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In their seminal paper published in 1979, Gould and Lewontin argued that some traits arise as by-products of the development of other structures and not for direct utility in themselves. We show here that this applies to the trident, a pigmentation pattern observed on the thorax of  Drosophila melanogaster . Using reporter constructs, we show that the expression domain of several genes encoding pigmentation enzymes follows the trident shape. This domain is complementary to the expression pattern of  stripe (sr) , which encodes an essential transcription factor specifying flight muscle attachment sites. We demonstrate that  sr  limits the expression of these pigmentation enzyme genes to the trident by repressing them in its own expression domain,  i.e . at the flight muscle attachment sites. We give evidence that repression of not only  yellow  but also other pigmentation genes, notably  tan , is involved in the trident shape. The flight muscle attachment sites and  sr  expression patterns are remarkably conserved in dipterans reflecting the essential role of  sr . Our data suggest that the trident is a by-product of flight muscle attachment site patterning that arose when  sr  was co-opted for the regulation of pigmentation enzyme coding genes.

Pigmentation pattern and developmental constraints: flight muscle attachment sites delimit the thoracic trident of Drosophila melanogaster