“Phylogenetic Classification of Cordyceps and the Clavicipitaceous Fungi”
Open accessAbstract
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. Taxonomic novelties: New family: Ophiocordycipitaceae G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora. New genera: Elaphocordyceps G.H. Sung & Spatafora, Metacordyceps G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora. New species: Metacordyceps yongmunensis G.H. Sung, J.M. Sung & Spatafora; Ophiocordyceps communis Hywel-Jones & Samson. New combinations: Cordyceps confragosa (Mains) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, C. ninchukispora (C.H. Su & H.-H. Wang) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora; Elaphocordyceps capitata (Holmsk.) G.H. Sung, J.M. Sung & Spatafora, E. delicatistipitata (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, E. fracta (Mains) G.H. Sung, J.M. Sung & Spatafora, E. inegoënsis (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, E. intermedia (S. Imai) G.H. Sung, J.M. Sung& Spatafora, E. japonica (Lloyd) G.H. Sung, J.M. Sung& Spatafora, E. jezoënsis (S. Imai) G.H. Sung, J.M. Sung & Spatafora, E. longisegmentis (Ginns) G.H. Sung, J.M. Sung & Spatafora, E. minazukiensis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung & Spatafora, E. miomoteana (Kobayasi & Shimizu) G.H. Sung, J.M. Sung & Spatafora, E. ophioglossoides (Ehrh.) G.H. Sung, J.M. Sung & Spatafora, E. paradoxa (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, E. ramosa (Teng) G.H. Sung, J.M. Sung & Spatafora, E. rouxii (Cand.) G.H. Sung, J.M. Sung & Spatafora, E. subsessilis (Petch) G.H. Sung, J.M. Sung & Spatafora, E. szemaoënsis (M. Zang) G.H. Sung, J.M. Sung & Spatafora, E. tenuispora (Mains) G.H. Sung, J.M. Sung & Spatafora, E. toriharamontana (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, E. valliformis (Mains) G.H. Sung, J.M. Sung & Spatafora, E. valvatistipitata (Kobayasi) G.H. Sung, J.M. Sung & Spatafora, E. virens (Kobayasi) G.H. Sung, J.M. Sung & Spatafora; infraspecific: E. intermedia f. michinokuënsis (Kobayasi & Shimizu) G.H. Sung, J.M. Sung & Spatafora, E. ophioglossoides f.alba (Kobayasi & Shimizu ex Y.J. Yao) G.H. Sung, J.M. Sung& Spatafora, E. ophioglossoides f. cuboides (Kobayasi) G.H. Sung, J.M. Sung & Spatafora; Metacordyceps brittlebankisoides (Z.Y. Liu, Z.Q. Liang, Whalley, Y.J. Yao & A.Y. Liu) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, M. campsosterni (W.M. Zhang & T. H. Li) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, M. chlamydosporia (H.C. Evans) G.H. Sung, J.M. Sung, Hywel-Jones & Spata
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The scientific name of the caterpillar fungus is now often written as Cordyceps sinensis (Berk.) Sacc. or Ophiocordyceps sinensis (Berk.).
“The scientific name of this fungus is now often written as ' Cordyceps sinensis (Berk.) Sacc.' or ' Ophiocordyceps sinensis (Berk.).'”
From Transnational Travels of the Caterpillar Fungus in the Fifteenth through Nineteenth Centuries (2017) · cites this paper as Sung et al. 2007
The Taxonomic Revision, under “Accepted names and new combinations for Ophiocordyceps,” lists Cordyceps sinensis (Berk.) Sacc. as an equivalent name. It gives the new combination as Ophiocordyceps sinensis (Berk.) G.H. Sung, J.M. Sung, Hywel-Jones & Spatafora, so the citing sentence omits the authors of that combination. The passage does not establish how often either name is used. (LLM, conf=0.86)
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Transnational Travels of the Caterpillar Fungus in the Fifteenth through Nineteenth Centuries
Lu, D. (2017) · Asian Medicine · doi:10.1163/15734218-12341387 · Cited by 3 · see citing works
- Authors
- Sung, G.-H., Hywel-Jones, N. L., Sung, J.-M., Luangsa-ard, J. J., Shrestha, B., & Spatafora, J. W.
- Affiliations
- Oregon State University (US)
- Kangwon National University (KR)
- National Center for Genetic Engineering and Biotechnology (TH)
- Corresponding author
- Sung, Gi-Ho
- Venue
- Studies in Mycology
- Published
- 2007
- Volume
- 57
- Pages
- 5-59
- ISSN
- 0166-0616 (ISSN-L)
- PMC
- PMC2104736
- OpenAlex
- W2117540003
- Cited by
- 1,077
- References
- 0
- Citations by year
- 2026: 72 · 2025: 68 · 2024: 81 · 2023: 70 · 2022: 65 · 2021: 79 · 2020: 60 · 2019: 56 · 2018: 55 · 2017: 71 · 2016: 58 · 2015: 63 · 2014: 49 · 2013: 47 · 2012: 48
- FWCI
- 40.55
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Studies in Mycology (publishedVersion, cc-by-nc-nd)
- PubMed Central (publishedVersion, cc-by-nc-nd)
- PubMed (publishedVersion)
- Funders
- National Science Foundation (DEB-0129212 and DEB-0 297 2)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Insect Science › Entomopathogenic Microorganisms in Pest Control
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Plant Pathogens and Fungal Diseases
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Yeasts and Rust Fungi Studies
- Keywords
- Cordyceps, Clavicipitaceae, Cordycipitaceae, Ophiocordycipitaceae, fungal taxonomy, phylogenetic classification, multilocus phylogeny, Elaphomyces, parasitic fungi, monophyly
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