Phylogenies and the Comparative Method

Abstract

Recent years have seen a growth in numerical studies using the comparative method. The method usually involves a comparison of two phenotypes across a range of species or higher taxa, or a comparison of one phenotype with an environmental variable. Objectives of such studies vary, and include assessing whether one variable is correlated with another and assessing whether the regression of one variable on another differs significantly from some expected value. Notable recent studies using statistical methods of this type include Pilbeam and Gould's (1974) regressions of tooth area on several size measurements in mammals; Sherman's (1979) test of the relation between insect chromosome numbers and social behavior; Damuth's (1981) investigation of population density and body size in mammals; Martin's (1981) regression of brain weight in mammals on body weight; Givnish's (1982) examination of traits associated with dioecy across the families of angiosperms; and Armstrong's (1983) regressions of brain weight on body weight and basal metabolism rate in mammals. My intention is to point out a serious statistical problem with this approach, a problem that affects all of these studies. It arises from the fact that species are part of a hierarchically structured phylogeny, and thus cannot be regarded for statistical purposes as if drawn independently from the same distribution. This problem has been noticed before, and previous suggestions of ways of coping with it are briefly discussed. The nonindependence can be circumvented in principle if adequate information on the phylogeny is available. The information needed to do so and the limitations on its use will be discussed. The problem will be discussed and illustrated with reference to continuous variables, but the same statistical issues arise when one or both of the variables are discrete, in which case the statistical methods involve contingency tables rather than regressions and correlations.