AR TICLE Phylogenetic-Based Nomenclatural
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IMA FUNGUS · VOLUME 5 · no 1: 121–134 doi:10.5598/imafungus.2014.05.01.12 Phylogenetic-based nomenclatural proposals for Ophiocordycipitaceae ARTICLE (Hypocreales) with new combinations in Tolypocladium C. Alisha Quandt1*, Ryan M. Kepler2, Walter Gams3, João P. M. Araújo4, Sayaka Ban5, Harry C. Evans6, David Hughes4,7, Richard Humber8, Nigel Hywel-Jones9, Zengzhi Li10, J. Jennifer Luangsa-ard11, Stephen A. Rehner2, Tatiana Sanjuan12, Hiroki Sato13, Bhushan Shrestha14, Gi-Ho Sung15, Yi-Jian Yao16, Rasoul Zare17, and Joseph W. Spatafora1 1Department of Botany and Plant Pathology, Oregon State University, Corvallis, OR 97331, USA; corresponding author e-mail: owensbyc@ science.oregonstate.edu 2USDA-ARS, Systematic Mycology and Microbiology Laboratory, Beltsville, MD 20705, USA 3Formerly CBS-KNAW, Fungal Biodiversity Centre, Utrecht, The Netherlands 4Department of Biology, Pennsylvania State University, University Park, State College, PA 16802, USA 5Biological Resource Center, National Institute of Technology and Evaluation 2-5-8 Kazusakamatari, Kisarazu, Chiba 292-0818, Japan 6CAB International, E-UK Centre, Egham, Surrey TW20 9TY, UK 7Department of Entomology, Center for Infectious Disease Dynamics, Pennsylvania State University, University Park, State College, PA 16802, USA 8USDA-ARS Biological Integrated Pest Management Research, Robert W. Holley Center for Agriculture and Health, Ithaca, NY14853, USA 9Milton Biotech Ltd, Krathum Rai, Nong Chok, Bangkok 10530, Thailand 10Department of Forestry, Anhui Agricultural University, Hefei, Anhui 230036, China 11Microbe Interaction Laboratory, National Center for Genetic Engineering and Biotechnology (BIOTEC), 113 Thailand Science Park, Phahonyothin Rd, Khlong Nueng, Khlong Luang, Pathum Thani 12120, Thailand 12Laboratorio de Taxonomía y Ecología de Hongos, Instituto de Biología, Facultad de Ciencias Naturales, Universidad de Antioquia, Medellín, Colombia and Laboratorio de micología y fitopatología, Departamento Ciencias Biológicas, Universidad de losAndes, Bogotá, Colombia 13Department of Forest Entomology, Forestry and Forest Products Research Institute, 1 Matsunosato, Tsukuba, Ibaraki 305, Japan 14Institute of Life Science and Biotechnology, Sungkyunkwan University, Suwon 440-746, Korea 15Mushroom Research Division, National Institute of Horticultural and Herbal Science, Rural Development Administration, Eumseong 369-873, Korea 16State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China 17Department of Botany, Iranian Research Institute of Plant Protection, P.O. Box 1454, Tehran 19395, Iran Abstract: Ophiocordycipitaceae is a diverse family comprising ecologically, economically, medicinally, and Key words: culturally important fungi. The family was recognized due to the polyphyly of the genus Cordyceps and the broad arthropod-pathogens diversity of the mostly arthropod-pathogenic lineages of Hypocreales. The other two cordyceps-like families, Article 59 Cordycipitaceae and Clavicipitaceae, will be revised taxonomically elsewhere. Historically, many species were new combinations placed in Cordyceps, but other genera have been described in this family as well, including several based on nomenclature anamorphic features. Currently there are 24 generic names in use across both asexual and sexual life stages Ophiocordycipitaceae for species of Ophiocordycipitaceae. To reflect changes in Art. 59 in the International Code of Nomenclature Tolypocladium for algae, fungi, and plants (ICN), we propose to protect and to suppress names within Ophiocordycipitaceae, and to present taxonomic revisions in the genus Tolypocladium, based on rigorous and extensively sampled molecular phylogenetic analyses. When approaching this task, we considered the principles of priority, monophyly, minimizing taxonomic revisions, and the practical utility of these fungi within the wider biological research community. Article info: Submitted: 31 March 2014; Accepted: 2 June 2014; Published: 19 June 2014. BACKGROUND (Hibbett & Taylor 2013). Groups have already begun to propose names which should be protected or suppressed The revision of Art. 59 in the International Code of within Hypocreales in accordance with the ‘one fungus Nomenclature for algae, fungi, and plants (ICN; McNeill one name’ policy (Geiser et al. 2013, Rossman et al. 2013, et al. 2012) has created a major task for