Figure 1. Number- and frequency-dependence in mimicry and aposematism.

Müller's number-dependent theory supposes that, while they learn to avoid the pattern of an unpalatable insect, predators kill a constant number, nk(i) of each morph i per unit time in a given area. Assuming the local population has constant size (N) and contains a novel pattern (A) and a "wild-type" pattern (a), Müller's theory can give the strength of frequency-dependent selection for or against the pattern A at different frequencies (qA) in the population. The fitness of A is WA = 1 - (nk(A)/qAN), while that of a is Wa = 1 - [nk(a)/(1-qA)N]. The measure of frequency-dependent selection acting on A relative to a used here is SA = (WA/Wa) - 1; if SA is positive, A is favored, if SA is negative, A is disfavored. The dashed and solid lines show the frequency-dependence for a low total population size (N = 10) and a high total population size (N = 100), respectively, relative to nk(A) and nk(a) (the fractions nk/N are important, rather than absolute values of nk and N). In contrast, linear frequency-dependent selection has been more normally used to study the population genetics of warning color and mimicry (37, 47, 54, 91, 93), for example where WA = 1 - sA(1 - qA), and Wa = 1 - saqA. This model gives the frequency-dependence shown in the dotted curve of the figure. In both number-dependent and frequency-depenedent selection, values of s and nk have been chosen to give an unstable equilibrium frequency of qA* = 0.4, which could be the case if A has 1.5× greater fitness than a.