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Sexual reproduction is of such fundamental importance in eukaryotes that, with unique exceptions, it appears essential for the long-term persistence of species (43). The benefits of sex stem from both the DNA maintenance and repair during meiotic divisions and the impact of various forms of mating upon genetic variation. Meiosis is retained by virtually all sex- ual organisms, and aside from the meiotic parthenogens, mat- ing is required between distinct and alternate forms, called sexes or mating types (but interesting cases exist in fungi of same-sex mating and sex without evidence of meiosis (17, 76)). Mating compatibility is determined by a great variety of mech- anisms. In reptiles, diploid males and females are determined by the incubation temperature during egg development (60). Some fish and plants can switch sexes during their lifetime in response to environmental conditions (4, 70). However, the irreversible differentiation into different sexes or mating types is genetically determined in the majority of species. In many animals and plants, sexual development is established by genes found on a dimorphic pair of sex chromosomes, where females and males differ with regard to being homogametic or hetero- gametic (i.e., whether or not a diploid carries two copies of the same sex chromosome or one copy of each alternate sex chro- mosome). The origin of sex chromosomes is believed to have involved suppression of recombination around the essential sex deter- mining genes on an ancestral autosomal pair (86) and progres- sive expansion of this region of suppressed recombination through the recruitment of other sex-related factors (92). Re- combination suppression and the bias in the extent of het- erozygosity between the homogametic or heterogametic sex chromosomes would then drive the accumulation of load loci and the degeneration of sex chromosomes via reduced effective population size and Muller's ratchet (21). The sex chromo- somes indeed not only diverge from each other, but they de- generate relative to autosomes, retaining few coding genes and sometimes drastically reducing in size (21).

Mating System of the Anther Smut Fungus Microbotryum violaceum : Selfing under Heterothallism