Comparative studies tend to differ from optimality and functionality s
tudies in how they treat adaptation. While the comparative approach fo
cuses on the origin and change of traits, optimality studies assume th
at adaptations are maintained at an optimum by stabilizing selection.
This paper presents a model of adaptive evolution on a macroevolutiona
ry time scale that includes the maintenance of traits at adaptive opti
ma by stabilizing selection as the dominant evolutionary force. Inters
pecific variation is treated as variation in the position of adaptive
optima. The model illustrates how phylogenetic constraints nor only le
ad to correlations between phylogenetically related species, but also
to imperfect adaptations. From this model, a statistical comparative m
ethod is derived that can be used to estimate the effect of a selectiv
e factor on adaptive optima in a way that would be consistent with an
optimality study of adaptation to this factor. The method is illustrat
ed with an analysis of dental evolution in fossil horses. The use of c
omparative methods to study evolutionary trends is also discussed.