The origin and evolution of the euarthropod labrum.

A widely (although not universally) accepted model of arthropod head evolution postulates that the labrum, a structure seen in almost all living euarthropods, evolved from an anterior pair of appendages homologous to the frontal appendages of onychophorans. However, the implications of this model for the interpretation of fossil arthropods have not been fully integrated into reconstructions of the euarthropod stem group, which remains in a state of some disorder. Here I review the evidence for the nature and evolution of the labrum from living taxa, and reconsider how fossils should be interpreted in the light of this. Identification of the segmental identity of head appendage in fossil arthropods remains problematic, and often rests ultimately on unproven assertions. New evidence from the Cambrian stem-group euarthropod Parapeytoia is presented to suggest that an originally protocerebral appendage persisted well up into the upper stem-group of the euarthropods, which prompts a re-evaluation of widely-accepted segmental homologies and the interpretation of fossil central nervous systems. Only a protocerebral brain was implicitly present in a large part of the euarthropod stem group, and the deutocerebrum must have been a relatively late addition.