A quasi-equilibrium theory of the distribution of rare alleles in a subdivided population

The conditional average frequency of rare alleles has been shown in simulations to provide a simple and robust estimator of the number of individuals exchanged between local populations in an island model (Nm). This statistic is defined as the average frequency of an allele in those samples in which the allele is present. Here, we show that the conditional average frequency can be calculated from the distribution of allele frequencies. It is a measure of the spread of this distribution, and so is analogous to the standardised variance, FST. Analytic predictions for the island model of migration agree well with the corresponding simulation results. These predictions are based on the assumption that the rare alleles found in samples have reached a "quasi-equilibrium" distribution. As well as relating the conditional average frequency to the underlying allele frequency distribution, our results provide a more accurate method of estimating Nm from the conditional average frequency of private alleles in samples of different sizes.