The origin and evolution of appendages.

Current awareness of gene expression patterns and developmental mechanisms involved in the outgrowth and patterning of animal appendages contributes to our understanding of the origin and evolution of these body parts. Nevertheless, this vision needs to be complemented by a new adequate comparative framework, in the context of a factorial notion of homology. It may even be profitable to categorize as appendages also gut diverticula, body ingrowths and 'virtual appendages' such as the eye spots on butterfly wings. Another unwarranted framework is the Cartesian co-ordinate system onto which the appendages are currently described and where it is supposed that one patterning system exists for each separate Cartesian axis. It may be justified, instead, to look for correspondences between the appendages and the main body axis of the same animal, as the latter might be the source of the growth and patterning mechanisms which gave rise to the former. This hypothesis of axis paramorphisms is contrasted with the current hypothesis of gene co-option. Recapitulationism is a common fault in current Evo-Devo perspectives concerning the origin of the appendages, in that the evolutionary origin of appendages is often expected to be the same as one of the key mechanisms involved in the ontogenetic inception of appendage formation. This unwarranted perspective is also evident in the current debate on the nature of the default arthropod appendage. Most likely, a default arthropod appendage never did exist, as the first appendages probably developed along the trunk of an animal already patterned extensively along the antero-posterior body axis.