Complex interactions with females and rival males limit the evolution of sperm offence and defence

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Abstract
Postcopulatory sexual selection favours males which are strong offensive and defensive sperm competitors. As a means of identifying component traits comprising each strategy, we used an experimental evolution approach. Separate populations ofDrosophila melanogasterwere selected for enhanced sperm offence and defence. Despite using a large outbred population and evidence of substantive genetic variation for each strategy, neither trait responded to selection in the two replicates of this experiment. Recent work with fixed chromosome lines ofD. melanogastersuggests that complex genotypic interactions between females and competing males contribute to the maintenance of this variation. To determine whether such interactions could explain our lack of response to selection on sperm offence and defence, we quantified sperm precedence across multiple sperm competition bouts using an outbredD. melanogasterpopulation exhibiting continuous genetic variation. Both offensive and defensive sperm competitive abilities were found to be significantly repeatable only across matings involving ejaculates of the same pair of males competing within the same female. These repeatabilities decreased when the rival male stayed the same but the female changed, and they disappeared when both the rival male and the female changed. Our results are discussed with a focus on the complex nature of sperm precedence and the maintenance of genetic variation in ejaculate characteristics.