Phylogenetic relationships of Erysiphales within Ascomycota were inferred from the newly determined sequences of the 18S rDNA and partial sequences of the 28S rDNA including the D1 and D2 regions of 10 Erysiphales taxa. Phylogenetic analyses revealed that the Erysiphales form a distinct clade among ascomycetous fungi, suggesting that the Erysiphales diverged from a single ancestral taxon. The Myxotrichaceae of the Onygenales was distantly related to the other onygenalean families and was the sister group to the Erysiphales clade, with which it combined to form a clade. The Erysiphales/Myxotrichaceae clade was also closely related to some discomycetous fungi (Leotiales, Cyttariales and Thelebolaceae) including taxa that form cleistothecial ascomata. The present molecular analyses as well as previously reported morphological observations suggest the possible exsistence of a novel evolutionary pathway from cleistothecial discomycetous fungi to Erysiphales and Myxotrichaceae.However, since most of these fungi, except for the Erysiphales, are saprophytic on dung and/or plant materials, the questions of how and why an obligate biotroph like the Erysiphales radiated from the saprophytic fungi remain to be addressed. We also estimated the radiation time of the Erysiphales using the 18S rDNA sequences and the two molecular clocks that have been previously reported. The calculation showed that the Erysiphales split from the Nlyxotrichaceae 190-127 myr ago. Since the radiation time of the Erysiphales does not exceed 230 myr ago, even when allowance is made for the uncertainty of the molecular clocks, it is possible to consider that the Erysiphales evolved after the radiation of angiosperms. The results of our calculation also showed that the first radiation within the Erysiphales (138-92 myr ago) coincided with the date of a major diversification of angiosperms (130-90 myr ago). These results may support our early assumption that the radiation of the Erysiphales coincided with the evolution of angiosperm plants.
Key WordsAscomycota; evolution; molecular clock; plant pathogen; powdery mildew.The fungi belonging to the Erysiphales (powdery mildews) are obligate biotrophs that infect a wide range of angiosperm plants and appear as white, powdery material on the leaves, stems, or fruits. Braun (1987) described one family, 18 genera, and 435 species of Erysiphales in his monograph. This fungal group has the following unique characteristics as plant pathogenic fungi. 1 ) Obligate biotrophy: Biotope of Erysiphales is completely restricted to living plants and lacks saprophytic life stages. Several plant pathogenic fungi such as rusts and downy mildews are also known as obligate biotrophs. Although the obligate biotrophic nature is not unique to Erysiphales, it is an important characteristic in considerring the evolutionary history of these fungi.2) Ectotrophy: All but a few genera of Erysiphales are ectotrophic fungi. Their mycelia usually distribute on host surface and put haustoria, a nutrient-absorbing organ, into the epidermal...