![]() ![]() Genetic correlations have been extensively documented for many morphological (and other) traits (examples in ref. The modularity of developing organisms has supposedly facilitated independent evolution of groups of traits belonging to different modules, but it also may have led to the concerted evolution of traits within one module ( 1, 2). The idea that in animals groups of traits are developmentally integrated within modules has been receiving much attention in evolutionary developmental biology ( 1–5). Our results are discussed within the context of the evolution of modularity and individuality of serially repeated morphological traits. The ease with which we have been able to produce independent responses to artificial selection on different eyespots may be linked to a legacy of natural selection favoring individuality. ![]() We argue that among-eyespot correlations are unlikely to have constrained the evolutionary diversification of butterfly wing patterns but might be important when only limited time is available for adaptive evolution to occur. We show that, in Bicyclus anynana butterflies, despite the evidence that all eyespots are developmentally coupled, the response to selection for increased size of one individual eyespot can proceed in a manner largely independent from selection imposed on another eyespot. Here, we use artificial selection to explore the modular organization of butterfly wing patterns and the extent to which their evolution is constrained by the genetic correlations among repeated pattern elements. These have often been postulated to explain patterns of morphological variation and have been examined theoretically but seldom analyzed experimentally. Correlations among traits describe potential developmental constraints on evolution. Modularity in animal development is thought to have facilitated morphological diversification, but independent change of those traits integrated within a module might be restricted.
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