Phylogeny and the hierarchical organization of plant diversity.

R. H. Whittaker's idea that plant diversity can be divided into a hierarchy of spatial components from alpha at the within-habitat scale through beta for the turnover of species between habitats to gamma along regional gradients implies the underlying existence of alpha, beta, and gamma niches. We explore the hypothesis that the evolution of alpha, beta, and gamma niches is also hierarchical, with traits that define the alpha niche being labile, while those defining beta and gamma niches are conservative. At the alpha level we find support for the hypothesis in the lack of close significant phylogenetic relationship between meadow species that have similar alpha niches. In a second test, alpha niche overlap based on a variety of traits is compared between congeners and noncongeners in several communities; here, too, there is no evidence of a correlation between alpha niche and phylogeny. To test whether beta and gamma niches evolve conservatively, we reconstructed the evolution of relevant traits on evolutionary trees for 14 different clades. Tests against null models revealed a number of instances, including some in island radiations, in which habitat (beta niche) and elevational maximum (an aspect of the gamma niche) showed evolutionary conservatism.