A single gene orchestrates androgen variation underlying male mating morphs in ruffs

Androgens are pleiotropic and play pivotal roles in the formation and variation of sexual phenotypes. We show that differences in circulating androgens between the three male mating morphs in ruff sandpipers are linked to 17-beta hydroxysteroid dehydrogenase 2 (HSD17B2), encoded by a gene within the supergene that determines the morphs. Low-testosterone males had higher HSD17B2 expression in blood than high-testosterone males, as well as in brain areas related to social behaviors and testosterone production. Derived HSD17B2 isozymes, which are absent in high-testosterone males but preferentially expressed in low-testosterone males, converted testosterone to androstenedione faster than the ancestral isozyme. Thus, a combination of evolutionary changes in regulation, sequence, and structure of a single gene introduces endocrine variation underlying reproductive phenotypes.