Phylogenetic classification of Cordyceps and the clavicipitaceous fungi

By G. Sung, N. Hywel-Jones, J. Sung, Janet Jennifer Luangsa-ard, Bhushan Shrestha, J. Spatafora. Studies in Mycology, 2007

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Cordyceps, comprising over 400 species, was historically classified in the Clavicipitaceae, based on cylindrical asci, thickened ascus apices and filiform ascospores, which often disarticulate into part-spores. Cordyceps was characterized by the production of well-developed often stipitate stromata and an ecology as a pathogen of arthropods and Elaphomyces with infrageneric classifications emphasizing arrangement of perithecia, ascospore morphology and host affiliation. To refine the classification of Cordyceps and the Clavicipitaceae, 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 (nrSSU and nrLSU), the elongation factor 1α (tef1), the largest and the second largest subunits of RNA polymerase II (rpb1 and rpb2), β-tubulin (tub), and mitochondrial ATP6 (atp6). Our results strongly support the existence of three clavicipitaceous clades and reject the monophyly of both Cordyceps and Clavicipitaceae. Most diagnostic characters used in current classifications of Cordyceps (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 Cordyceps and the Clavicipitaceae to be consistent with the multi-gene phylogeny. The family Cordycipitaceae is validated based on the type of Cordyceps, C. militaris, and includes most Cordyceps species that possess brightly coloured, fleshy stromata. The new family Ophiocordycipitaceae is proposed based on Ophiocordyceps 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 Elaphocordyceps is proposed for a subclade of the Ophiocordycipitaceae, which includes all species of Cordyceps that parasitize the fungal genus Elaphomyces and some closely related species that parasitize arthropods. The family Clavicipitaceae s. s. is emended and includes the core clade of grass symbionts (e.g., Balansia, Claviceps, Epichloë, etc.), and the entomopathogenic genus Hypocrella and relatives. In addition, the new genus Metacordyceps is proposed for Cordyceps species that are closely related to the grass symbionts in the Clavicipitaceae s. s. Metacordyceps includes teleomorphs linked to Metarhizium and other closely related anamorphs. Two new species are described, and lists of accepted names for species in Cordyceps, Elaphocordyceps, Metacordyceps and Ophiocordyceps are provided.

Subfields: Ecology, Evolution, Behavior and Systematics, Pharmacology, Plant Science

View Paper: https://www.semanticscholar.org/paper/7f0cf9ad62d8feaa28ed4b58129de8eb3ccd1e92

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Citations in the corpus

Refined families of Sordariomycetes (2020, methodology): “The genus infects insects and produces bright, stipitate, fleshy stromata, superficial to embedded perithecia, ascospores which break or do not break into part spores (Sung et al. 2007).”
Steps towards sustainable harvest of Ophiocordyceps sinensis in Bhutan (2009, background): “Despite claims to the contrary (e.g. Pegler et al. 1994; Li et al. 1999) the ascospores of Ophiocordyceps species are aseptate and do not fragment following dispersal in common with true Cordyceps species (Sung et al. 2007).”
Entomopathogenic fungi as biological controllers: New insights into their virulence and pathogenicity (2012, background): “A critical question is whether species of Metarhizium, Beauveria, Lecanicillium and Isaria (Luangsa-ard et al., 2005; Sung et al., 2007) function in nature as ecologically obligate insect parasites or make use of additional sources of nutrition.”
Yartsa Gunbu (Cordyceps sinensis) and the Fungal Commodification of Tibet’s Rural Economy (2008, background): “A recent molecular study by Sung et al. (2007) concluded that Cordyceps should be divided into three genera, with C. sinensis belonging to the largest of the three, Ophiocordyceps.”
AJOM new records and collections of fungi: 1–100 (2020, methodology): “Known hosts (based on molecular data) – Cocoon of a Limacodidae insect (Meng et al. 2014, Zha et al. 2019); Iragoides fasciata (Lepidoptera) (Spatafora et al. 2007, Sung et al. 2007).”
Fungal diversity notes 603–708: taxonomic and phylogenetic notes on genera and species (2017, methodology): “Ophiocordyceps Petch Quandt Ophiocordyceps was introduced originally by Petch (1931), and revised as the type genus under the family Ophiocordycipitaceae (Sung et al. 2007), with the type”
Challenges in modelling complexity of fungal entomopathogens in semi-natural populations of insects (2009, background): “Phylogenetic analysis suggests that the Cordyceps are not monophyletic but occur in three families: Clavicipitaceae, Cordycipitaceae and Ophiocordycipitaceae (Sung et al. 2007; Blackwell 2009).”
Myconet Volume 14. Part One. Outline of Ascomycota—2009. Part Two. Notes on Ascomycete Systematics. Nos. 4751–5113 (2010, background): “(4842, 5000) In a multi-gene phylogenetic study by Johnson et al. (2009), this genus was shown to be highly polyphyletic, with species clustering in Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae.”
Fungi as Biological Control Agents of Plant-Parasitic Nematodes (2020, background): “The teleomorphs of Pochonia species are located within Metacordyceps [172].”
Insect pathogens as biological control agents: Back to the future. (2015, neutral): “…Ranjard et al., 1190 2001; Sung et al., 2001; Bidochka et al., 2002; Enkerli et al., 2005; Huang et al., 2005; McGuire 1191 et al., 2006; Nielsen et al., 2005; Rehner et al., 2006; Hibbett et al., 2007; Inglis et al., 2007; 1192 Sung et al., 2007; Meyling et al., 2009; Enkerli and Widmer, 2010).”
Genome Sequencing and Comparative Transcriptomics of the Model Entomopathogenic Fungi Metarhizium anisopliae and M. acridum (2011, background): “Cordyceps taii has been described as the sexual type (teleomorph) of Metarhizium taii [7,85] but the sexual stages of M.”
What is the Chinese caterpillar fungus Ophiocordyceps sinensis (Ophiocordycipitaceae)? (2010, background): “Hirustella is one of the common anamorphic genera of Ophiocordyceps (Sung et al. 2007; Kurihara et al. 2009) Chen et al.”
Fungal diversity notes 491–602: taxonomic and phylogenetic contributions to fungal taxa (2017, background): “The families Clavicipitaceae, Cordycipitaceae and Ophiocordycipitaceae were recognized by Sung et al. (2007) based on phylogenetic analyses.”