Developmental integration in a complex morphological structure: how distinct are the modules in the mouse mandible?

The mouse mandible has long served as a model system for studying the development and evolution of complex morphological structures. We used the methods of geometric morphometrics to reassess the hypothesis that the mandible consists of two separate modules: an anterior part bearing the teeth and a posterior part with muscle attachment surfaces and articulating with the skull. The analyses particularly focused on covariation of fluctuating asymmetry, because such covariation is due exclusively to direct interactions between the developmental processes that produce the traits of interest, whereas variation of traits among individuals also reflects other factors. The patterns of fluctuating asymmetry and individual variation were only partly consistent, indicating that developmental processes contribute differentially to variation at different levels. The results were in agreement with the hypothesis that the anterior and posterior parts of the mandible are separate develop-mental modules. Comparison of all alternative partitions of the landmarks into two contiguous subsets confirmed the hypothesis for the location of the boundary between modules but also underscored that the separation between them is not complete. Modularity is therefore manifest as the relative independence of parts within the framework of overall integration of the mandible as a whole-it is a matter of degrees, not all or nothing.