Sexual reproduction is a common reproductive system among many groups of organisms (Rice, 2002). A prime advantage of sexual over asexual reproduction is that sexual reproduction introduces genetic variation among individuals (Otto, 2009). Sexual reproduction and recombination integrate beneficial mutations that have independently occurred in different strains (the Hill-Robertson effect: Hill & Robertson, 1966), thereby facilitating adaptation in sexually reproducing organisms (Fisher, 1930;Muller, 1932). Recombination can also give advantages to sexual reproduction in a co-evolutionary arms race with parasites (the