Cryptic Persistence of Truncated Abdominal Legs in Insects Enabled Diverse Outgrowths with Novel Functions

An iconic feature of insects is the apparent lack of legs on the abdomen, which is believed to be due to the repression of the leg-patterning gene Distalless (Dll) by abdominal Hox genes. However, in contrast to these molecular observations, it is not widely appreciated that the embryos of most insect groups do in fact form paired protrusions on most abdominal segments that appear to be homologous to the thoracic legs. However, these degenerate before hatching to form the abdominal body wall. To resolve this discordance between molecular and morphological observations, the expression patterns of pannier and araucan, genes known to distinguish proximal leg segments in all arthropods, are examined in embryos of the flour beetle Tribolium castaneum. In Tribolium embryos, all pregenital abdominal segments develop leg-like paired protrusions, and the stripes of pannier and araucan expression that delineate the proximal leg segments of the thorax are also expressed in the same configuration around these abdominal protrusions. This suggests that insect abdominal legs are homologous to only the proximal portion of the thoracic legs, which in insect adults forms the body wall (lateral tergum and pleura). These cryptic, truncated abdominal legs – likely inherited from their crustacean ancestors – appear to be an important wellspring for new functions in insects, such as caterpillar prolegs, gills, and structures for camouflage¹⁶ and aposematic warning.