“…Mutator alleles can also be favored by strong selection for phenotypic variation, such as that imposed by immunological attack against pathogens, together with stable linkage to beneficial mutations, provided by haploidy in microorganisms. But the special conditions required for these examples highlight two assumptions that have framed discussion of mutation-rate evolution for most of the past century (for example, Bataillon, 2000;Bell, 2005;Cotton and Pomiankowski, 2007). First, although close linkage may allow a mutator to hitchhike on selection for a beneficial allele, recombination, at least in sexually reproducing populations, will eventually separate the two.…”