Extensive trans-specific polymorphism at the mating type locus of the root decay fungus Heterobasidion

Incompatibility systems in which individuals bearing identical alleles reject each other favor the maintenance of a diversity of alleles. Mushroom mating type loci (MAT) encode for dozens or hundreds of incompatibility alleles whose loss from the population is greatly restricted through negative frequency selection, leading to a system of alleles with highly divergent sequences. Here we use DNA sequences of homeodomain (HD) encoding genes the MAT locus of five closely related species of the root rot basidiomycete Heterobasidion annosum sensu lato to show that the extended coalescence time of MAT alleles greatly predates speciation in the group, contrasting loci outside of MAT that show allele divergences largely consistent with the species phylogeny with those of MAT which show rampant trans-species polymorphism. We observe a roughly six-fold greater genealogical depth and polymorphism of MAT compared to non-MAT which argues for the maintenance of balanced polymorphism for a minimum dur...