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THE EVOLUTION OF CLUTCH SIZE: A REPLY TO WOOTTON, YOUNG, AND WINKLER

Wootton et al. (1991) claim to have derived from the Lotka-Euler equation a clutch-size equation similar to the Murray-Nolan (M-N) clutch-size equation (Murray and Nolan, 1989) and that their equation tells us nothing about the evolution of clutch size. I agree. They claim further that these results indicate that neither my theory nor the M-N clutch-size equation (Murray, 1979, 1985, 1991a; Murray and Nolan, 1989) tells us anything about the evolution of clutch size. I disagree. My theory and the M-N equation cannot be fairly evaluated outside the philosophical, theoretical, and factual environment of which they are a part. (1) Philosophically, my hypothesis that selection favors those females that lay as few eggs as are necessary for replacement because they have the highest probability of surviving to breed again, their young have the highest probability of surviving to breed, or both (Murray, 1979, 1985, 1991a; Murray and Nolan, 1989) is an assumption of a hypothetico-deductive argument. Such assumptions are evaluated by comparing the predictions that follow from them with observation (see, for example, Newton, 1729; Popper, 1968, 1979). As Feynman (1985; p.1 0, original italics) points out, physicists have "leamed to realize that whether they like a theory or they don't like a theory is not the essential question. Rather, it is whether or not the theory gives predictions that agree with experiment." Wootton et al. (1991) do not question the predictions that I drew from the assumptions of my theory nor show that the predictions are contrary to fact. Nevertheless, they do not like my theory. (2) Wootton et al. (1991) claim that my clutch-size theory implicitly assumes that populations are neither increasing nor decreasing in size. This is not true. The dynamics of an evolving, genetically diverse population are complex. I think we have to sort through these complexities if we are interested in developing unifying, predictive theory. Let me try. (a) A more general form of the M-N equation, which will predict the clutch size (CS) of any genotype, whether increasing, decreasing, or in steady-state, is