Morphological Integration and Developmental Modularity

Biological systems, from molecular complexes to whole organisms and ecological interactions, tend to have a modular organization. Modules are sets of traits that are internally integrated by interactions among traits, but are relatively independent from other modules. The interactions within modules rely on different mechanisms, depending on the context of a study. For morphological traits, modularity occurs in developmental, genetic, functional, and evolutionary contexts. A range of methods for quantifying integration and modularity in morphological data is available, and a number of comparative and experimental designs can be used to compare the different contexts. How development produces covariation between traits can have substantial implications for understanding genetic variation and the potential for evolutionary change, but research in this area has only begun and many questions remain unanswered.