ESTIMATING HIERARCHICAL F-STATISTICS.

This paper presents an analysis of variance (ANOVA) approach by which estimation of F-statistics can be made from data with an arbitrary s-level hierarchical population structure. Assuming a complete random-effect model, a general ANOVA procedure is developed to estimate F-statistics as ratios of different variance components for all levels of population subdivision in the hierarchy. A generalized relationship among F-statistics is also derived to extend the well-known relationship originally found by Sewall Wright. Although not entirely free from the bias particular to small number of subdivisions at each hierarchy and extreme gene frequencies, the ANOVA estimators of F-statistics consider sampling effects at each level of hierarchy, thus removing the bias incurred in the other estimators that are commonly based on direct substitution of unknown gene frequencies by their sample estimates. Therefore, the ANOVA estimation procedure presented here may become increasingly useful in analyzing complex population structure because of increasing use of the estimated hierarchical F-statistics to infer genetic and demographic structures of natural populations within and among species.