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        Phylogenetic classification of <i>Cordyceps</i> and the clavicipitaceous fungi

        Sung, Gi-Ho,Hywel-Jones, Nigel L.,Sung, Jae-Mo,Luangsa-ard, J. Jennifer,Shrestha, Bhushan,Spatafora, Joseph W. CBS PUBLICATIONS 2007 STUDIES IN MYCOLOGY Vol.- No.57

        <P><I>Cordyceps</I>, comprising over 400 species, was historically classified in the <I>Clavicipitaceae</I>, based on cylindrical asci, thickened ascus apices and filiform ascospores, which often disarticulate into part-spores. <I>Cordyceps</I> was characterized by the production of well-developed often stipitate stromata and an ecology as a pathogen of arthropods and <I>Elaphomyces</I> with infrageneric classifications emphasizing arrangement of perithecia, ascospore morphology and host affiliation. To refine the classification of <I>Cordyceps</I> and the <I>Clavicipitaceae</I>, the phylogenetic relationships of 162 taxa were estimated based on analyses consisting of five to seven loci, including the nuclear ribosomal small and large subunits (<I>nrSSU</I> and <I>nrLSU</I>), the elongation factor 1α (<I>tef1</I>), the largest and the second largest subunits of RNA polymerase II (<I>rpb1</I> and <I>rpb2</I>), β-tubulin (<I>tub</I>), and mitochondrial ATP6 (<I>atp6</I>). Our results strongly support the existence of three clavicipitaceous clades and reject the monophyly of both <I>Cordyceps</I> and <I>Clavicipitaceae</I>. Most diagnostic characters used in current classifications of <I>Cordyceps</I> (e.g., arrangement of perithecia, ascospore fragmentation, etc.) were not supported as being phylogenetically informative; the characters that were most consistent with the phylogeny were texture, pigmentation and morphology of stromata. Therefore, we revise the taxonomy of <I>Cordyceps</I> and the <I>Clavicipitaceae</I> to be consistent with the multi-gene phylogeny. The family <I>Cordycipitaceae</I> is validated based on the type of <I>Cordyceps</I>, <I>C. militaris,</I> and includes most <I>Cordyceps</I> species that possess brightly coloured, fleshy stromata. The new family <I>Ophiocordycipitaceae</I> is proposed based on <I>Ophiocordyceps</I> Petch, which we emend. The majority of species in this family produce darkly pigmented, tough to pliant stromata that often possess aperithecial apices. The new genus <I>Elaphocordyceps</I> is proposed for a subclade of the <I>Ophiocordycipitaceae</I>, which includes all species of <I>Cordyceps</I> that parasitize the fungal genus <I>Elaphomyces</I> and some closely related species that parasitize arthropods. The family <I>Clavicipitaceae</I><I>s. s.</I> is emended and includes the core clade of grass symbionts (e.g., <I>Balansia</I>, <I>Claviceps</I>, <I>Epichloë</I>, etc.), and the entomopathogenic genus <I>Hypocrella</I> and relatives. In addition, the new genus <I>Metacordyceps</I> is proposed for <I>Cordyceps</I> species that are closely related to the grass symbionts in the <I>Clavicipitaceae</I><I>s. s.</I><I>Metacordyceps</I> includes teleomorphs linked to <I>Metarhizium</I> and other closely related anamorphs. Two new species are described, and lists of accepted names for species in <I>Cordyceps</I>, <I>Elaphocordyceps</I>, <I>Metacordyceps</I> and <I>Ophiocordyceps</I> are provided.</P>

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