Ultrastructural markers and systematics in smut fungi and allied taxa

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Abstract
To identify monophyletic groups within the smut fungi and related taxa, characteristics of hyphal septation and zones of host–parasite interaction were analyzed by serial-section electron microscopy of 139 species belonging to 50 smut and 10 allied genera. Our results support the hypothesis of the existence of two phylogenetically separate lines of smut fungi. The first line, the Microbotryales, is composed of Aurantiosporium, Fulvisporium, Liroa, Microbotryum, Sphacelotheca, and Ustilentyloma, which are distributed into the Ustilentylomataceae and Microbotryaceae. The second monophyletic line, the Ustilaginomycetes, is formed by the remaining smut fungi studied here together with the Exobasidiales, Graphiolales, and Cryptobasidiales (in the traditional sense). The ultrastructural analysis identifies three lineages within the Ustilaginomycetes. The Entorrhizomycetidae are represented by Entorrhiza. The Ustilaginomycetidae consist of the Urocystales and Ustilaginales. The Exobasidiomycetidae are composed of the Doassansiales, Entylomatales, Exobasidiales, Georgefischeriales, Graphiolales, Microstromatales, and Tilletiales. The Entorrhizomycetidae, Exobasidianae, Entorrhizales, Entylomatales, Doassansiales, Georgefischeriales, Microbotryales, Microstromatales, Tilletiales, Urocystales, Entorrhizaceae, Entylomataceae, Georgefischeriaceae, Mycosyringaceae, Rhamphosporaceae, and Ustilentylomataceae are proposed as new taxa. The descriptions of the Exobasidiomycetidae, Ustilaginomycetidae, Exobasidiales, Ustilaginales, Doassansiaceae, Tilletiaceae, and Ustilaginaceae are emended. Some species of Ustilago are transferred to Microbotryum. Key words: basidiomycetes, classification, Exobasidiales, Graphiolales, phylogeny, smut fungi, ultrastructure.