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In this review, we combine theoretical results, recent genomic data, and documented changes in mating systems among ciliate, fungal, plant, algal, and animal lineages to identify general patterns in the evolution of mating classes and their underlying genetic architectures. We emphasize how interactions among distinct coexisting \u201clayers\u201d of mating classes within a species (e.g., anisogamy plus self-incompatibility systems) have shaped their evolution. 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