The ontogeny of the insect labrum, or upper lip, has been debated for nearl
y a century. Recent molecular data suggest a segmental appendage origin of
this structure. Here we report the first arthropod mutation associated with
a homeotic transformation of the labrum. Antennagalea-5 (Ag-5) transforms
both antennal and labral structures to resemble those of gnathal appendages
in Tribolium castaneum. This labral transformation suggests that the labru
m is a fused structure composed of two pairs of appendage endites, and is s
erially homologous to the gnathal appendages.