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Quantifying integration and modularity of evolutionary changes in morphometric traits is crucial for understanding how organismal shapes evolve. For this purpose, comparative studies are necessary, which need to take into account the phylogenetic structure of interspecific data. This study applies several of the standard tools of geometric morphometrics, which mostly have been used in intraspecific studies, in the new context of analyzing integration and modularity based on comparative data. Morphometric methods such as principal component analysis, multivariate regression, partial least squares, and modularity tests can be applied to phylogenetically independent contrasts of shape data. We illustrate this approach in an analysis of cranial evolution in 160 species from all orders of birds. Mapping the shape information onto the phylogeny indicates that there is a significant phylogenetic signal in skull shape. Multivariate regression of independent contrasts of shape on independent contrasts of size reveals clear evolutionary allometry. Regardless of whether or not a correction for allometry is used, evolutionary integration between the face and braincase is strong, and tests reject the hypothesis that the face and braincase are separate evolutionary modules. These analyses can easily be applied to other taxa and can be combined with other morphometric tools to address a wide range of questions about evolutionary patterns and processes.

Evolutionary Covariation in Geometric Morphometric Data: Analyzing Integration, Modularity, and Allometry in a Phylogenetic Context