HETEROCHRONY AND ALLOMETRY: LESSONS FROM THE WATER STRIDER GENUS LIMNOPORUS.

Heterochrony and allometry both deal with evolutionary modifications of ontogenies. Although data about both morphology and age are required to identify heterochronic processes, age data are not needed to study allometry. Using a simple graphical model, we show that allometric patterns cannot be used to infer the underlying heterochronic processes. We present a case study of the water strider genus Limnoporus Stål (Heteroptera: Gerridae) to illuminate the distinct roles that allometry and heterochrony play in integrated studies of the evolution of form. Multivariate analyses reveal several evolutionary modifications of growth trajectories (changes in direction, lateral transposition, and ontogenetic scaling), which are fairly consistent with the hypothesized phylogeny of the genus. Because there is no positive correlation between instar durations and size increments, size cannot be used as a proxy for age data in studies of heterochrony. In fact, a measure of overall size itself shows a remarkable variety of heterochronic changes among the six species. Mixtures of several heterochronic processes predominate over the more unitary reflections of "pure" processes. Heterochronic changes in different branches of the phylogeny, apparently independent of size scaling, suggest considerable potential for adaptive evolution. "Local" differentiation of ontogenetic traits within small clades may be at least as important as "global" evolutionary trends in large clades and will often be missed in "global" analyses.