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The fungal species  Cryptococcus neoformans  and  Cryptococcus gattii  cause respiratory and neurological disease in animals and humans following inhalation of basidiospores or desiccated yeast cells from the environment. Sexual reproduction in  C. neoformans  and  C. gattii  is controlled by a bipolar system in which a single mating type locus ( MAT ) specifies compatibility. These two species are dimorphic, growing as yeast in the asexual stage, and producing hyphae, basidia, and basidiospores during the sexual stage. In contrast,  Filobasidiella depauperata , one of the closest related species, grows exclusively as hyphae and it is found in association with decaying insects. Examination of two available strains of  F. depauperata  showed that the life cycle of this fungal species shares features associated with the unisexual or same-sex mating cycle in  C. neoformans . Therefore,  F. depauperata  may represent a homothallic and possibly an obligately sexual fungal species. RAPD genotyping of 39 randomly isolated progeny from isolate CBS7855 revealed a new genotype pattern in one of the isolated basidiospores progeny, therefore suggesting that the homothallic cycle in  F. depauperata  could lead to the emergence of new genotypes. Phylogenetic analyses of genes linked to  MAT  in  C. neoformans  indicated that two of these genes in  F. depauperata ,  MYO2  and  STE20 , appear to form a monophyletic clade with the  MAT   a  alleles of  C. neoformans  and  C. gattii , and thus these genes may have been recruited to the  MAT  locus before  F. depauperata  diverged. Furthermore, the ancestral  MAT   a  locus may have undergone accelerated evolution prior to the divergence of the pathogenic  Cryptococcus  species since several of the genes linked to the  MAT   a  locus appear to have a higher number of changes and substitutions than their  MAT α counterparts. Synteny analyses between  C. neoformans  and  F. depauperata  showed that genomic regions on other chromosomes displayed conserved gene order. In contrast, the genes linked to the  MAT  locus of  C. neoformans  showed a higher number of chromosomal translocations in the genome of  F. depauperata . We therefore propose that chromosomal rearrangements appear to be a major force driving speciation and sexual divergence in these closely related pathogenic and saprobic species.

Morphological and Genomic Characterization of Filobasidiella depauperata: A Homothallic Sibling Species of the Pathogenic Cryptococcus Species Complex