The Drosophila spineless (ss) gene is regulated downstream of the appendage gene Distal-less (Dll) and is involved in leg and antenna development. Specifically, loss of ss leads to the homeotic transformation of the arista, the distalmost antennal segment, into tarsal identity, and the loss or fusion of distal leg segments. Here we show that the ss homolog from the red flour beetle Tribolium castaneum also homeotically transforms the beetle antenna into leg, but the extent of the transformation is significantly larger than in Drosophila, as the entire antenna (except for the basal antennifer) is transformed into pretarsal, tibiotarsal, and femoral identity; i.e., the transformation comprises the Dll positive area in both appendages. We interpret the antennal phenotype in Tribolium as evidence for a more exclusive role of ss in antennal determination downstream of Dll in the beetle. By contrast, the fact that, in Drosophila ss mutants, only a small portion of the Dll positive area in the antenna is homeotically transformed indicates that Dll uses additional targets to govern the development of the other antennal segments in the fly.