Why are arthropods segmented?

Segmentation as an attribute of organisms is being increasingly discussed in the recent literature because (1) new phylogenies suggest that organisms classically considered to be segmented may lie in separate clades; (2) the molecular basis of segmental development has been much studied; (3) various theories of bilaterian origins place weight on segmentation as a primitive character; (4) there has been recent stress on the importance of modularity as an evolutionary topic. However, the definition and extent of segmentation are highly ambiguous and usually typological. Here, segmentation is regarded as an attribute of organs, not organisms. The evolution of just one system, the arthropod epidermis, is examined on the basis of the fossil record and the extant euarthropods, tardigrades, and onychophorans. It may be seen to have become segmented in a complex pathway that necessitated shifts in function, redundancy, and changes in associated organs. This complexity must inevitably reflect on, and to an extent have primacy over, the genetic basis for the changes involved. Evolutionary functional morphology has been relatively little considered in the context of the evolution of development, but may play an important role in defining the framework within which this evolution occurs.