to field studies persists, a quarter-century after its introduction (Grafen 1982, 1984; Brown 1987). For instance, strict application of the original (and currently accepted) definition of inclusive fitness predicts that no one should ever attempt to breed among obligately cooperative breeders. Much of this confusion may have arisen because Hamilton's (1964) original verbal definition of inclusive fitness was not in complete accord with his justifying model. By re-examining Hamilton's original model, a modified verbal definition of inclusive fitness can be justified.