Morphometric integration and modularity in configurations of landmarks: tools for evaluating a priori hypotheses.

Identifying the modular components of a configuration of landmarks is an important task of morphometric analyses in evolutionary developmental biology. Modules are integrated internally by many interactions among their component parts, but are linked to one another only by few or weak interactions. Accordingly, traits within modules are tightly correlated with each other, but relatively independent of traits in other modules. Hypotheses concerning the boundaries of modules in a landmark configuration can therefore be tested by comparing the strength of covariation among alternative partitions of the configuration into subsets of landmarks. If a subdivision coincides with the true boundaries between modules, the correlations among subsets should be minimal. This article introduces Escoufier's RV coefficient and the multi-set RV coefficient as measures of the correlation between two or more subsets of landmarks. These measures can be compared between alternative partitions of the configuration into subsets. Because developmental interactions are tissue bound, it is sensible to require that modules should be spatially contiguous. I propose a criterion for spatial contiguity for sets of landmarks using an adjacency graph. The new methods are demonstrated with data on shape of the wing in Drosophila melanogaster and the mandible of the house mouse.