Sperm show patterns of rapid and divergent evolution that are characteristic of sexual selection. Sperm competition has been proposed as an important selective agent in the evolution of sperm morphology. However, several comparative analyses have revealed evolutionary associations between sperm length and female reproductive tract morphology that suggest patterns of male-female coevolution. In the dung beetle Onthophagus taurus, males with short sperm have a fertilization advantage that depends on the size of the female's sperm storage organ, the spermatheca; large spermathecae select for short sperm. Sperm length is heritable and is genetically correlated with male condition. Here we report significant additive genetic variation and heritability for spermatheca size and genetic covariance between spermatheca size and sperm length predicted by both the ''goodsperm'' and ''sexy-sperm'' models of postcopulatory female preference. Our data thus provide quantitative genetic support for the role of a sexually selected sperm process in the evolutionary divergence of sperm morphology, in much the same manner as precopulatory female preferences drive the evolutionary divergence of male secondary sexual traits.male-female coevolution ͉ postcopulatory female choice ͉ sperm length F emale mating preferences are widely recognized as being responsible for the rapid and divergent evolution of male secondary sexual traits (1). Indicator or good gene models envisage genetic coupling between male sexual trait expression and offspring fitness, so that females with a preference for male traits produce offspring of greater viability (2). Once a preference becomes established, females choosing males with elaborate secondary sexual traits will produce sons that carry alleles for the trait and daughters that carry alleles for the preference, generating genetic coupling that will drive self-reinforcing coevolution of both trait and preference because of the mating advantage to males with the trait. Thus, the original viability benefits associated with the preference can be undermined by a runaway Fisherian sexy sons process (2). Analogous models have been proposed for postcopulatory female preferences (3). ''Good-sperm'' models predict positive genetic associations between a male's sperm competitiveness and the general viability of his offspring (4), whereas ''sexy-sperm'' models predict that multiply mating females produce sons successful in sperm competition and daughters that incite sperm competition through multiple mating (5, 6). As with precopulatory processes, postcopulatory models predict that the trait in males that determines fertilization success will become genetically coupled with the mechanism by which females bias sperm use toward preferred males (7).Although spermatozoa are well known for their rapid and divergent morphological variation (8, 9), little is known of the selective processes that drive sperm evolution. Patterns of divergent evolution are characteristic of strong sexual selection, and researchers have sugges...