Species selection on variability

Most analyses of species selection require emergent, as opposed to aggregate, characters at the species level. This "emergent character" approach tends to focus on the search for adaptations at the species level. Such an approach seems to banish the most potent evolutionary property of populations--variability itself--from arguments about species selection (for variation is an aggregate character). We wish, instead, to extend the legitimate domain of species selection to aggregate characters. This extension of selection theory to the species level will concentrate, instead, on the relation between fitness and the species character, whether aggregate or emergent. Examination of the role of genetic variability in the long-term evolution of clades illustrates the cogency of broadening the definition of species selection to include aggregate characters. We reinterpret, in this light, a classic case presented in support of species selection. As originally presented, the species selection explanation of volutid neogastropod evolution was vulnerable to a counterinterpretation at the organism level. Once this case is recast within a definition of species selection that reflects the essential structure and broad applicability of hierarchical selection models, the organism-level reinterpretation of variability loses its force. We conclude that species selection on variability is a major force of macroevolution.