mycologists, who Leuchtmann et al. 2014, Johnston et al. 2014, Kepler et al. must now reconcile under one name various possible names 2014) and others are in progress. Here, we seek to retain existing for different morphs of the same species of fungus names in Ophiocordycipitaceae with the goal of harmonizing © 2014 International Mycological Association You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. VOLUME 5 · NO. 1 121 Quandt et al. priority, monophyly, simplicity of taxonomic revisions, and and Podonectria, members of the Agaricomycetes and minimization of disruption to the research community. Dothideomycetes, respectively (Rossman 1978, Stalpers et The family Ophiocordycipitaceae was described by Sung al.1991, Hughes et al. 2001, Boonmee et al. 2011). Despite et al. (2007) to accommodate species that were determined the large number of taxa associated with Ophiocordyceps, to be phylogenetically distinct from Cordycipitaceae a lack of support for internal nodes resulting in equivocal ARTICLE and Clavicipitaceae s.str. Asexual morphologies in topologies has limited inferences about relationships within Ophiocordycipitaceae show a tremendous range of variation, the genus in previous studies (Sung et al.2007). some of which are restricted in their phylogenetic distribution The most notable species in the Ophiocordyceps clade while others are often found in disparate lineages. For is O. sinensis, which is nearly double the price of gold by example, Verticillium is a common asexual morph of weight (Stone 2008, Shrestha & Bawa 2013) and the subject many species in several hypocrealean families, including of intense research, especially in China (Shrestha et al. 2010, Ophiocordycipitaceae, Cordycipitaceae and Clavicipitaceae Hu et al. 2013, Ren & Yao 2013, Bushley et al. 2013a, etc.). (see Zare et al. 2000, Sung et al. 2001, 2007, and Gams & Almost exclusively found parasitizing the larvae of ghost Zare 2001). moths (Hepialidae: Thitarodes) in the alpine and sub-alpine Ophiocordyceps is the most speciose genus of the pastures of the Tibetan plateau and the Himalayas, this family, and was described originally by Petch (1931a) species is undergoing heavy, possibly unsustainable, and for species of Cordyceps that have septate ascospores destructive harvesting (Cannon et al. 2009, Shrestha & Bawa that do not disarticulate into part-spores at maturity and 2013). asci with inconspicuous apical caps (Petch 1931a, 1933). The recently described genus Elaphocordyceps is Kobayasi (1941) later used Ophiocordyceps as a subgeneric typified by E. ophioglossoides, one of the first Cordyceps classification of the genus Cordyceps, but Sung et al. (2007) species to be described. Species in Elaphocordyceps restored Ophiocordyceps to the rank of genus to include are mostly parasites of the ectomycorrhizal truffle genus those Cordyceps species within Ophiocordycipitaceae Elaphomyces (Ascomycota, Eurotiales). The majority of forming a sister clade with the genus Elaphocordyceps (see Elaphocordyceps species have no known asexual morph, below). The type of the genus is O. blattae, a rarely collected but where known they produce ones which are verticillium- cockroach pathogen for which no culture or molecular data like or Tolypocladium (Sung et al. 2007). There are a few are available. Elaphocordyceps species known to be entomopathogens, Asexual generic names associated with Ophiocordyceps including three cicada pathogens (E. inegoensis, E. include Sorosporella, the oldest name still in use for species in paradoxa, and E. toriharamontana), and one beetle the clade, Hirsutella, Hymenostilbe, Stilbella, Syngliocladium, pathogen, E. subsessilis (syn.Tolypocladium inflatum) and Paraisaria. Hirsutella species typically produce one to (Hodge et al. 1996, Sung et al. 2007). Tolypocladium inflatum several conidia in a limited mucus droplet borne on basally (a name conserved by the rejection of Pachybasium niveum; subulate phialides that taper into slender necks (Gams & Zare Dreyfuss & Gams 1994), is a medicinally important fungus 2003). Hymenostilbe was proposed by Petch (1931b), and and the subject of much research due to its production there is some evidence to support restricting its use within of the immunosuppressant drug, cyclosporin A (Survase the genus Ophiocordyceps to the ‘O. sphecocephala clade’, et al. 2011, Bushley et al. 2013b). The other species of most species of which sporulate from adult insects (Sung Tolypocladium